PSYB51 - Sensation and Perception (5th Edition) PDF

Brief Contents
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CHAPTER 1 Introduction 2
CHAPTER 2 The First Steps in Vision:
From Light to Neural Signals 30
CHAPTER 3 Spatial Vi sion: From Spots to Stripes 52
CHAPTER 4 Perceiving and Recognizing Objects 88
CHAPTER 5 Th e Perception of Color 122
CHAPTER 6 Space Perception and Bi nocular Vision 156
CHAPTER 7 Attention and Scene Perception 200
CHAPTER 8 Visual Motion Perception 236

CHAPTER 9 Hearing: Physiology and Psychoacoustics 260
CHAPTER 10 Hearing in the Environment 290
CHAPTER 11 Music and Speech Perception 320
CHAPTER 12 The Vestibular System and Our Sense

of Equi librium 348
CHAPTER 13 Touch 388
CHAPTER 14 Olfaction 426
CHAPTER 15 Taste 466
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About the Authors
JEREMY M- WOLFE is Professor of Ophthalmology and Radiology at Harvard

Medical School. Dr. \'Volfe was trained as a vision researcher/ experimental psy-
chologist and remains one today. His ea rly work includes papers on binocular
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vision , adaptation, and accomnuxla tion . The bulk of his recent work has dea lt
with visual sea rch and visual attention in the lab and in rea l-world settings sud\
---"- as airport securi ty and cancer screening. He taught Introductory Psychology fo r
over h venty-five years at the tvlass.achu sett.s In stitute of Tedmology, w here he
\>\>' O il the Baker Memorial Prize for und ergraduate teachin g in 1989. He directs the
\ .. Visual Attention Lab a.nd the Center for Ad van ced Med ical fmaging of Brigham
and \Vomen's Hospital.
DENNIS M. LEVI has ta ug ht at the University of California, Berkeley since 2001. He
is Dean/ Professor in the School of Optometry and Professor at the Helen \ Vills
Neuroscience Institute. In the lab, Dr. Levi and colle.Jg ues use p sythophysics,
comp utntional m od eling. and brain imaging (fMRI ) to study the neura l mech a-
nisms of normal p a ttern vision in htmtans, and to leam how they rue degraded
by abno rma l visual exp erience (amblyopia ).
KEITH A . KLUENDER is Department Hea d, Professor of Speech, language, and

Hea ring Sciences, and Professor of Psych ological Sciences at Purdue University.
His research encompnsses: how people h e..i.r complex sounds s ud1 ;;is speech; how
experience shapes the w :.iiy we hear; how w ha t \.Ve hear guides our a ctions and
conuiHmication j dinical problerns of hearing irnpai rmen t or lan g uage delay; and
practical concerns about compute r speed1 recognition a nd hearing aid design..
Dr. Kluend er is deeply conunitted to teaching, a nd h3S taught a wide array of
com ses--phiJ osoph.ic.Jl, psych ological, cmd physiological.
LINDA M. BARTOSHUK is Bus hnell Professor, Department of Food Science and

Huma n Nutrition a t the University of Florida. Her research on taste h as opened
up broo d n tJW aven ues for htr th er establis hing the impact of both genetic
a nd pa thologic.JJ variation in tas te on food preferences, die t, and health. Sh e d is-
covered tha t t..lSte n om1a.lly inhibits other oral sensa tions s uch that da mage to
taste lead s to unexpected consequences like weight gain and intensified oral pain .
Mos t recently, working \.vi th coJleagues in Hor ticulture, he r group fo und tha t a
considerable am0tmt of thesweetr'less in fm.it is actually produced by interactions
between taste and o lfaction in the brain. TIUs may lead to a nevv way to reduce
su gar in foods and beverages.

RACHEL s. HERZ is a n Ad jtmd Assistant Professor in the Depa rtme nt of
Psychiatry' and Human Behavior a t Brmvn Universit)'rs Wa rren Alpert Med ical
Sch ool and Pa rt-ti1ne Faculty in the Psychology Depa rhnent a t Boston College.
Her research focuses on a number of facets of olfactory cogni tion and percep-
tion and on e motion, me n1ory1 and motivated behavior. Using an experimental
approach grmmd ed in evolutionary theory and incorporating bo th cogn:itive-be-
ha vioral and ne uropsyd1ological techniques, Dr. Herz ahns to lmde rs tand hm·\'
b iological mechan isms and cogn itive processes interact to influence percep tion,
cognition , a nd beh avior.
ROBE:RTA L. KlATZKY is the Charl es J. Q ueena n, Jr. Professor of Psychology at

Carnegie Mell on University, w here sh e also holds faculty appoinbnent.s in the
Center for the Ne ural Basis o f Cognition a nd the Htunan-Compute r hl terac tion
Ins titute. Sh e has d one exte ns ive research on haptic and visual object recogn:ition 1
sp ace perception and sp atial thinking, and mo tor p erformance. Her work has ap-
plication to haptk interfaces, nav igation aids for the blind, itnage-guid ed surgery,
teleoperntion, and virtual environments.
SUSAN J . LEDE:RMAN is Pro fessor Emerira of Psychology a t Q ueen 's Uni versity,
w ith cross-ap pointments in the Centre fo r Neu roscience and in the Sch ool of
Computing . He r research in terests span both perception and cognition , wi th
particular em.phases on p syLhophysics, ha.p tic perception and recognition o f
objects a nd their underlying ne ural p rocesses representa tions, mul tisen.sory
pe rception, and sensory-g uid ed motor control.She has applied the results of he r
resea rch to a nunlber of real -world problems, including the design of ha ptic and
multisen.sory inte rfaces for virtual en vironments and
OANIE:L M. is a Professor of O tology and Laryn gology at the Harvard

Med ical Sch ool with appoinhnentsa t the Harvard-NUT Hea lth, Science, and l\ppa
Tech nology p rogram and tl1e Harvard School of Engineering a nd Applied
Scien ce. He is also the Director of the Jenks Vestibular Physiology Laborato ry at
the Eye and Ea r lnfinrui ry. Mudi. of his research career has been
spen t studying how th e brain combines info rma tion from multiple sources, w ith
a speci fic focus on how the brain p rocesses ambigu ous senso1y information from
the vestibular system in the presen ce of noise. Tran slational work includes re-
se..u ch d eveloping new meth ods to help d iagnose p a tients experien cin g vestibu-
lar symptoms a nd research d evelopin g vestibular implants for pa tien ts \vho have
severe p roblem s ,..,;_th their vestibuki.r organs .

Contents
u ...... u..,J6 AmmJ App.l
Introduction 3 Signal Detection Theory 12

Fourier Analysis 15
Welcome to Our World 3
Sensation and Perception 3 Sensory Neuroscience and the Biology
of Perception 1B
Thresholds and the Dawn of Neuronal Connections 21
Psychophysics 5
Neural Firing: The Action Potential 22
Psychophysical Methods 8
Neuroimaging 24
Scaling Methods 9
Summary 29
CHAPTER 2• · • • .• •
The First Steps in Vision: From Convergence and Divergence of Information via
Bipolar Cells 42
Light to Neural Signals 31
Communicabng to the Brain via Ganglion
A Little Light Physics 31 Cells 43
Eyes That Capture Light 32 Dark and Light Adaptation 47
Focusing 1.Jght onto the Retina 33 Pupil Size 48
The Retina 35 Photopigment Regeneration 48
What the Doctor Saw 36 The Duplex Retina 49
Rebnal Geography and Function 38 Neural Circuitry 49
Retinal Information Processing 40 Sensation & Perception in Everyday
Ught Transduction by Rod and Cone Life: When Good Retina Goes Bad 50
Photoreceptors 40 Summary 51
Lateral Inhibition through Horizontal and
Amacrine Cells 42

CONTENTS ix
Spatial Vision: Simple and Complex CeDs 72

From Spots to Stripes 53 R.Jrther Complications 73
Visual Acuity: Oh Say, Can You See? 53 Columns and Hypercolumns 74

A Visit to the Eye Doctor 57 Selective Adaptation :
Types of Visual Acuity 57 The Psychologist's Electrode 77
Acuity for Law-Contrast Stnpes 59 The Site of Selective Adaptation Effects 80
\.Vhy Sine Wave Gratings? 61 SpatiaJ Frequency-Tuned Pattern Analyzers in
Retinal Ganglion Cells and Stripes 62 Human Vision 81
The Development of Spatial Vision 83
The Lateral Geniculate Nucleus 64
Development of the Contrast Sensitl\/ity
The Striate Cortex 65 Function 84
The Topography of the Human Cortex 67 Sensation & Perception in Everyday
Some Perceptual Consequences of Cortical Life: The Girl Who Almost Couldn't See
Magnifleation 68 Stripes 65
Receptive Fields in Striate Cortex 70 Summary 87
Crientation Selectivity 71
Other Receptive-Field Properties 72
CHAPTER 4
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/ -' ·--.' 1
I
1\
Perceiving and Sensation & Perception in Everyday
Recognizing Objects 89 Life: Material Perception: The Everyday
Problem of Knowing What It Is Made
What and Where Pathways 89 Of 111
The Problems of Perceiving and Object Recognition 112
Recognizing Objects 95 Templates versus Structural Descnptions 114
Middle Vision 97 Problems with Structural-Description
Finding Edges 97 Theories 115
Texture Segmentation and Grouping 101 Multiple Recognition Committees? 116
Perceptual Committees Revisited 104 Faces: An Illustrative Special Case 117
Agure and Ground 106 The Pathway Runs in Both D1iections:
Dealing with Occlusion 108 Feedback and Reentrant Processing 118
Parts and \.Vho/es 109 Summary 119
Summarizing Middle Vision 109
From Metaphor to Formal Model 110

X CONTENTS
The Perception of Color 123 Sensation & Perception in Everyd ay

Life: Picking Colors 133
Basic Principles of Color
Perception 1 23 The Umits of the Rainba;v 134
Three Steps to Color Perception 123 Opponent Colors 135
Color in the Visual Cortex 138
Step 1: Color Detection 124
Individual Differences in Color
Step 2: Color Discrimination 124 Perception 140
The Principle of Univariance 124 Philosophical Problem of "Inverted Qualia" 140
The Trichromatic Solution 126 Language and Color 141
Metamers 127 Genetic Differences in Coior Vision 143
n<e History of Trichromatic Theory 128
From the Color of Lights to a World of
A Brief Digression into Ughts, Rlters, and Rnger
Paints 129
Color 145
From Retina to Brain: Repackaging the Adaptation and Afterimages 146
Information 130 ColorConstancy 147
Cone-Opponent Cells in the Retina and The Problem with the Illuminant 149
LGN 131 Physical Constraints Make Constancy
A Different Ganglion Cell Helps to Keep Track of Fbssible 149
DayandNight 131 What Is Color Vision Good For? 151
Step 3: Color Appearance 132 Summary 155
Three Numbers, Many Colors 132
Space Perception and Binocular Random Dot Stereograms 179

Vision 157 Stereo Movies, TY, and Video Games 180
Using Binocular Stereopsis 181
Monocular Cues to Three-Dimensional Stereoscopic Correspondence 182
Space 160
The Physiological Basis of Stereopsis and Depth
Occlusion 161 Perception 184
Size and Position Cues 161
Aerial Perspective 165
Combining Depth Cues 186
Unear Perspective 166 The Bayesian Approach Revisited 186
Pictorial Depth Cues and Pie tures 16 7 Illusions and the Construction of Space 188
Motion Cues 169 Binocular Rivalry and Suppression 190
Accommodation and Convergence 171 Development of Binocular Vision and
Stereopsis 192
Binocular Vision and Stereopsis 172
Abnormal Visual Experience Can Disrupt
Stereoscopes and Stereograms 175
Binocular Vision 195
Sensation & Perce ption in Everyday
Life: Recovering Stereo Vision 1 78 Summary 197

CONTENTS Xi
Attention and Scene Attention and Single Cells 217

Perception 201 Disorders of Visual Attention 219
Selection in Space 203 Neglect 219
The "Spotlight" of Attention 205 Extinction 220
Sensation & Perce ption in Everyday
Visual Search 205
Life: Selective Attention and Attention
Feature Searches Are Efficient 207 Deficit Hyperactivity Disorder (ADHD) 221
Many Searches Ate Inefficient 208
In Real-World Searches, Basic Features Guide Perceiving and Understanding
Visual Search 209 Scenes 222
In Real-World Searches, Properties of Scenes Two Pathways to Scene Perception 222
Guide Visual Search 210 The Nonselective Pathway Computes Ensemble
The B1i1ding Problem in Visual Search 211 Statistics 222
Attending in Time: RSVP and the The Nonselective Pathway Computes Scene
Gist and Layout- Vety Quickly 223
Attentional Blink 212
Mem0ty for Objects and Scenes Is Amazingly
The Physiological Basis of Goad 226
Attention 215 But, Memory for Objects and Scenes Can Be
Attention Could Enhance Neural Activity 215 Amazingly Bad: Change Blindness 227
Attention Could Enhance the Processing of a VVhat Do We Actually See? 229
Specific Type of Stimulus 215 Summary 233
Attention Could Coordinate the Activity of
Different Brain Areas 217
Visual Motion Perception 237 Avoiding Imminent Collision: The Tao of

Tau 250
Computation of Visual Motion 238
Apparent Motion 240
Eye Movements 250
The Correspondence Problem 241 Physiology and Types of Eye Movements 252
Eye Movements and Reading 254
The Aperture Problem 242
Detection of Global Motion in Area MT 243 Saccadic Suppression and the
Comparator 254
Motion Aftereffects Revisited 245
Updating the Neural Mechanisms for Eye
Second-Order Motion 246 Movement Compensation 256
Using Motion Information 247 Development of Motion Perception 257
Going with the Row: Using Motion lnfotmation
Sensati o n & Perceptio n in Eve ry-
to Navigate 247
Something in the Way You Move: Using Motion
day Life: The Man Who Couldn't See
Information to Identify Objects 248
Motion 257
Summary 258

Xii CONTENTS
CHAPTER 9 __ '· Ar;/

Hearing: Physiology and TheAuditoryNerve 274
Psychoac oustics 261 Auditory Brain Structures 279
The Function of Hearing 261 Basic Operating Characteristics of the

Auditory System 281
What Is Sound? 262 Intensity and Loudness 282
Basic Qualities of Sound Waves: Frequency Frequency and Pitch 283
and Amplitude 262
Sine Waves and Complex Sounds 264 Hearing Loss 285
Treating Hearing Loss 28 7
Basic Structure of the Mammalian
Auditory System 265 Sensation & Perception in Everyd ay
Life: Electronic Ears 288
Outer Ear 266
Middle Ear 266
8230331.1 A Su{l)mary 289
Inner Ear 268
Hearing in the Environment 291 Auditory Scene Analysis 308

Spatial, Spectral, and Temporal
Sound Localization 291
Segregation 309
lnteraura/ Time Difference 292 Grouping by Timbre 311
lnteraura/ Level Difference 295
Grouping by Onset 312
Cones of Confusion 296
When Sounds Become Familiar 313
Pinna and Head Cues 297
Auditory Distance Perception 301 Continuity and Restoration Effects 314
Restoration of Complex Sounds 315
Complex Sounds 303
Harmonics 303 Auditory Attention 317
Timbre 304 Summary 318
Sensat ion & Perceptio n in Everyday
Life: Auditory "Color" Constancy 306
Attack and Decay 307

CONTENTS Xiii
CHAPl'ER 1 1''11 • ; , -y-· r ...

Music and Speech Speech 328
Perception 321 Speech Production 328
Speech Perception 333
Music 321
Leaming to Usten 339
Musical Notes 321
Speech in the Brain 342
Sensation & Perception in Everyday
Life: Music and Emotion 325 Summary 346
Making Music 326
CHAPTER 12
The Vestibular System and Our Sensory Integration 370
Sense of Equilibrium 349 Visual-Vestibular Integration 3 70
Vestibular Contributions to Active Sensing 371
Equilibrium 351 Reflexive Vestibular Responses 373
Sensation & Perception in Everyday Vestibulo-Ocular Responses 373
Life: Evolution and Equilibrium 351 Vestibulo-Autonomic Responses 377
Modalities and Qualities of Spatial Vestibulo-Spinal Responses 378
Orientation 352 Spatial Orientation Cortex 381
Sensing .Angular Motion, Linear Motion, Vestibular Thalamocortical Pathways 381
and Tilt 352 Corocal Influences 382
Basic Q.ialities of SpatJaJ 0-ientation:
Amplitude and Direction 353 When the Vestibular System Goes
Bad 383
The Mammalian Vestibular System 356 Mal de Oebarquement Syndrome 383
Hair Cells: Mechanical Transducers 356 Meniere 's Syndrome 384
Semicircular Canals 358
Sensation & Perception in Everyday
Otolith Organs 363
Life: Amusement Park Rides -Vestibular
Spatial Orientation Perception 366 Physics Is Fun 384
Rotation Perception 367 Summary 386
ifanslation Perception 368
Tilt Perception 369

Xiv CONTENTS
Touch 389 Haptic Perception 410

Perception for ActJon 410
Touch Physiology 390
Action for Perception 410
Touch Receptors in the Skin 390
Role of Rngerprints in Perception and
Kinesthetic Receptors 396
Action 412
From Skin to Brain 396 The What System of Touch: Perceiving Objects
Pain 402 and Their Properties 413
Tactile Sensitivity and Acuity 405 The Where System of Touch: Locating
How Sensitive Are We to Mechanical Objects 417
Pressure? 405 TactJJe SpatJal Attention 418
How Rnely Can We Resolve Spatial Social Touch 420
Deta17s? 407 Interactions between Touch and Other
How Rne/y Can We Resolve Temporal Modalities 420
DetaUs ? 409 Sensation & Perception in Everyday
Do People Differ in Tactile Sensitivity? 409 Life: Haptic Virtual Environments 423
Summary 424
Psychophysical Methods for DetectJ'on and

Discrimination 447
Olfactory Physiology 427 Identification 448
Odors and Odorants 427 Individual Differences 449
428
The Human Olfactory Apparatus Adaptation 450
Sensation & Perception in Everyday Cognitive Habituation 453
Life: Anosmia 432
Olfactory Hedonics 454
Neurophysiology of Olfaction 433 Familiarity and Intensity 454
The Genetic Basis of Olfactory Receptors 436 Nature or Nwture? 455
The Feel of Scent 438 An Evolutionary Argument 456
From Chemicals to Smells 439 Caveats 457
Theories of Olfactory Perception 439 Associative Learning and Emotion:
The Importance of Patterns 441 Neuroanatomical and Evolutionary
Is Odor Perception Synthetic or Considerations 458
Analytical? 442 The Vomeronasal Organ, Human Pheromones,
The Power of Sniffing 445 and Chemosignals 459
Odor Imagery 445 Sensation & Perception in Everyday
Olfactory Psychophysics, Identification, Life: Odor-Evoked Memory and the Truth
and Adaptation 446 behind Aromath erapy 462
Detection, Discrimination, and Recognition 446 Summary 464

CONTENTS XV
CHAPTER 15 • .- ,. .
Taste 467 Genetic Variation in Bitter 480
Supertasters 481
Taste versus Flavor 467
Health Consequences of Taste Sensation 482
Localizing Flavor Sensations 468
Sensation & Perception in Everyday Wisdom of the Body: How Do We Solve
Life: Volat ile-E nhanced Taste: A New Way the "Omnivore's Dilemma"? 483
t o Safely Alter Flavors 469 How Do We Regulate Nutrients? Early Belief
in "Specific Hungers" Gave Way to /den-
Anatomy and Physiology of the Gustatory tif1Cation of Conditioned Preferences and
System 470 Aversions 485
Papillae 471 The Special Case of Umami 487
Taste Buds and Taste Receptor Cells 4 72 Tile Special Case of Fat 488
Taste Processing in the Central Nervous is All Olfoctory Affect Learned? 488
System 474
The Nature of Taste Qualities 489
The Four Basic Tastes 475 Taste Adaptation and Cross -Adaptation 490
Salty 476 Pleasure and Retronasal versus Orthonasaf
Sour 477 Offaction 490
Bitter 477 O-u7i Peppers 490
Sweet 478 Summary 492
Glossary 495
References 513
Photo Credits 537
Index 539
8230336 Am111a Appa

CHAPTER 1

Introduction
Welcome to Our World

YOU'VE TAKEN THE PLUNGE TO READ AT LEAST PART OF A TEXTBOOK on
"sensa tion and Yo u maX' be majoring in psyd1ology or s tudying
an allied field, silch as'nei.rro ice or1' o logy, or you may be simply curi ous.
N o matter w ha t interests you most, yotu un derstanding w ill be info rme d by
sensation an.cl pen:eption.
11'tis first chapte r provides an introd ucti on to the sorts of questions tha t
captivate the a uth ors of this book and the sorts of me thods that resea rch ers
have d eveloped to answer those questions. These ar e only examples and we
could h ave picked man y o thers. The res t of the book \¥ill introd uce- you to a
pan ora ma of questions that have occupi ed a nd continue to occupy the a tten-
tion of thousa nds of scientists.
Sensation and Perception

Wha t does your smartphon e feel as you run your fin ger d ovvn its touch screen
(Figure 1 .1)? Wha t does it hear as you w h isper into its receiver? \Ve assume
that these are s illy thoug h it would n o t be s illy to ask about the
lightest touch that the screen could sense or the faintest vibration in the a ir that
the microphone could sense. What does your ca t feel as you run your finger
d o\'\-1l its back? Tha t seems a more reasona ble questi on, though you have no
access to the priva te experie nce of the cat. You d on' t even h ave access to the
private sensations of a person whose back you mig ht stroke. Your own sen sory
experience is directly accessible only to you .
This book is title d Se11satio11 & Pi,rcep l io11. The ability to d etect the press ure
of a finger and, perhaps, to turn that detection into a priv.:J. te experi ence is an
example of sensation. Perception can be thought of as the act o f giving m ean -
ing and/ or purp ose to those d etected sensa tions. How do you unders tand the
fin ge r that mns d own ym.1 r back? ls it a gesture of affection ? 1s it a n offi cer a t
an airpo rt security checkp oint looking for weapons? TI,is book "'";11 trace the
path from stimuli in the world, through you r sense organ s, to the understand-
ing of th e world that you perceive.
Every thin g we feel, th.ink, and d o d epends on sensa tions and perceptions.
For this reason, philosophe rs have thought, talked , and w ritten about the topic
in p rofound. and syste rna tic ways for over h<1i·o m.illemlla. (See Web Essay 1.1:
Senses of Reality through the Ages.) Tile idea that sensation a nd percep-
tion a re central to mental life has deep roots. 1lle eighteenth-century Fren ch sensation The ablllty to detoct a
philosopher Etienne Bo1mot d eCondillac (1715-1780) (Figure 1.2) fa mo us ly stimulus anJ , perhaps, to turn that
asked his readers to imag ine th e m ental life of :a statue w ith no and he detection Into a pr1vate experience.
con cl uded that the s tatue would have no men tal life. TI1en Condillac imagined perception The act of giving mean -
opening the s ta tue's n ose and giv ing it :a whiff o f the scent of a rose. Then , he ing to a detected sensation.

4 CHAPTER 1
thought, the entire rnental life of the statue would consist o f tha t srnell. Wi th
more senses and more experience, Condillac imagined, a real mental life would
d evel op. If our m ent,,l life relies on information from our senses, then it fol-
lows tha t the p lace for the study of the senses is lvi thin the science of human
be havior and human m ental life-tha t is, within psychology. Of course, this
placement is n ot a bsolute. Researchers studying topics in sens..,tion and percep-
tio n can be fo und in biologyi computer science, m edicine, netuosc:ien ce, an d
many othe r fi elds. Cri tically, however, we approach the s tudy of sensation and
perception as a scientific p urs uit As sud 11 it needs scientific method s. TI1a t's
why the next sections of this chGpter are d evoted to the variety of methods
used in thesh..1dy of the senses.
METHOD 1 : THRESHOLDS What is tl'e faintest sound you can hear? H ow

wou ld you know?\ Vh at is the loudest sound you can hear? This last question
is not as stupid as it may so und, th ough it could be rep hrased like this: What
is the loudest. sound you can hea r safely? If you listened to sounds above that
limit, pe.rhcips by b!Gsting your music too enthu siastically, you wo uld change
FIGURE 1.1 Wou ld It make sensi::! to th e t.lnswer to the first q uestion. You would damage your a uditory sys tem.
ask what a c011 phone feels vvhen you Then you would be un.:,bl e to hear the fain tes t smUld tha t you used to be able
stroke its screen? to hear. You r thresh old would have changed (for the worse) How would you
measure that? As we1.l leam in this cha pter, a variety of methods are avail able
for measurin g sensory thres hold s of this sort.
METHOD 2: SCALING-MEASURING PRIVATE EXPERIENCE W hen you

q.ialla (sing. qua/e) In philosophy, say tha t you uhear" or "taste" someth in g, are those experiences--what the
prtvate conscious experlerces of sen- philosophers at ll qualia (singular qr1ale)-th e same as the experien ces of th e
satlm or perceptlcn person you're tal king to? We can' t really ans\·ver the question of whether your
qualitative experience o f " red" is like my qualitative experience of "green" or,
for tha t matter, "middle C." \ Ve stilJ have no direct way to experience someone
else' s experiences. H owever, we can demonstrate tha t different people d o, in
some cases, inhabit different sensory worlds. Our d iscus.sion in this chap ter
wi ll sh ow h ow scaling me thods can be used to perfonn th.is act of mind re<'ld-
ing. F'urther discussion of qu alia can be fo und in Chapter 5_
METHOD 3: SIGNAL DETECTION THEORY-MEASURING DIFFICULT DECI-

SIONS A radiologist looks a t a ma mmogra m, the X-ray test u sed to screen
for breast cancer. There's something on the X-ray tha t might be a s ign of ca n-
cer, but it is n ot perfectly d ear. \ Vhat s h ould the radiologist d o? Sup pose sh e
d ecid es to call it ben ign, not cancerous, and suppose she is wrong. Her patient
might die. Suppose she d ecid es to treat it asa s ign of malignancy. H er patient
will need m ore tests, perhaps involving surgery. 1l1e patient and her family
will be terribly worried. If the radiologist is wrong and the spot on the mam-
rnogram is, in fact, benign, the consequen ces m ay be less dire than those of
missing a can cer, but there \vill be consequences. This is a perceptual decision,
rnade by an expert, that has real consequ en ces. Our d iscussion of sign...11d etei.."-
tion theory w ill sh ow h ow d ecisions of this sort can be studied scientifical ly,
METHOD 4: SENSORY NEUROSCIENCE Grilled peppers appear on your

They have an appea ling. s moky s mell. When you bite
into one,, tt has a complex that includes some of that sm okiness. Fai rly
quickly you also experience a burning sensation. There is n o actual change in
FIGURE 1.2 !:tienne Bonnot de the temperature in your mouth, and your tong ue is no w arm er th._,_n it was, but
Condil lac imagined how a s tatue oouk:i the "bum" is unmistakable. How does the pepper fool your nervo us system
OOvftlop a tn€.ntal life. into thinking that your tongue is on fire? This chap ter's of sensory

INTROOUCTION 5
neuroscience vvill introduce the ways in which sensory receptors and nen·es
unde rgird your pe rcep tual experien ce.
METHOD 5: NEUROIMAGING-AN IMAGE OF THE MIND Suppose you ar-

range to view cornpletely different images with each of your two eyes. We
ntight p resent a picture of a h ouse to one eye and a face to the other (Tong
et al., 1998). The result would be a n interesting effect known as " binocular
rivalry u (see Chapter 6). 11"'1e h vo images would cornpete to do minate your
pel'C'eption: some times you would see a house, and sornetimes you wou ld
see a face. You would not see the h vo together. One reason binocular rivalry
is interesting is that it represents a d issocia tion o f the stimuli, presented to
the eyes, and your private perceptual experien ce. Even if we cannot share the
experience, modem brain-imaging techniques enable us to see traces of tha t
experien ce as it takes place in the brain . Metho ds of neuroimaging ""'ill be o ur
final top ic in th is chapter.
Thresholds and the Dawn of Psychophysics

FIGURE 1.3 Gustav Fechner
The stud y of the sens es was always a mix of experimental science and phi- lnv9nted p sychophysics and is thought
losophy. It is very interesting to look for p recursors of the modem , scientific by som9 to b9 the truQ founder o f
study of sensa tion and perception. We wi ll s tart wi th the very inte resting and experirr..:intal psychology. Fechner Is
versatile nineteenth -<'E'n tl1ry German scientist-philosop her Gustav Fechner best krmo11n for his plon99flng work
relating changes In the physical world
(1801-1 887) (Fig ure 1 .3). Fechner is s om e times consid ered to be the true to changes in our psycho loglcal
fotmder of experimental psychology (Boringr 1950),e\'en if that title is USL1<.illy 8Xperi91108S-
given to Wilhelm Wundt (1832-1920), w ho began h is work sometime later.
Before making his first contributions to psychology, Fechner had an eventti.tl
personal hi story. Young Fedm erean"'!ed his degree in mediciner but his interests
turned from biological scien ce to physics and mathematics.. Though this might
seem an unlike ly way to get to psychology, events proved o therwise. Fechner
was a very hardworking yo ung scientis t. He worked himself to exh a us tion_
ln tlddition to being ovel"V\>·orked, he s uffered severe eye damage from gazin g
too mud"'! tlt the s un while performin g vision expe riments (a n ot uncommon
problem for curi ous vision researchers in the da ys before reliable, bright1
artificial Light sources). As a conseq uence, Fechner fell into deep depression. Amma Appil
Not only did he resign from his posi ti on at the he a lso \.vithd rew
from a lmost all his friends and colleagues. For 3 yea.rs he spent almost all of
his time alone with his though ts.
Then Fechner experienced w hat he believed to be a miracle when his vision
began to recover quickly. His spiritual con victions d eepened , and he became
absorbed \-v·i th the rela tionship beh-veen mind and tnatter. This pursuit placed
him in the middle of a classic philosophical debate between adherents of dual·
Ism and materialism. Dualists hold tha t the mind has an existen ce separate
from the material world of the body. Materialists ho ld th at the mind is n ot
separa te. A modem mate rial ist position, probably the majority view in scien- dualism The <lea that the mind has
tific psydwlogy1 is that the mind is wha t the bra in d oes. Fechner p roposed to an existence separate from the mate-
effectively s plit the difference by imagining that the mind, or conscio usness, is n<> wa'ld of the body.
present in al l of nahue. TI1is panpsychism-the idea that the mind exists as a material ism The Idea that the onty
property of a ll matter-extended not only to animals, but to inanim.a te things thing that exists Is matter, and that all
things. Including the mind and con-
as well. Fechne r described his philosophy of panpsydtlsm in a provocative sciousness, a.re the results of Interac-
book entitled Nanna, or Concerning the Mental Life of Plants. This title alone tion between bits of matter.
gives a pretty good idea of what Fechner had in mi nd.
panpsychism The Idea that
Ins pired by what we rnight ron.sid er to unronventional ideas, the mind exists as a property of all
Fechner took on the job of explainin g the relation beh\.·een the spirihm1 and matter-that Is, that all matter has
material wor lds: n-Und and body. Fro m his experience as a physicist, Fed1ner consdcusness.

6 CHAPTER 1
FIG U RE 1.4 Ernst WebM discovg-00 that th9srnallest d9tecta.ble change in a stim-
ulus, such as the weight of an object, is a constant proportion of the stimulus level.
This reilationship later bGcamQ known as "Weber's law."
thought it should be possible to describe the relation beh.veen mind and body
using rnathematics. His goal was to forma lly d escribe the re lationship bet>.veen
sensation (mind ) and the energy (matter) that gave rise to that serisation. He
called both his me thods a nd his theory psychophysics (psycho for mind, and
physics for matter).
In his e ffort, Fechner was inspired by the findin gs o f one of his Leipzig col-
leagues, En1st \"leber (1 79&-1878) (Figure 1.4), an anatomist and physio logist
who was interested in touch. \•Veber tested he of our sense of touch by
using a device much like the compass onenui;ht use w hen learning geometry. He
used this d evice to rne..'lsure the sm.-tllest
. distance betv,reen two poi.n ts that \Vas
required fora person to fee.I toudl on two p oints instea d of one. Later, Fechner
would call the dista nce behveen the points the two-point touch threshold. We
wi ll discuss t\'lo-point touch thresholds, and touch in general, in Chapter 13.
For Fechne r, Weber's most important findings involved judgments of
lifted weights. Weber ,..,.·ou ld ask p eople to lift one s tandard weight (a weight
that stayed the sa me over a series of experimental trial s) and one comparison
weight that differed from the standard. \ Veber increased the compru-ison weight
in increm ental over the series of trials. He found tha t the ability of a
s ubject to detect the differe nce be hveen the standard and comparison weights
depended greatly on the weight of the standard. \\'hen the s ta ndmd was rela-
tively light, people were much better at detecting a s mall difference when they
lifted a weight. \ Vhen the standard was heavier, p eople need ed rt
psychophysics The science of bigger difference before they could de tect a change. He called the difference
defining quantitative relationships
between pfW;lcaJ and i:sl'<hologlcaJ required for deta"ting a change in weight the just noticeable dtfference, or
(subjective) ewnts. JND. Another te rm. for JND, the s ma llest in a sti mulus that can be
two-point touch threshold ll1e d etected, is the difference threshold.
minimum distance at which two stimuli \ Veber noticed that JNDs changed in a systematic \·..-ay. The smalles t chan ge
(e.g., two s lmultanoous touches) are in weight that coukl be d etected was a lways d ose to one-fortie th of the standard
just p€fceptlble as separate. ""'eight. Thus, a 1-gram change could be d etected when the s t(mdard weigh ed
just noticeable difference (JND) or 40 grams, but a 10-gram change was required w hen the s tandard weighed
difference threshold The smallest 400 gram s. Weber went on to test JNDs for a few other kinds of stimuli, sud1
detectable difference between two as tbe lengths of two lines, for which the d etectable change ratio was 1:100.
stimuli. or the minimum dhange In a For virtual ly every measure-whethe r brightness, pitd11 or time-a constant
stlmulus that enables it to be CCfrEcily
judged as different from a reference ratio between the change and what was being d1anged could describe the
stimulus. threshold of detectable change quite well. Th is ratio rule holds true except
when intensities, s ize, and so on are very small o r very large, nearing the
Weber fraction 1he constant of
proportionality In Weber' s law. min.imum and m aximum of our senses. [n recognition of Weber's discovery,
Fechner ca lled these ratios Weber fractions. He also gave Weber's observa-
Weber' s law The prlndpe describ-
ing the raatlmshlp bet>t.oon stimulus tion a formula. Fechner named the general rule--that the size
and resulting sensation that says the of the difference (6..l) is a constant proportion (K) of the level of the
Just noticeable dltterence (JNDI Is a s timulus (D-Weber's law.
constant fractbn of the ocmparlscn In Weber's observations, Fechner found what he was looking a way
stimulus.
to describe the relationship beh"1een mind and matter. Fedmer asslm1ed that
Fechner's law A principle describ- the s mallest de tectable change in a s timulus (6.I) could be con sidered a unit
ing the relatlcnship betvJeen stimulus of the mind becau se this is the smallest bit of change that is perceived. He
and resulting sensation that says the
magnitude of subjective sensation then mathe mati cally ex tend ed Weber's law to create what became knovvn as
Increases proportionally to the loga- Fechner 's law ( Figure 1.5):
nthm o f the stimulus Intensity. klog R

INTROOUCTION 7
FIGURE 1.5 This illustratbn of Fechner' s law shows

that as stimulus intensity grows large r, larger changes are
required for the changes to OOdatected by a percelv«.
w h e re S is the psychological sen s'1 ti on , which is

equal to the logarithm of the physical s timulus leve l
(log R} rnul tiplied by a cons tant, k. This e quation
d escribes the fact that o ur p sych ologica l experi-
ence o f the intensity o f light, sound, s rne ll, taste, or
toud1 increases less quickly than the actual physi-
cal stimulus increases. With th is equation, Fechner
provided a m a them atical expression tha t formally
d em onstrated a re la ti onship between psyche and
physics (psychophysics).
Consider the si milarity between Fedmer 's law
and Albert Einstein's famous equ a tio n:
E
Like mind and body, energy (f)nnd mass(m) had, before Eins tein , been thought absolute tlTeshold The minimum
o f as d istinct things. CT"he le tter c corresp onds to the speed o f light, a lm os t a amount of stlrnulatloo necessary fcr a
billio n feet per second .) Just as Einstein sh o\lved how to equa te energy and persoo to detect a stimulus 50% of the
time.
m ass, Pechner provided us w ith at lea.s t one way to think abo ut mind and
ma tter as equivale nt. As you learn about the senses lvhen reading thi s book,
you w ill find that we typicaUy make a distinction between un.its of physical
entities (light, sound ) a nd measures of people's perception . For exa mple (as
we' ll learn in 01apter 9), we ineasure the physical intensity of a squn"d'(sound a Appa
pressure lev el) in d ecibels, but we refer to o ur sensation as "lo udness."
Feclmer invented new l·v ays to measure what people see, hea_r, and feel.
All of his me thods are s till in u se today. ln explain.1.ng these rne t:hods here, we
will use absolute threshold us an example becau se it is simpler to understand,
but we would use the s..1.rne methods to d etermine difference threshold s s uch
as An absolute threshold is the minimum intensity of a s timulu s tha t
can be detected (Table 1.1 ). This returns us to the question we raised earlie r;
W hat is the faintest sound yo u can hea r? Of cou rse, we could ask the same
ques ti on about the faintest light, the lightes t touch, and so forth . (See Web
Activity 1.1 : Psycho physics.)
TABLE 1.1
Some commonsense absolute thresholds
sense Threshold
Vision Stars at night, or a candle flame 30 miles away on a dark, clear night
Hearing A ticking watch 20 feet away, with no other noises
Ves.tibular A tilt of less than half a minute on a clock face
Taste A teaspoon of sugar in 2 gallons of water
Smell A drop of perfume in three rooms
Touch The wing of a fly falling on your cheek from a of 3 inches
source: From Galanter. 1962.

8 CHAPTER 1 8230336 AmMa Appa
(a) (b)
100 JOO
"I hear it:'

!!. 75
lfil"
50
____________ _
"Cl
.W 25
"I don't hear it:' J

JO 11 12 JO II 12
Stimulus le\'el (a rbitrary units) Stimulus level {aibitr.uy units)
FIGURE 1.6 The mg.thod o f constant stimuli . (fl) We might expect the threshold to
b9 a sharp change in detection from never reported to always r.;:ported , as depicted
hGr9, but this is not so. fP) In rQ(l.lity, QXP"1flments ITl9asuring absolute thrashcid pro-
duce shallower functions rtilatlng stimulus to response. A somewhat arbitrary point on
the rurve. oftm 50% detection , is designated as the threshold.
Psychophysical Methods
H ow can we measure an absolute threshold in a valid a nd reliable milnner?
O ne method is known as the method of constant s·ttmull. This me th od re-
quires creating m a ny stimuli with d ifferent intensities in order to find the tini-
est intensi ty tha t ca n be d e tected (Figure 1.6). lf you 've had a hearing test, you
may recall ha ...ing to report you could a nd could n ot hear a tone that the
m1di ologistpl.ayed to you over headpho nes. In this test, intensities of all of the
tones would be relatively low, n ot too far above or below the intensity where
your th resh old was expected to be. TI1e tones, varying in intensity, '"''ou.ld be
presented randomly, and tones would be presented multiple times at each inten-
sity. The "multiple times" piece is important. Subtle perceptual judgments (e.g.,
thresh o ld judgrnent.s) are variable. TI1e stimulus va ries for physical reasons. Tile
observer varies. Attention waivers and sensory sys tems fluctua te for all sorts
of reasons. As a con seque nce, one measu re is almost never enough . You need
to repeat the measure over and over and the n average responses or othenvise
describe the pattern of re.s Lt! ts. Some experiments require thousands of repetitions
(thous ands of "tr ials'1 to establish a s ufficiently reliable data point. Re turning
to our auditory exarnple, as the listene r you would report w he ther you hemd
a tone or not. You would always report hearin g a tone that was relatively fur
above threshold, and almost never re p o rt hearing a tone that was well below
threshold . In ben.veen, h owever, you would be more likely to hear some tone
method of constalt stimuli A intensi ties than not hear the m, and you \Vould h ear other, lower intensities on
i:Gychopl1yslcal method In which many only a few presentations. In generat the intensity a t w hich a sti mulus would
stimuli , from rareti,i to almost be detected 50% of the time would be chosen as your thresh old.
always perceivable (or rarely to almost That 50% d efi nition of absolute thresho ld is ra ther interesting. Weren' t
always perceivably different from a we looking for a way to measure the \Veakest detectable s timulus? Using the
reference stimulus), are presented one
at a time. Participants respond to each hearing exa mpl e, .shouldn ' t that be a val ue below w hich \.Ve ju st can 't hear
presentation: "yeafno," •same/differ- an ything (.see Figure l.6a)? It turns o ut ti-Mt no .sud1 hard lxnmdary exists .
ent," and so on. Because of variability in the n ervous system, stimuli near thre.shold will be

INTROOUCTION 9
d etected sometimes and missed at other times, As a resu.lt, the re la t- Trial series
ing the probability of d etection ·with the s timulus level MJI be more grad ual
(see Figure l.6b), an d we mus t settle for a somewha t arbitra ry d efinition
11 j2 13 14 Is f6 j7 ta
20 y y
of an absolute tluesh old,. (We wi ll re turn to this issue when we talk abo ut
signa l detedion theory.) 19 y y y y
The rneth od of constant stinnili is simple to use, but it can be inefficient lS y y y y
in an experiment because much of the s ubject's time is spent wi tll stimuli
17 y y y y
that are clear ly well above or below threshold . A som ewhat more efficient
approach is the method of limits (Figure 1.7) . Wi th this method, the exp eri- 16 y y y y y
me nter begins with the same set of s timuli-in this case, tones that vary in y y y y y y y y
15
inte nsity. Instead of random presentations, tones are presented in orde r of
increasing or d ecreasin g intens ity. When tones are p resented in ascending 14 y N y N y N y y
-- --- - -- - - -- - -- -- --- --
order, from faintest to loudest, listeners are asked to repor t w hen they first 13 N N y N y N N y
h ear tile tone. \ Vith descending order, the task is to report w hen the tone is
l2 N N N N N N
n o longer audible. Da.ta from a..n e xperiment s ud1 as this s how tha t there is
some "overshoot" in jud gm e n ts. [t us ual.ly takes more inten sity to re port 11 N N N N
hearing tile tone whe n in tensity is increas ing, a nd it takes more decreases 10 N N N N
in intensity before a listene r reports tlla t the tone cannot be heard_\ Ve take
13.5 14.5 1251 4.5 ! LS 14.5 13.5 12.5
the average of tllese crossover points-\'\-hen lis tene rs shift from reporting
hearing the tone to not hearing the tone, and vice versa-to be the tlue.sho ld . Crossover v allles (average-= 13.5)
The third a nd final of these classic measures of thresh olds is the method
FIGURE 1.7 Th9 method of limits.
of adjustment This me thod is just like the method o f limits, excep t the s ubject Hef9 the subject attends to multiple
is the one who steadily increases or decreases the intensity of the s timulus . The series o f trials. Fo r each s9ries, the
method of adjust men t m.1y be the easiest me thod to understand becau se it is intensity o f the stirnulus Is gradualty
mudl like d ay-to-day activities s uch as adjusting the volume dial on a stereo incroosed or decreased until the sub-
or the dimmer 5"\.'itch for a light. Even tho ugh it's tile easiest to unde rstand, ject d.:atacts M or fails to detect (N),
respectlw ty. s titnulus. For each
the me thod o f adjus tm ent is not us ua lly used to measure thresh old s. 11le series. an esti rnate of the threshold (red
m etl,od wo uld be perfec t if tluesh old data looked Uke tlw.se p lotted in Figure dashed line) Is taksn to be the avqage
L 6a.. Bu t, real data look m ore like Figure l.6b. The same person will adjust a of the stimulus level jus t before and
dial to different p laces on diffe rent trials, an d m easurements ge t even m essier aft9r the dlang9 In peroeption.
w hen we try to combine the da ta from multip le persons .
Scaling Methods
Mov ing beyond absolute thresholds a nd difference thresh olds, s uppose we
'""' anted to kn ow about tile magni tude of your experiences. For examp le, \Ve
might give you a li ght and ask how muc h additional light you would need to
make a nothe r light that looks hvke as bright? Though that might seem like
an odd ques tion , it tunlS out to be a nswerable. We could give you a knob to
adjust so that you could set the second light to appear twice as bright as the
first, and you could do it
We don 't need to give the observers a light to adjust. A s urprising! y s traight-
forward way to ad d ress the question of the s trength or size of a sensa tion is
to simply ask observers to ra te the experience. For exa mple, we could give method of limlts A psychophysical
observers a series of s ugar solutions and as k the m to assign numbers to each method Jn which the particular dllnen-
sample. We would jus t tell o ur observers that S\'/eeter solutions should get big- eJcn of a stimulus . er the dlffererce
ger nun1bers, and if solution Aseen.lS twice as sweet as solution B, the number betvveen two stimuli, Is varloct Incre-
mentally until the participant responds
assigned to A should be twice the number assigned to R This m e thod is called d ifferently.
magnitude estimation. l11is approadl actua lly works well, even w hen observ-
method of adjustment A method
ers are free to choose their ovro r.1nge of numbe rs . More typically, we might of limits In which the subject controls
begin the experiment by presenting one solution at a n inte nnediate level and tre change In tre stimulus.
telling the taste r to label this level as a specific val ue-10, for insta nce. All of
magnitude estimation A psycho-
the responses should then be scaled sensibly above or below this stand a rd of physical method In which the
10. H you d o this for s ugar solutions, you ,viU get d a ta tk1t look like the blue pant assigns values accord ing to pef -
"sweetness" line in Figure 1.8. celved magnitudes of the stlrnull.

10 CHAPTER 1
FIGURE 1.6 Magnitude eistimatio n. The lines on 100

this graph represent data from magnitude estimation
QXpel'"iITIQflts using electric shocks o f diffgrent cutnaots, 90
lines o f diff9t"ent lengths . solutions o f different sweet-
n9Ss leveils. and lights o f diffQ1'"'11nt brightneSS9S. Th9
exponents that describe these lings are 3.5. 1.0. 0.8. 80
and 0.3, respectively. Fer exponents greater than 1,
such as for electric shock, Fechner' s law does not hcid , 70
and Stevens' power law must be used instead.
§"' 60
.s
1l 50
40
30
8230336 Arnrna Ap 20
10
30 40 50 60 70 80 00 100
Stimulus energy
stevens' power law A prln::::lple Harva rd psychologis t S.S. Stevens (1962, 1975) in vented magni tude esti-
describing the relationship between mation . H e, his s htdents, and successo rs. measured functions like the one in
stimulus and resulting sensatrn that Pigure 1.8 for many different sensations. Even tho ugh observers were asked
says the mag nitude of subjectt-le sen-
satlm Is proportional to the stimulus
to assign num bers to priva te experien ce, the resul ts were m derly and lawful.
magnitude raised to an exponent. H m,.,•ever, they were not the sam e for every type of sensation. Thzit rebtionship
betv..·een stimulus intensity and sensation is described by w hat is n ow know n
as Stevens' power law:
w h ich states tha t the sensation (S) is rela ted to thes'timulus intensi ty(!) by an
exponent (b). So 1 for ex.ample, experienced sen sation might rise with intensity
squared (Ix l). That would be an exponent of 2.0. If the exponent is less than
1, this mea ns that the sensat ion grows less rapidly than the s timulus. 11-U.S is
w ha t Fechner's law and Weber's law would predict.
Suppose you have some lit ca ndles and you li ght 10 more. If you started
\.vi th 1 candle, the change from 1 to 11 candles must be quite drainatic. lf we
add JO to 100, the change will be modest. Adding JO to 10,000 won°t even be
notice1ble. In fact, the exponen t for brightness is about0.3. The exponent for
swee tness is about0.8(Bartoshuk, 1979). Properties like length h ave e..xponents
n ear 1, so, reasonably enough.1 a 12-inch-lon g s tick looks twice as lon g as a
6-in ch-long s tick (S.S. Stevens and Galanter, 1957). Note tha t th is relation-
s hip is true over only a m oderate range of sizes. An inch added to the size
of a spider changes your sensory experience mud\ m o re than an ind1 added
to the height of a giraffe. Som e stimuli ha\.-e exponents g reater tha n 1. In the
painful case of e lectric sh ock, the pain grows wi th f3.S (Stevens, Ca rton, and
Shickman, 1958), so a 4-fo.ld increase in the electrical current is experienced
as a 128-fold increase in painJ
At this p oint in ou r discussion of psychophysics, it is worth taking a mo-
ment to compare the three laws that ha\·e been p resented: Weber's, PedUler 's,
and Stevens'.

INTRODUCllON 11
• VVeber's fmt1 involves a d ear objective measllrement. \Ve know how cross-modality matching The
much ,.,,.e va ried the stimulus, and eithe r the observers can te ll that the to match the Intensities of
stim ulus changed or they cannot. sensations that come from different
sensory mod811tles. This ability allows
• Fedmer's fnw begins \v:ith the sam e sort of objecti ve mei.1Sureme nts Insight Into sensay differenoes. For
as 1Neber 's, but the law is actually a calculation based on some example. a listener might adjust the
assumptions :about how sensati on works. Jn particular, Fechner 's bnghtness of a light until It matches the
law assumes tha t all JNDs are perceptually equi valent. ln foct, this loudness of a tone.
a ssumption turns ot.1t to be incorrec t and feads to some places w here supertaster Supertasters are th:::E;e
the "law'' is viola ted, such as in the e lectric shock example just given. lndMduals who experience the most
• Sff?ivms' power law describes rating da ta quite well, but notice that rating Intense taste soosatlons: fcr some
stimuli, they are dramatically mae
data are qualitatively differe nt fro m the data tha t s upported \¥eber 's
intense than fa rnedlurn tasters or
law. \Ve can record the subjects' ratings a nd we can d)eck w he ther nontasters. Supertasters atso tend to
those ratings are reasonable and. consistent, but the re is n o way to know experience more Intense oral burn and
w hether they are objectively right or wrong. cral tc:uci1 sensatlcn s.
A use ful varian t of the scaling method can s h ow us tha t different indi-
vid uals can live in different sensory worlds, even if they are exposed to the
same stimuli. The me thod is cross-modality matching Q. C. Stevens, 1959).
In cross-modality matching, a n observer adjusts a s timulus of one sort to
m atd1 the perceived magnitude of a s timulus o f a completely different sort.
For examp le, we might ask a listener to adjust the brighh1ess of a light until it
matches the loudne:..s of a particular tone. Again, though the task might sound
odd, people c..m do this, a nd for the most part, everyone with "nomKtl" vision
hearing will produce the same pattern of matches of a sound to a light
We still can 't exa nUne someone else's private experience, but a t Matching sensations
least the relationship of visua l experience and auditory experien ce
appears to be si milar across individuals.
Not so w hen it comes to the sense of taste. There is a molecule
cal led p ropylthiouracil (PROP) that some people experience as
l.oudaot"""""
Brightut light
Supa-tasters-
very bitter, '"'hil e others expe rie nce it as almos t tu.steless. S till
oth ers fall in between. This relationship can be examined formally
with cross-modali ty matching (Marks et al., 1988). If observers
Brightness of the sun
ore asked to match the bitterness of PROP to oth er .sensations
comple tely unrelated to taste, we do not find the sort of agreement
Heat of scalding water
that is found w hen observers match sounds and lights (Figure
1.9 ). Some people- we'll call them nontasters-match the taste
Sound of a fire engine
of PROP to very weak sensation s like the solUld of a watch or
a w hisper. A group o f supertasters assert that the bitterness o f
PROP is simila r in intensity to the bri°'tness Qf the. sun o r the
most inten se pain ever experienced. Medium tasters matdl PROP pa
to weaker stimuli, such as the smell of frying bacon o r the pain of
a mild headache (Bartoshuk, Fast, and Snyder, 2005). As \\Te """'ill
see in Chapter 15, there is a genetic basis for this variation, and it
h .:1.S wide implications for our food preferences and, conseciuently,
for health. For the present discussion, this example s hows tfo:tt Medium__.,.
we can use sca ling methods to quantify '""hat appear to be rea l Brightness of low-beam headlights
Smell of bacon frying
d ifferences in individuals' taste experiences. Pain of a mild h eadache
Brightne.ss of the moon/ loudne.ss

of a convers.:1tion
FIG U RE 1.9 Cross- rTOdality matching. Th9 lwels of bitt9rn8SS o f
cortCQntrated PROP perc eived by nontasters, rredium tastgrs, and
supartasters of PRO P are shO\M"'l on th9 l9ft. Th9 perceiv9d intensities
of a varigty o f everyday SQnsatb ns arg shown ori the right. The arraw Nonlast<:'rs --..
from Qach taster typ9 indicates the o f sensation to wl1 ich thosQ
No sensat ion
tasters matc hed th9 tast9 o f PROP. (Data from Fast, 2004.)

12 CHAPTER 1
Signal Detection Theory

rehll'n tq to the fact tha t they are not absolute.
An important way to think abou t this fact and to d eal vvith it experi mentally
is known as signal detection theory (D. M. Green and Swets, 1966). Signa l
detection theory ho lds that the s tirnulLLS you're trying to d etect (the "signalN)
is always being detected in the presence of " noise.'' If you si t in the quietest
p lace you can find and you wear ymu best noise-canceling heLldphones, you
W"tiU find that you can stUl h e.:ir sametlriug. Simila rl}'i if you dose your eyes in a
dark room , you still see som ething-..'\ mottled pattern of gray with occasional
brighter flashes. This is internal n oise, the static in your n ervous system . \Vhen
you're trying to detect a faint sound or flash of light, you mu st be ab le to de-
tect it in the presence of that in ternal n oise. Do\vtl near threshold, it vvill be
hard to tell a real stimulus from a particula rly vigorous surge of internal noise.
11"1ere is external noise too. Consider that radiologist, introd uced earlier,
reading a mammogram looking for signs of breast cancer. In Figure 1.10, it is
the marked fuzzy white region th.a t is the danger sign.. As you can see, however,
the mamm ogram contains lots of other s inlilar regions. We can th.in k of the
ca ncer as the signal. By the time it is presented to the radiologist in an X-ray,
the re is a signal plus n oise. Elsewhere in the image, a nd in other images, a re
stim uli that are just noise. The radiologis t is a visual expert, trained to find
these particular signals, but sometimes the signa.l will be lost in the n oise an d
missed, and sometimes some noise wi ll look enough like ca ncer to generate
FIGURE 1.10 Marnmogrruns, X- rays
a ialse a lam1 (Nodine et al., 2002).
o f the breast. are used to screen
women for breast cancer. Reading O f course, sometimes n either in terna l no r exte rna l noi se is much of a
such imagQS Is a d ifficult psrooptual problem. When you see this dot, • , you are seeing it in the presence of inter-
task, even for a trained radiologist. na l noise, but the magni tude of that noise i.s so much.smaller than the signa l
(Court9Sy of Dr. Robyn BirdW91 1, 1-iar- generated by the dot that it has n o real impact. Similarly; the do t m ay no t be
vard MOO icaJ Sctool.)
exactly the sam e as other dots, but that varia tion, the external noise, is also
too s mall to have an impact. If asked about the presence o f a dot here, • ,and
its absence here, , you will be correct in youl" ans\ver essentia lly every time.
Signal detection theory exists to help us understand w hat's going on when
we make d ecisions tmder conditions of uncertainty.
BecatISe we are not expert m amm ographers, let's introduce a diffe rent
example to ill ustrate the worki n gs of s ignal d etection theo.ry. You 're in th e
sh ower. 11le wate r is m aking a noise that we '"rill imaginatively call "noise."
Sotn etimes the noise sow"!ds louder to you; som etimes it seem.s softer. We could
plot the distribution of your perception of noise as shown in Figure 1.11 a. On
the x-axis, we h ave the magni tude of your sensation from "less" to "more.''
Imagine that we asked you, over and over again , about yo ur sensa tion. Or
imagine we took many repea ted measmes of the response in your n ervous
system to the so und. For som e instan ces, the response woul d be less. For
some, it would be more. On average, it would lie somewhere in behveen. If
we tabu lated a ll of the responses, we would ge t a bell-shaped {or "normal")
dis tribution of answers, with the peak of that dis tribution sho\'Ving the average
answer that you gave.
Now the phone rings. Tha t w ill be our "signnT." Your perceptual task is to
d etect the signal in the presence of the noise. Wha t you hear is a combina tion
of the ring and the sh°'"'·er. Tha t is, the signal is add ed to the noise, so we can
signa detection theory A psy- imagine tha tnow we hm·e two distributions of responses in yow nervotlS system:
chcphyslcal theoty that quantmes the a noise-alone d istributi on and a signal-pl us-noise distribution (Figure 1.11b).
respcnse of an observef to the pre- For the sake of simplkity; let1ssupp ose that"more" response mea ns tha t it
sentation of a signal In the presence
of noise. Measures cl:ltalned frcm a smmds more like the phone is ringing. So now your job is to decide whether
series of presentations are sensitivity it's time to jump ou t of the shm,·erand answer w hat might be the phone. The
(<l'I and criterion of the obsaver. problem is that you hm·e n o way of knowing Many given m omen t whethe r

INTRODUCTION 13
(c)
(• )
li!:
- Sh ow er "'n ojge" alone
- Ring+noise
f Less - - Mo 1'l!
Your perceptk>n Sounds like phone Sounds like ph one
(cf) Co1Yect rejection (e) Hit (fl False almn (g) !\fas
NO Criterion YES NO Ci"it'2rion YES NO Criterion YES NO Criterion YES
el rh I rh I rh I
Less - - More LeM> - - More Less - - More Less - - Mon

SoWld s like phone Sounds like phonll? Sounds like phone Sounds like phone
FIG URE 1. 11 Det9Cting a s timulus using signal d et ection theory (SDl). (a) SDT
assumes that al l pgrceptual decisions must be made against a background of noise
(the red curv e) g.:nerat:ed In the w-orkl Of ln thie nervou s system . tp) Your job is t o
distinguish neivous systgn rQS?Onsss du.e to noise alonQ (r.ed) o r to signal plus nolS9
(b lue). (c) The best you can do is 9StabHsh a criteflcrl (dotted line) and declare that
you detect something if the response is abOV9 that c riti9flon . SDT indud9s four c lass-
es of responses: rejections (you say "no' and there Is, Indeed, no signa O:
(9) hits (you Sf£f and then;1 is a sig nal) ; rn
fal se-alarm i:<rors {you $8-f "yes· to no
signal): and (g) m iss errors (you say ··no• to a r9aJ sign al).
you 're hea.ring noi.se aloneorsigna l plus noise. 11le best you ca n d o is to decide
on a criterion le ve l of response (Figure1 .11 c). lf the resp onse in your nervous
system exceeds tha t c rite rion , you w ill jtm1p o ut of the s hower and nm naked
and drippin g to find the phone . If the level is be low the criteri on, you will
decid e tha t it is not a rin g and s tay in the s hO"\f\.·er. Note that this "decision " is
mad e a utomatically; it's n ot that you s it d o"",' to ma ke a conscious (soggy)
cho ice. Thus a criterion, in signal de tection theory, is a va lue t±ki.t is someh ow
by the observer. A response, ins ide the observer, a bove criterion
w ill be taken as evidence tha t a signal is present. A response below tha t level
w ill be treated as noise. criterion In slgna.I detection theory,
The re a re four p ossible outcomes (Figure 1.11d-g): You rnight sa y 11 non an Internal threshold that Is set by
the observer. If the Internal response
w hen there is no ring; tha t's a co rrect rejection (Fig ure 1.ll d) . You might say Is above crlterlon, the observer gives
''yes" w hen. there isa ring; that's known as a hit (Figure 1.ll e). Then there a re m e response (e.g., "yes, I hear than.
the e rrors. If you jtmlp o ut of the sh mve r w hen there's no ring, that's a false Below criterlon, the observer gtves
ala rm (Figure 1.llj). If you m iss the call, that's a miss (Figure 1.lls). another res1x:mse (e.g., •no, I hear
nothing").
H mv sensitive are you to the ring? ln the graphs of Figure 1.11 , the sensitiv-
ity is shown as the separation between the noise-alone and signal-plus- noise sensi!Mty In signal detootlm
di stributio ns . If the d istributions a re on top of each o ther (Figure 1.12a), you theory. a val ue that defines the ease
with which an obs€f'Ver can tell the
can' t tell n oise a lone fro m signal p lus noise. A fa lse alarm is just as likely as difference betwoon the presence ard
'1 hit. By knowing the relationship of h its to false alm"ms , you can calcul a te absen::::e of a stimulus cr the differerce
a sensitivity measu re kn crwn as d' (d-prime), w hich would be- a bout zero in bet1N00n stimulus 1 and stimulus 2.

14 CHAPTER 1
(a) No sensitivity <P) Mo<:ler,1 cmitivit,y (c) High sensitivity
llAIA lQ
- ShO\ver "noise" ailone
- Ring+noise
Les& . _ - + - :h.1ore Less . - - . More l..ei!>S , . _ _ . More

Sounds like phone Sounds like phone Sounds like phone
FIGURE 1. 12 Ya..1r sensit Mty to a stimulus Is illustrated by th9 S9paration bet'W99n

the dis tributions o f your respcnse to noise alone (red curve) and to signal plus no ise
(blu9). This S0paratbn is captured by the d' (d· (a) If the dis tributions
cornpletet-J overlap , d' = O and you ha\19 n o ability to detect the signal. fp} If d' is inter-
med iate, you have so1Tl9 s ensWv ity but your perfamanc9 w ill be imperfect. (c) If d' Is
big, then distinguishing signal from noise Is easy
receiver operating characteristic Fi gu re 1.1 2n. In Figure 1 .12c we see the case of a large cl' . He.re you could
(ROC) cuive In studies of signal detect essen tia lly all the rings an d never a false a.larm. The si tu a ti on \ve've
detection, the graphloo plot of the hit been discussing is in behveen (Figure 1.12b).
rate as a function of the false-alarm
rate. these are the same. points tan Now suppose you 're wai ting fo r an important call Even though you really
en the diagonal. lndbatlng that the dor{t want to miss the ca IL you can' t nrn.gically ma ke you rself tnore sensitive.
cbserver cannot tell the difference All you can do is move th e cri terion level of response, as shown in Figure1 .13.
between the presen:::e and absence of H you shift yom criterion to the left, you won' t m iss many ca Us, but you \vill
the signal. As the cbserver's sensitiv- lots of fa lse alarm.s (Figure l.13a). 11lat's annoying. You're nmning around
ity increases, the curve bows upward naked, dripping on the fl oor, and traumatizing the cat for no good reason.
toward the upper len corner. That
pdnt represents a perfect abl llty to dis- Ii you s hift you r criterion to the right, you won 't have those an noying false
tinguish signal trom noise (100% hits, alarms, but you will m iss m ost of the ca lls (Figu re 1.13c). For a fixed va lue
0% false alarms). of d', changin g the cri te rion chan ges the hits and false a larms in predictable
ways. lf you plot false ala rms on the x-axis of a grap h against hits on the y-axis
for different criterion values, you get a curve know n as a re ceiver operating
characteristic (ROC) curve (Figure 1.14).
Suppose you were guessing (the Figu re l .12a si h1ation ); then you mi ght
guess "yes" on 40% of the occasions when the phone ran g, but you would
also guess "yes" on 40% of the oo::asions when the phone did n o t ring. If you
moved your cri teri on and g uessed ''yes" o n 80% of phone-present ocrnsion s,
you would also g uess "yes" on 80% of phone -absent occasion s . Your d a ta
(a) "Gott.1getth.1tcaJl1" (b) "ls that the phone?" (c)
l&
- S.lCM"er"noise" alone
- Ring+noise
Less - - More
SoundEi like phone Sounds like phon e Sounds like phone
FIGURE 1.13 f or a fixed d ', a ll ycu can do is c hange the pattem of your errors by
shifting the respons9 c ritEfion. If you don' t w ant to miss any signals. you m ove your
crite rion to the left (a), but then you hav,g, m ore false alarms. If you don't lik.tl f alse
alarms, you move the rooponse c riterion to the tig ht (c), b ut then you make morn
miss erTors. In all thBSecas9s your SQnsitivity, d', rarnains the same.

INTRODUCTION 15
_ Correct rejectionPr (N/n) O FIGURE 1 .14 Th1oretical reciBiver OPQrating c haract9ristic

10
{AOC) curves fa differMtvaJuesof<f. Note that d'= 0 w hen p9r-
1.0 0 formance is at dlancei. \tVhtin d' ircreastis, the probability of hits
and com;iict rejection s inc r98.8Qs, and 1h9 probability o f misses
and faJse alarms decreases. Prf/\l/n) = probability o f response
signal pr9Sent• when n o sgnal is pr9Sen1: (oorrect rejection) :
Pr(Nls) = "n o signal p-eisent• when signal
is present {miss); Pr(S/n) = probability of response pres-
9flt.. when no s gnal is present (falS9 alarm): Pr(S/s) ,.. probabmty
of responS9 presenr w heo slgnru is (hit;\.
,'
oo' - - -- '- - --'-- - --'-- - -1""'_01.o

False a1rrm Pr (S/11) sine wave A slmple, smoothly
cl1anglng oscillation that repeats
across space. Higher frequency sine
wcwes have more oscillations and
would fall on that ''chance performance" diagonal in Figure 1.14. If you were l<l'Mlf trequencloo have fewer oocilla-
perfect (the Figure 1.1 2c situa tio n}, you would have lOOo/u hits and 0% false tions over a given distance. 1. In hear-
alarms and your d ata point would lie a t the upper left corne r in Figure 1.14. ing, a waveform for whbh variation as
a !Unctlm of time Is a sine !Unction.
Situations in behveen (Figure 1.12b) produce curves beh-veen g uessin g and AJso called pure tone. 2. In \lisb1, a
perfection (the green, purple, and blue curves in Figure 1.14). If your data lie pattern for vmlch variation In a property
below the cha n1."'e l.ine, you did the experitne nt wrong! like brQhtness or cdcr as a function of
Let's return to o m radiologist. She has an ROC curve whose closeness to space Is a sine furictlon .
perfection reflects her expe rtise. On th..'lt ROC, her criterion can s lide up and
to the right, in which case s he w W make m ore hits but also more false alarms,
or down and to the le ft, in \vhich case she '1'-ill have fewe r false alarn1s but
m o re misses. \,Vhere s he places her cri te rion (consciously or unconscio us ly)
w ill depend on many factors. Does the patient have fact ·l !l that make her more
or less like ly to have cancer? What is the perceived :;,cost,,., of a mfssed ca.ricer?
What is the perceived cost of a false alarm? You can see that what started out
as a query about the lack of absolute thresholds ca n become, quite literally, a
1natter of life and death.
Signal detection theory can become a rather complicated topic in detni1. To
learn about how to calculated' and about ROC cUNes, you can take advan tage
of ma ny usefu l website.sand several texts (e.g., f\ilaanillan and C reelrnan,
2005;see Burgess, 2010, if you' re interested in the application t o radiology).
Fourier Analysis
While we're on the top ic of signals, there's just on e m ore tool in the research-
er's arsenal that will prove helpful to you as you lea rn about sensation and
perception. French mathematician Joseph Fourier (1768-1830) (Figure 1 .15)
d eveloped an alyses that permit mode m perception scientists to be tter under-
stand how complex sounds s uch as mu.sic and speed11com p lex head motions,
and complex images such as objects and scenes ca n be d ecomposed into a set
of s impler s ignals. To tmderstand Fourier 's analytical tedm.ique, le t's begin FIG U R E 1. 15 Joseph Fourier
shOWQd how veiy complex signals
1..vith sounds, beca use they're relatively easy to describe. could be und9rstood more easily as
One of the si1nplest kind s of sotmds is a sine wave (in hearing,ap1're to11e) . a combination of simple sine wave
The air pressure in a sine \.1;"ave chan ges continuously (sinusoidally) a t one components.

16 CHAPTER 1
(a) FIGURE 1 .16 SinQ wal/9S. (a) A \librating tuning fo rk produces sinusoidal varla-
One wavelength,
tbns in air pressure - variatio ns that are the stimulus for hearing. We can plot those
i;;G.deg.ree cycle, or p eriod
variatio ns as wcrvfJJS. The perbd is th9 time taken for o ne oomplet9 cydQ o r
ph.>" "-....._ f<-------..i the r;.::essage o f one wavelength. The amplitude is the height o f the wave. Th9 p osi-
shift ltil tb n of ths l/l/ave relative to a fixed is Its phase-measured in degrees out
of a total of 360 deo.gregs, like the 360 degrges aro und a circle. The red ond blue
ill!Vlill{w.t011Amplaud• sine waves shown here a.re separated by 90 degrees of phase. rp) This tuning fork
produ:::es a si ne wave that has half the amplitude and half the \Na'/Qlsngth o f the
waws in (a). In hearing, the frequi:incy of a sine wave Is the number o f cycles pgr
second. In visic:n. you might have sinusoidal vaiiation of light ove< spacs: then the
(b) 'freq uency W"Ould be in cycles pg degree o f visual Mgle (siee C ho.pt9t" 3),
il)'i
{\ {\ {\ {\ { CT reduced
v v v v v
Amplitude
by half
H
V•iaveJe:ngth
frequency (Figure 1.16). Th e time taken for one complete cycle of a s ine wave,
o r fora wavelength to pass a point, is the period o f the sine wave . The height
reduced by half,
frequen cy do ubled of the wave is its amplihtde. T11e phase of the wave is its p osition rela ti ve to a
fi xed marker. Phase is meas ured in degrees, with 360 degrees of phase across
one period, like the 360 degrees around a circle, Thus, in Figure 1.1 6 the red
and blue s ine waves differ by QO d egrees in phase.
Sine ,..;aves are no t common, everyday sounds, because few vibrations in
wavelength The distance required the world are.so pure. U you've taken a heming tes t or used hming forks, you
for cne full cycle of osdll3tlon fcr a sire may heard s ine waves. Flutes can produ ce m usical notes that are close
wave.
to pure tones. but o ther musical ins trumen ts. hum.an voices, birds, cars, an d
period Fcr heanng, t11e time required almos t all otha..SOund SOlU'CeS ln thew d produce complex sounds.
for a 1ull wavelength of an acoustlc Sne
wwe to pass by a point In space. lf pure tones are so uncommon, yo u may wonder why we' re bothering
to discuss them. It hU'JlS o ut all sounds, no matter how complex, can be
phase A fractlcn of the cycle of the d escribed combination of sine wa ves (Figure 1.17). Pourier proved that
sine wave described In degrees (0° to
360C? or radians (0.."t to 2.rr:). In hearing, even the cacophony of room full of people talking or the swelling soLmd of
phase can be Uled to describe frac- a full orches tra can be broken down into combinations of sin e waves a t ntany
tions of a period that re8.te to time. different frequenci.es wi th different amplitudes mid phases. Any complex solmd
Frequency= 1 Frequency= 3 Frequency = 5 Prequency = 7 Fn:-quenc:y = 9 And soon .. •

Amplitude = I Amplitude= 113 Amplitude= 'ts Amplitud.:: = 1h Amplitude= 'f9
FIGURE 1.17 Every sound wai1e can be anatyzed as a combination of

sine waJJes, ooc h with its o wn frequi8tlcy, amplitude, and phase. H9re, multiple. spe·
ciflc sine W<Nes (top) ar9 addOO togethQr to form m ore compl9X waV9forms (bottom).
When infinitety rrcrn siM wa\113S with even highQr frequencies are addOO, a square
waw (bottom right) can be constructQd.

INTRODUCTIO N 17
c.a n be broken dmvn into individual sin e wave compone nts through th.is p rocess, Fourier analysts A mathematical
w hich is cal led Fourier analysis. (See Web Activity 1.2.: Fourier Analysis.) procedure by w hich any signal can be
separated Into component sine waves
Fourier analysis is a P°'"'-erhU tool that is used in m any research
al different l"equencloo. Ccmblnlng
fields. As we vvi.l l see in Chapter 12, Fourier ana lysis is used exte nsive ly by these sine wwes will reproduoo the
vestibtUar and spatial orienta ti on researchers. Vision researchers use Fourier alglnal signal.
analysis to explore the v;sual sy.s te m in \vaysquitesimi lar to th ose employed spaUal frequency The number of
by hearin g research e rs. While sounds are described as changes in pressure cyc"'6 of a gratlrg per unit of 'Jlsual
across time, images c.a n be d escribed as cha nges in light a nd dark across space. angle (usually spoolled In cycles per
Images can be broken d own into com ponents that captu re h o'"'' often d1anges degree).
fro m light to dark occur over a p articuJar region in space, called spatlal fre- cycles per degree The number
quencies. Spa tial frequencies are d efined as the nurnber of th ese light/ d ark of pairs of dark and bright bars per
d1angesacross 1 degree of a person's ";sual field . There would be360 degrees deg ree o f 'Jlsual angle.
around the head . On e degree is about the size of a thumbna il a t arrn ' s length .
Thus, in vision the units of spatial frequency are cycles per degree of visual
.m gle (Fig ure 1.18 ). Jus t as a comp lex sound can be broken down into a set of
sine wave pure tones, a visu al s timulus can be broken d own into componen t
spatial frequencies. \Ve will have more to say a bout this in Chapter 3.
(a) High-frequency (b) Low-frequency (c) High-contrast (tf) Low-contrast

square wave .squa.reweve sinusoidal spatial grid sinusoidal sp<1tlal grid
r-...,
I 10 cycles
11111 . S-cycle
·-
(e) Normal (/) High fr equencies filtered out (g) Low frequencie; filtered out
FIGU RE 1. 18 Thg spacing b9tW99n dark and light stripes in {a) has a spatial fre-
qufficy that is t wice spatlaJ frgquency \Nt1en spatial frequency is rep.-eserrt -
ed as sinusoids (c, d), thr;ire are gri98.t6f" amplitude differences for high contrast
(c) than for bw contrast (d), A photcgraph o f pe!lQlJins (e) ill ustratgs how images
appear when high spatial frequto::::les are taken aw8J (filtQ"OO out) (f) ard wl1m low
spatial frequ'1lncio9S 1ilten:1d out (g). (From & 99dloVQ and Watson, 2013.)

18 CHAPTER 1
Fourie r aru:"\lysis is m ore than a ma thematical curiosity. To a firs t approxi-

1:nationr yom a uditory and visua l systems appear to break down real-world
sotmds and i1nages into sine wave components. When we study how individual
n eurons respond to sotmds and irnagesr we find many neurons that have s trong
p referen ces for som e frequency components over o thers, a nd th.is is especially
true for ea rly s tages of auditory and visual processing. Unde rs tanding h ow
simpler sounds and images are e ncode d p rovides essential insights into h ow
we hear and see real e vents and objects in our world.
Sensory Neuroscience and the

Biology of Perception
docbine of speclftc nerve Most s tude nts taking this course w ill h.1 ve had an introduction to neuroscience.
gles A doctrine . formulated by Our discussion of some of the ne uroscien ce that is releva nt to the stud y of
Johannes MC1lle(. stating that the sensation and perception should serve as a reminder of w ha t you have leam ed
nature of a sensatlcn d€f)ends on elsel"'"'here. lf this is your first encounter with neuroscience, you may want to
which sensory fibers are stimulated,
not on how fibers are stimulated. consult a neuroscience text to g ive yourself a more de tail ed background than
we \vii.I provide here.
cranial nerves Twelve pairs of
nerves (one for each side of the txxl\? Dming the nineteenth century, when\ lkber and Fechne r were initiating the
that orl;Jlnate In the brak1 slem and experi me nta l s h1d y of perception, physiologists were hard a t work learning
reach sense organs ard muscles how the senses and the brain operate. Much of this work involv--ed research on
thrrug h openings In the skull. anima ls. It's wor th spending a moment on a key ass umption here: that s tud-
ies o f animal senses tell us something about huma n senses. That may seem
obvious, but the as.surnption requires the belie f tha t there is some continuity
ben"1een the way ani ma ls work a nd the way humans work.
1l-1e 1lloot powerful argument for a continui ty between humans and animals
CCIUle from Darwin's theory of evolution. During the 1800.s, C harles Darwin
(1809-1882) proposed his revoluti onary theory in Tl1e Origi11 of Species (1859).
Although maw Ideas fo u1,d in Q1a t q<lpk had been brewing for some
time, controversy expanded \vith vigo r fo llowing Dan"in's provoca tive sta te-
.. ments in Tire Desceftt of Mnu (1871), where he a rgu ed thnt human s evolved
from apes. If there was continui ty in the structure of the bones, heart, nnd
kidneys of cov..'S, dogs, monkeys, hum.:ms, then why wouldn't there be
,,
continuity in the structure and ftmction of thei r sensory and ne rvous systems?
An inescapable implication of the theory of evolution is that we can learn
nrnch about human sensation and pen:::eption by s tudyin g the s tructure and
functi on of our nonhwnan rela tives.
At the same time that Danvi.n was at work in England, the Germa n
gist Johannes M iiller(l 801-1858) (Figure 1.19) was \>Ti ting h is very influential
Hn11dbook of Physiology during the early 1830s. In this book, in addition to
covering most of w hat was then known about physiology,lvli.ille r formulated
the doctrine of specific nerve energies. The central idea of this doctrine is
that Vv""e cannot be directly aware of the world itself, and we are only 3ware
of the activity in our nerves. Further, wha t is most impor t.ant is which nerves
are s timulated, and no t haw they are stirnulated. For exa mp le, we exp erience
vision because the op tic n erve leading fr om the eye to the brain is s timulated,
but it d oes not ma tter whethe r light, or .something else, s timulates the nerve.
FIGURE 1.19 Johannes MOiier fCf- To prove to yourself that this is true, close yom eyes and press very gently on
rnulated the doctrine of specific nerve the outside com er of one eye through the lid. (This works better in a darkened
energias, w hich says that we [11"9 aware room.) You wi ll see a spot of light toward the insid e of your \·isual field by
only o f the a.cti\otty In o ur nerves . and your nose. Desp ite the lack of s timulation by light, yo ur brain interp rets the
WQ cannot be aware of the 1NOrld itself.
For this reaso n, 'What is most important input from y01rr optic nerve as in for min g you about some thing visual.
is which r;erves a re stimulated, n ot how The cranial nerves leading into and out of the skull illus trate the d octrine
th.gy are stimulated. of specific nerve energies (Figure 1.20). The pair o f optic nerves is one o f 12

INTRODUCTION 19
FIG URE 1.20 Twetv9 pairs of cranial nerv9S pass through small openings in the
bone at the base of the skull, AH of therSe nerves conduct for sensation,
motorbehavicr. Cf b oth. (After Breedlove and Watson. 20 13.)
VIL F.1ci.1 l
Tongue,
soft palate:
pairs of crani..'ll ne rves that pass thro ugh s mall ope nings in the bone atthe base
of the s kull . The cra nial nerves o:u e d edicated m ainly to sensory a.n d m otor
systerns. Crnnial nerves are labeled both by names and by Roman numerals
thilt roughly correspond to the order of their loc.a tions, beginning from the

20 CHAPTER 1
oHactory (I) nerves The first pair of front of the s kull. Th ree of the cranial ne:rves---Olfactory (I), optic (II), and
cranial nerves. The axons of the olfac- vestibulocochlear (Vlll}-are exclusively dedicated to sensory informati on.
tory sensoiy neurcns bundle together The ve.stibulocochlear n erve serves n.-.·o sensory modali ties: the vestibular
after passing throu;ih the crlbrlform
plate to fam the dfactory nerve, sensa tions that support onr sense of equilibrium (see Chapte r l 2) and hearing
which 001ducts impulses fran the (discussed in Chapter9). TIU'ee more cranial nerves-oculomotor (Ill), trochlear
olfactory eplthella In the nose to the (IV), a nd abducens (Vl)- are dedicated to mus cles that move the eyes. The
olfactory bulb. other six cranial nerves either are exclusively motor (spinc1l accessory [XI] and
optic (IQ nerves The ooccnd pair hypoglossal [XII]) or convey both sensory and motor sign als (trigeminal [\'],
of cranial nerves, which arise trom the facial [VII], glossopharyngeal [IXJ, and vagus [X)). Wi th respect to our study
retlra and c arry ;;sual Information to of perception, in later chap te rs we w ill return to the first six cranial nerves
the thalamus and other parts of the
brain.
d escribed here, because each one plays an important role in our ability to use
o ur senses to lean1 about the '"'orld around us.
vestlbulocochlear (VllQ nerves
1l1e doctrine of specific nen..-e energies extend s beyond the cranial nerves,
The p air of cranial nerves,
which ocnnect tlie Inner ear ,.;111 the as illus trated especially ,.1;ell by our senses o f hot and cold on the skin. Two
brain , transmitting Impulses concerned special types of nerve cells a re warmth fibers and cold fibers, which respond
with hearing ard spatial e<lentatlon. to increases and decrenses in te rnperah1 re on the skin. Capsaidn, a chemical
The vestlbulocochlear """"1is ocm- thatnaturtilly occurs in chili pep pers, ca uses warmth fibers to fire, crea ting a
posed of the cochlear nerve branch sense of increasing heat even thou gh the tempera ture has not changed. On the
ard tl1e vestibular nerve branch.
othe r hand , menthol, \vlUch imparts a minty flavor in cough drops, sti mubtes
oculomolor (Ill) nerves The thrd cold fibers (Ba u tistn e t al., 2007), so s kin feels coo ler wi thout gettin g physically
pair of cranial nerves. v.t'jch Inner-
vate all the extrinsic muscles of the colder. In sufficiently high amounts, both capsa id n and menthol stimula te
"fe except the lateral rectus and the pain receptors in the s kin . Paradoxically; th is is w hy ointments often contain
superior oblique muscloo, and which caps.aici n or menthol, \vhere their effect is to mask real physical pain.
Innervate the elevator muscle of the Just as different nerves are dedicated to individual sensory m otor tasks,
upper "lei Id, the clllary muscle, and areas of the brain s tem and ce rebral cortex are si milarly dedimted to particular
the spl1l ncter musde of the pupll.
tasks . Areas of the cortex d ed ica ted to perception actually are m uch larger than
trochlear PVl nerves The frurth the darkened areas in Figure 1.21 . The areas depicted here are primary sens01y
pair of cranial nerves, v4hlch innervate
areas; m ore complex p rocessin g is accomplished across cortical regions tha t
the supene< c:Oli::l ue muscles of the
"f0balls. spread \.'\fell beyond these primary areas. For example, visual perception uses
cortex th at ex tends both an teriorly (forward) into P..""'rietal cortex and venh·ally
abducens (Vl) nerves The sixth
pair of cranial neivee, which Inner-
(lower) into regions of tl1e temporol lobe (see Figures 4.2 and 4.4 in C hapter 4).
vate the lateral rectus muscle of the In addition, processing extends beyond primary areas, cortex often becomes
"fOballs. potysensory, 1neaning th at information from rnore tha n one sense is being
polysensory Referring to blending combined in som e manner. {See Web ActMty 1.3: sensory Areas In the Brain.)
multiple sensory system s. Herma nn Ludwig Ferdinand von Helmholtz (1821-1894) (Figure 1 .22),
on e of the greates t scientists of the nineteenth century, was greatly influenced
by Millier a t the Free University of Berlin. Although initially inspired by Mu l-
ler, He hnholtz truly disliked one of MUller' s beliefs:
Central vitalism, the idea tha t there is a force in life that is
fissure d istinct from physical entities. By contra.st, Helmholtz
thought that all behavior shottld be explained by only
Parietal physical forces. Vitalism do lated the physical law of
lobe
Frontal conservation of energy, a nd Helmh oltz \¥anted the
Job;, bra in and behavior to obey purely physica l la\vs. He
Oo.:ipital chose a very s mart place to begin his attack on vital-
lobe ism. Mi.U !er had ckUmed that the nerve impulse could
never be measured experimenta lly. So, Helmholt z
se t out to sh ow tha t the activity of ne urons obeys
norm al ntles of physics and chemis try by being th e
Visual
COTfel(
FIGURE 1 .21 C:Orto9X o f th9 human brain. The dar\.<Qned

Sylvian Temporal Auditory ar93S show wl1ere Information from fcur of our sensory
fisrurn lob<i? co rtex m odalities fi rst reaches the oort&"X..

INTRODUCTION 21
first to e ffectively measure hm.v fas t ne urons trans mit their s ignals. (See Web
Activity 1.4: Neurons.)
ln an ear ly effort, Helmholtz estirnated that thespeed of signaJ transmission
in the nerves in frog leg.< was about 9o I P"" 5eoond , cqn1fort:1bly within the
range of ordinary physical events . Lute r he concluded that sensory nerves in
people transmitted signals at speeds of between 165and 330 feet per second. In
all cases, this transmiss ion was s lower than mi.'lny people believed at the fune.
Helmholtz emphasized this point w hen he noted tha t a "whale prOOably feels
a wound near its tai l in about one second, a nd req uire5 another second to send
back orders to the rail to defend itself" (Koenigsberger, 1906/1965, p. 72). As you
may already have guessed, not all ne urons are equal w hen it comes to speed;
som e are fas ter than others. It's still inte resting to realize th...'l.t when you stub
your toe, a measmable ammmt of time elapses before you feel the consequences.
Neuronal Connections
During the second h alf of the nine teenth century, \·vhen Hel mholtz was mak-
ing sh.inning d iscoveries co ncerning vision and heari ng, othe r scientists were
lea ming a grea t d eal about how neurons a nd brains work. After nearly dying
of malaria in Cuba, the Spaniard Santiago Ra m6n y Cajal (1852-1934) (Figure
1 23a) re turned to his h ome land to develop some of the most painsta king and FIGURE 1.22 Herrnannvon HQlm-
hottz was onQ of the greatest sdQOtists
b reathtaking insights into the organiza tion of n eu rons in the brain. Spendin g o f all tlmQ. H& made rnail'f impor-
rrumy h ou rs peering in to a microscope, he made s pectacularly d etailed draw- tant disooV9ties in physlciow and
ings of ne urons and their connec tions tha t remain useful 100 years afte r they perception.
were created . F1gure 1.23b s hows an example.
R.am 6n y Cajal's draw ings s uggested that neurons d o not achmlly tou ch one vitallsm The Idea that there Is a
ano the r. Ins tead , he d epicted nemons as separate cells w ith tiny gaps behveen foroe In that Is dlstk1ct !rem ph\1llcal
them. Sir Charles Sherrington (1857-1 952) (Frgu re 1.24) named the tiny gap entitles.
(•) (b)
--'I""<'!-.<!(
Z!Ti"'1'i:li-H"""'""<.L.;t
FIGURE 1.23 (a) Santiago Ca}al {pi9rhaps not on his h appiest day in the
lab). f,b) Rarn6n y Cajal creatOO thes.e dravvlngs of bro.in ne1.1rons whil& pegring into a
microscope fcr many hours . Because of his painstaking care and accurao,i, his &arty
drawings are still c ited tcx:Jay.

22 CHAPTER 1
FIGURE 1.24 Sir O'larloo Scott Sher--

rlngton was an English ngurophyslolo -
gls t 'Mlo earned a Nobel Prize for his
pbneering discoveries concerning neu-
ral activities.
FIGURE 1. 25 A synapse. An axon terminal in the presynaptlc call communicates

w ith a d 9ndrit9 of the p o stsynaptic cell. Neurotransmitter molecules are r919asGd by
synaptic vesicles in the axon and tit Into receptors m the dendrite on the other side
of thQ synapslil , thus communica.tirl9 from the axon of th9 firs t neuron to the d end rite
ot_th<> 5'!COfld n'*on·
synapse The Junction between neu- behveen the axon of on e neuron and the dendrite o f the next a synapse (Ftgure
rons that permits Information transfer. 1.25), from the Greek word mean ing "to fasten together." Sherrington made
neurotransmitter A chemical sub- very careful measurements demonstrating that the speed of neural transmission
stance used In neuronal communlca- decreases at synapses, and this work helped us understand that something
tbn at S)NpOOS. special h ap pens at this junction w here neurons meet to comm unicate.
Initially, people thought that some sort o f electrical wave traveled across
the synapse fro m one neuron to the n ex t. Otto Loewi (1873-1961) (Figure
1.26)1 however, was con vinced. tha t this could not be true. One reason is tha t
.some n e urbns increase the resp onse of the n ext n e uron (they are excitatory) but
o ther n euron s d ecrease the response of the n ext neuron (they a re itihibitory ).
Loewi prop osed tha t some thing d 1e mica l, ins tead o f electrical, might be at
work a t the synapse. This insight launched many studies about molecules tha t
travel from the axon across the synapse to bind to receptor m olecules on the
de ndrite o f the next neuron . The mo lecules, released by the axon, are called
neurotransmitters. There are many different kinds of neurotransmitters in the
brain, and ind hidua l neurons are selective with respect to whid 1 n eurotra ns-
mitters excite them or inhibit them from firing. Drugs that are psychoactive,
s uch as amphetamines, work by increasing or d ecreasing the effectiveness of
different neurotransmitters. Today, scientists use chemica ls that influence the
effec ts of ne urotransmitters in e fforts to understand pathways in the bra in,
includin g those used in perception.
FIGURE 1 .26 Otto Loewi was a Neural Firing: The Action Potential
German pharmacologist vvho received After Loewi' s discovery of neurotrnns mitters, scientists learned w hat it really
a Nob91 Prize for dgmons trating
m eans to have a neu ron "fire." Investigators m ade the greatest early advances
that neurcns communicate with ong
another b y releaslng chemicals called by takin g ad vantage o f the fact that some squids have giant ne urons as thi ck
MUrotransmltt9"S. as 1 millime ter. At the laboratory o f the Ma rine Biological Associa tion in

INTRODUCTION 23
(c) Hodgkin and Huxley's s quid
FIG URE 1.27 Sir Alan Hodgkin ta)

Plymouth, England, Sir Alan Hodgkin (l914--1998)and Sir Andrew Huxley and Sir Andrgw Huxley f/J} were E<lt-
(1917-2012) (Figure 1.27) seized their opportunity when trawlers opera ting ish physiologists who made the first
rec.:crdlngs from inside a neuron. They
beyond Plyinouth Sound brought ca tdies of fresh .sqtt.id to port. Hodgkin and
mowd from c.ambrldge to Plymouth,
Huxley conducted experim ents in w hich they co uld isolate a single ne uron England , to rQCOrd voltages from lnsid.9
from the sq uid and tes t hm..· the ner ve impulse travels along the axon. \Vith the giant axons of fr€1Sh Atlantic squid
su ch large axons, they could pierce the axon with an electrode to meas m e volt- (c) . They eam8d a NobQI Prlz8 for
'1ge, and they could even inject different chemicals in.side. They learned that d lscowring h ow the action ix>tentiaJ
n eural firing is actually e lectroch e1nical (Agure 1 .28). Voltage increases along 1A10rks to conduct signals along the
axons of neurons.
the axon are caused by changes in the membrane of the neu ron tha t permit
positively charged soditun ions (Na•) to rush very quickly into the axon froro
Axon Axon tem\ill.:11
·---.. Im-=--=----_---__ _ _,-'

\ --0. 1 ,(IO-O m•)
.. · ·"' I "' · ·· · -.
entry locally depolarize& axon. ..

ii ii ii O ii 11 ii ij [jF ltfl II II 11 ii U ILJLJI ii II Ii ii fl II U ii o II
... which sufficiently depolarize& the

adjacent region of the axon to.open
more of the volta,ge--gated
cfumru:>ls, re<re,1ting the action
potential there.
K'
1MJ=1MMt1Mt1J n=o::u::u::n:::,l 't't""USiSAHr-o--o--u--u-u- The> proceli6 continues down the
length of the axon.
K'
FIG URE 1.28 An action pot,mtial (tiring) of a neuron is cnmted wh911 the membran9
o f th.g, neurcn pe<mits sodium ions (Na*) to rush into the cell . thus lncrnasing the volt-
ag9. V9ry quickly aft9rvvard, pot assium (K•) flo\NS out of the oell, bringing voltage
back to resting voltage. This process oocurs along the length o f the axon until the
action potential reaches the axon terminal.

24 CHAPTER 1
outside. TI1en the membrane very quickly changes again in a way that pushes
positively charged potassiwn ions (K+) out of the axon, restoring the neuron
to its initial resting voltage.All of this-sodium in and potassium. out- occurs
in about a thousandth of a second every time a neuron fires.
Because even the biggest axons in mammals are much, much smaller
than the giant squid axon, it i.s difficult to insert an electrode inside a neuron.
Usually we measure electrical changes from just outside ma1IU11alian neurons
(Figure 1.29). By measuring different aspects of neurons firi.ng, we can learn
about how individual neurons encode and transmit information from sense
organs through higher levels of the brain.
One waytoinvestigatewhata neuron.encodes is to try to identify the stimulus
that makes it fire the most vigorously. For example, a neuron in the primary
visual cortex (see Chapter 3) might respond best to lines that are vertical, less
FIGURE 1.29 N9Urosci9ntlsts rQCOrd
to lines tilted to the left or right, and not at all to horizontal lines. When you
the activity of slnQle nrurons with
electrodes close to the axons. Mod-
get to the auditory system in Chapter 9, you will see th.at the precise timing
ern tgchnlques p9rmit the lnsgrtlon of of action potentials can be as important as the number of action potentials.
mu!tlple el9ctrodeis to measure activ- Sometimes we learn a lot by simply finding the threshold for getting a
ity in multiple ngurons so that we can neuron to fire. For example, you will learn that different neurons in the auditory
better understand hO'N neurons \l\l'Or1< nerve, on the way from the ear to the brain, are most sensitive to particular
In concart when encoding sensory
Information.
frequencies of sounds. In Figure 1.30, you can see curves obtained by measur-
ing responses of six different neurons to sounds of different fre<Juencies and
intensities. These are called tuning curves because they show how patterns
of neural firing are selectively LltWled" to different frequencies that vary from
0 to 50 kilohertz 100J hertz, or 1CXXJ cycles per second; plotted on the
x-axis). TI1e minimum sound intensity that is required for a neuron to respond
is shown in decibels (dB) on the y-axis. Thus, the best frequency for the neuron
neurolmaglng A set of methods
that generate Images of the structure
plotted in red in Figure 1.30 is about 1.3 kHz. To get that cell to respond to
and/or function of the brain. In many another frequency, the sound will need to be louder. The mange p lot is for a cell
cases, these methods allow us to that responds to a completely different, markedly higher range offrequencies.
E!XM1lne the brain In IMng, behaving
humans. Neuroimaging
electroencephalography (EEG) Modem perception researchers use other tools to understand how thousands
A technique that, usng many elec- or millions of neurons work together within the human brain. In many cases,
trodes on the scalp, measures electri- we look at the results of these methods by making pictures of the brain that
cal activity l'om populations of many
neurms In the brain. reveal its structure and hmctioru. TI1ese methods can be collectively referred
to as neurolmaglng methods. Of course, neuroanatomists have long described
the structure of thehwuan brain, and neuropsychological studies of individuals
with brain damage have tilught us mud1 about the functions
of different parts of the brain. However, a great advance in
.,,'" -JD recent decades has been the advent of neuroimaging meth-
.5 ods that allow us to look at the structure and hmction of the
Jl -40 human brain in heal thy, very much living human ob.servers.
j -60
For example, electroencephalography (EEG) measures
electrical activity through dozens of electrodes placed on
l1l
-!D
the scalp (Figure 1.31a ). EEG does not allow researchers
to leam what individual neurons are doing or to pinpoint
the exact area of neural activity. However, EEG can be used
§ -10'.l to roughly loca.lize whole populations of neurons (Figure
JI 1.31d} and fb..measure tfieir adivJties with excellent tem-
0.2 0.5 10 20 50 paral accuracy.
Frequency af tone stimuli {kHz) Like a single behavioral measurement, a single EEG signal
FIGURE 1.30 Tuning curves of auditory neurons. See the recorded for a sing.le event is usually not tembly informative.
tQXt for detalls. {After Br&edlove, Watson, and RoS9nZ'IN9ig, U you want to know the brain's response toa prick.of the skin
2010.) or a flash of light, Yo" record the respanses to many, many

INTRODUCTIOI" 25
(b) EEG segments
St imuh.us onset
. r...
(c) Averaged ERP wavefom\
(•)
Stimulus onset
!
P3
200
T1me(ms)
Time (ms) after s timulus pro.>se:nldtion
(d)
50-75 nts 75-100 ms 100-125rns 125-"ISOms

FIG URE 1.31 Electroencephalography, (a) Electrical activity from the brain c an be recorded 1rcm the
scalp by using an array o f electrodes. (/))The activity frcm one electrode is quite variable. even if the
sam9 s timulus (PQrhaps a fl.:lsh o f light) is pr'6Gnt9d multiple tirnGS.. (c) HoW"9VQr, if many such signals
are alJ€ct"ag9d, a regular pattern of positive (PJ and negative (N) IMN9S can 00 seen. Elec tri-
caJ actMty {Voltage) Is measured In m b rovolts (µV). (d) Different signals frcm different electrodes can be
us9d to create a rough map o f scalp topography o f thie actMty gJicited by a stimulus. HQl'"e Is an illus -
tration of the t }'Pical changes one see from 50 to 150 ms aftflf' the onset of a visual stimulus.
Not be that th9 signal is lnitiallyfocuS9d over 9ar1yvisual cort9X (50- 100 ms), then shifts to more ante·
rior areas of visual cortex (100-126 m s) and then begins to activate 1rontal cortex (125-1 50 ms). (Parts
lt-C after Breedlo\"9, Watson, and Fbsenzwelg, 201 O; d courtesy o f Steven Luck.)
event"related potenUal (ERP)

A rneasure of electrical activity from a
repetitions of that s timultLs (Figure 1.31 b). You then avernge all the responses subpopulatlon of neurons In response
alig ned to the moment that the s timulus present. The resulting averaged to particular stlmull tha1 requires aver-
waveform is known as .an event-related potential (ERP) (Figure 1.31 c). For aging many EEG reccrdings.

26 CHAPTER 1
FIGURE 1.32 Ev9nt-ri91ated pot911- 15

t iaB (ERPs. measured in microvolts,
µV) produced in response to viery briEif
flashes. o f llght. (After Yee.il)'1Jrt et ol .,
10
2009.)
0 5-0 100 300

magnetoencephaloSTaphy The event
Time(ms}
(MEG) A tsohnlque, similar to elec- happens
troencephalography, that measures attimeO
changes In magnetic activity across
p:pulatlons of many neurons In the
brain. example, Figure 1.32 s hows the ER.Ps from an experirnent in which observers
saw veJy brief flashes of light lasting for 0.1 toS milliseconds (ms); a millisecond
is a thousandth o f a second. A flash of each duration wets sh own to the observer
400 times. 11-ie EEGs for the 300 ms after eadl flash were averaged together to
FIGURE 1.33 (a) The ffi.'.l'isive device produce the i:n the figme. Fo r the millions of
used for n ons-c'dntributing to the partic ular signal, no thing happens for the first 50
(MEG), What looks likeahugehQ!met ms or so. Then there is a s mall negative deflection of the signal, followed by a
contains superoonducting magnets. '1;>)
larger positive deflection. You c.:ui. see how this sort of recording could tell us
The output of MEG reoordlng can be
used to \'lsuallze acttvlty In th& brain. 1n about the timing o f nen-ous sys tem responses to stim uli in the world.
this cas9, th9 indMdual saw pJctures of A related me thod known as magnetoencephalography (MEG) (Figure
obJ€ci:s ...Ho tterRcolors lndlcata mor9 1.33) maint.lins a good measure of the timing of populations of neurons
actMty. The hot spot at the back o f the while providing a be tter idea of w here in the brain neurons are most active.
brain is th9 primary \'isual cortex. 1119
floating squares represent the array
MEG takes advantage of the fact that neurons make very smaJI changes in
of MEG detectors. (Part b courtesy of their local magnetic fie lds in addition to s mall electrical changes. \ Vith MEG,
Daniel Baldauf.)
(b) Reconstmction of brain responses to a \ris:i.1.tl stimulus

INTRODUCTION 27
research e rs use extrem ely sen s iti ve devices to m ea s ure

th ese tiny magnetic field changes. \.Yh.y use EEG if you
can have lvffiG's better s patial resolution? EEG recording
is re latively simple and relatively cheap. fvrEG devices use
s upe rconducting quantum interference devices (SQUIDs,
but different from the squids in Figure 1.27). These are much
1nore expensive and compl ex.
As noted above, one of the most exci ting and irnportant
chan ges in neuroscience in the last generation has been the
advent of methods that enable us to see the brain w hile it
is still in ib )Ve had X-rays for
over 100 , but an imttge like the X-ray in Flgure 1.34
gives only a ghostly impression o f soft tissues like those
makin g up the brain. Compare that image to the images in
Figure 1.35. Figure l.3Sa shows a computed tomography,
or CT, image. To create CT images, s tandard X-rays are used
in a different ma nner. A beam of X-rays is passed through
the target-the head in this case. The d evire e mitting that
beam spins around the head , sending the beam on many
paths throu gh the sarne "slice" o f the head . This process
is repeated for many slices, until the whole hea d has been
FIGURE 1.34 A standard X -ray does
sca nned. Each time the bean' passes through the hea d, some amou nt of the
not f-:fovide muc h o f a look at the struc-
emitted energy is absorbed by the interven ing tissue. Dense tissue absorbs ture of the brain.
more energy li ghter tis.sue. The detector on the o ther side of the head
measures the amount of energy tha t has been lost on the way through the h ead.
If yo u have this information h orn many, man y passes through the head from
many diffe rent posi tions, you 1.."'an put it together to create a three-dim en sional
picture of the head.
Figure l.35b shows an image produced by magnetic resonance Imag-
ing, or MRI. us.es a very different method to m a ke its often spectacul•f f
images. The brain a nd its own er are p laced in a ve ry powerful magnetic
(a) CT (b) MRI
computed tomogr"'hy <en An

Imaging technology that uses X-rays
to create Images of sllces throUJh
volumes of material (e.g .. the human
t:od)?.
magnetic resonance Imaging
FIGURE 1.35 CT and MRI. {a) This CT image is o ne horizontal throug h the (MRI) An Imaging technology that
head, from the gyes at the top o f the im;)Qe to the bac k of thQ hQad . The dark arQ....-,s uses the responses of atoms to strong
are the fluid-filled ventricles in the brain, the bright white Is b one. and the g ray is the magnetic nelds to fonri Images of
brain its&lf. f.P) Thls s tructural MRI image shows a vertical slice through the middle structures like the brain. The method
of the heed. You c an S99 the folded cortsx of the brain. the splnal cord descending can be adapted to measure actMty In
down the neck, and rnuc h m::>re. the brain. as well.

28 CHAPTER 1
field that influences the way the a toms spin. The physics is complicated and
beyond the scope of ou r text, but by pulsing the magnetic field, it is p ossible
to m easure a signal that can indicate the presence of specific elemen ts in the
tissue. lf you ask about h yd rogen in the brain, you a!'e (mostly) as king about
wa ter, and yotu cornputer can use that s ignal to recons tn1ct the s tl'ucture of
the water-rich tis.sue in sid e your head.
Fo r the study of the senses and, indeed, for the s tudy of ma ny topics in
psychology, the m ost remarkable use of MRI technology is kn mvn as functional
magnetic resonance imaging, or fMRI . With we can see the activity
of the li ...i n g brain.. Here the critica l factor is that active brain tissu e is hLmgry
brain tissue. It needs oxygen. Oxygen and other s upplies are delivered by the
blood, so a n active brain d em ands rnore blood. The result is tha t there is a
blood oxygen leveklependent (BOLD) signal that ca n be measured by tl'e
MRl device. Ins tead of indicating the presence of water, the m agnetic pulses
and recording a re used to pick up evidence of the demand for more resources.
There are sorne dm,vbacks. It takes a fe\o\-' seconds for the BOLD signal to rise
after a bit o f bmin becomes rnore active, so the temporal resolution of the
FIGURE 1.36 Functio nal MRI. The method is quite s low compared with EEG / ERP methods, for example. The
warm colors show places where th9
machines are noisy, making auditory experiments difficult Moreover, acquiring
BOLD signal was .el1Nat9d by the pres-
ence of a visual stimulus. Blues show and running these madtl.nes is expensive. Still , none o f these drawbacks hcive
decreases in BOLD rtcUvity. The big preventa.1 the method from revolution izing the study of the brain.
region of si;Jnal elevation corresponds Figure 1 .36 s h ows some of the results from a very bask fMRI experirnent.
to the pt"irnary visual areas of the brain The observer was \vatching a screen. A visual sti mulus was o n for30 seconds
(SGO Cl>apter 3).
and off for 30 seconds in a ltemation. Regions of the brain that are made rnore
active by the presen tation of a visua l sti mulus dernand m ore supplies in
response to the presence of th e s timulus, so the.re is a difference between the
BOLD responses to stimulus-on and stim ulus-off conditions (wi th th.ci t lag of
several seconds)_ Subtracting the stimulus- off fMRI signal from the stimulus-
on signal reveals the difference in BOLD responses. Typically, these are sh mvn
as colors superi m posed on a s tructural r.tt.Rl of the
Positron emission tomography is an imaging teclmique in whid1
a small amount of a biologically active, radioactive mater ial (a tracer) is
introduced into the participant's b loodstream, and a specialized camera detects
gamma rays emitted fr om brain regions where the tracer is being used most
(Phelps, 2000) (Figure 1.37). Th e logic is similar to that o f fMRl: the idea is to
functlonal magnetic resonance
Imaging (IMRI) A variant of mag-
detect activity in neurons by looking for increased metabolic activi ty. lhe m ost
netic resonance Imaging that makes com mon tracer used in perception experi1nents is oxygen-15, an unstable form
It pooslble to measure bcallzoo pat- of oxygen that has a half-life of only<'\ little 1nore than 2 minutes. Neurons
terns of activity In the brain. Activated that are most active in the brai n s hould have the greatest req uirem ent for
neurons provcke Increase:! blood fiow. this oxygen. Although PET is a somewJ1atinconvenien t method because you
which can be quantlflecl by measuring
have to inject a tracer, it ha.s theadvaritb.ge of-being whichjs helpful in
changes In the response of oxygen-
ated and dooxygenatoo blood to s rudies o f brain activity related to hearing.
strong magnetic Modern labs o ften use multiple methods in the same e>..-perimen t o r series
blood oxygen level-dependent of experiments. You might use behavioral methods to determine the nature
(BOLD) signal The ratio of oxygen- of a particular perceptual but s uch methods mjght take hollfS and
ated to deoxygenated hemoglobin that htrndreds or thousands of tria ls. If your observers can't spend that much
permits the lccallzatlon of brain neu- time in the scanner, you might h ave them perform a shorter, stripped-do\vTl
rons that are most In a task. version of the study while being imaged. You might do yet an o ther version
posrt:ron errisslon tomography of the task, specialized for the de mands of BEG recording. By means of these
(PET) An Imaging techndogy that "con vergin g operations," you co uld build up quite a d eta iled picture of what
enablee us to delne locations In trn perceives and how her brain gives rise to those perceptions.
brain where neurons are especialty
actr.<e by measuring tlie metabolism Now tha t yo u've compl eted this brief tour of the methods of the trade, you
of brain cells using safe radloactWe .sho uld be ready to ern.bark on n m ore comprehensive examina.t.ion of sen.""3tion
lootopes. and perception. We h ope you enjoy the journey.

INTRODUCTION 29
Looking Li stening Thinl<ing Remembering \Vorking
FIG URE 1.37 Positron enlssion tomography (PE'l) is a form of nooroimaging that
uses p ositron-emitting radioisotopes to create images of biological proc9sS9s in the
living brain. Here we seg difftf"ent patterns of gluoos9 usage in the brain while an indl-
\'idual is peirforming dlffE:Vent tasks. (From Phelps, 2000.)
Summary
1. Sensation and perception are cen tral to, and often pre:.:ede, almost <ill
aspects of human behavior and thought. There are many prachcal applica - f1efer to the
tions of o ur increased understanding of sensation and perception. Sensation and Perception
2. G u stav Fechner invented several clever me thods fo r measuring the relation- Companion Website
ship beh'leen physical ch<inges in the world and consequent psychological snes.slnauer.comtwotte4e
changes in observers. T11ese methods remai n in u se today . Using Fechner's for aotivities, essays, study
researchers ca n measure the smallest levels of shmulus that can
be detected (absolute threshold) and the smallest differenCES tha t can be qwslons, and otrer study aids .
detected (difference thresholds, or just no ticeable differences).
3. A more recent development fo r understanding performance--signal detec-
tion theory-permits us to si mulate changes in the perceiver (e.g., internal
noise and biases) in order to under.stand perceptual performance better.
4. Studying simple stimuli (such as pure to nes) is usefu l because complex
stimuli can be decomposed into simple components. In vision and audi tion,
Fourier analysis is a mathem<itical tool thi.lt helps resei.\rch ers break d mYn
complex images and sollllds in ways that permit better understanding of
how they are sensed and perceived.
5. We leam a great de..-=tl about perception by understanding the biological
structures and processes involved. One early observation-the doctrine of
specific nerve energies---e:r..-presses the fact that people are aware o nly of
the activity of their nervous systems . For this reason, wh at matters is \vhich
nerves are stimulated, not how they are s timuL."'l.ted. The central nervous
system reflects specializations for the senses from cranial nerves to areas of
the cerebral cortex involved in perception.
6. The essential act ivities of all neurons, including those involved in sensory
processes 1 are chemic.al and electrochemical. Neurons communicate w ith
each oth er through neurotransmitters, molecules that cross the synapse
horn the axon of one ne uron to the dendrite of the nex t. Nerve impulses are
electrochemical; voltages change along the axon as e lectrically charged sodi-
um and potassium ions pass in and out of the membranes of nerve cells.
7. Recordings of individual neUTQnSefiablE! 'us the lowest level of
sti mulus required for a neuron to fire (absolute threshold). Both the rate and
the timing pattern of n eural firing provide additional infomiatio n about
how the brain encodes stimuli in the world .
8. Neuroimaging methOOs have revolutionized the .s tudy of sensa tion <ind
perception by allowing us to study the brain in healthy, living hum<in
observers. Useful m ethods include e lectroencephalogmphy (EEG), mag-
netoencephalography (MEG); positron emission to mography (PET), and
functional magnetic resorumce imaging (fMRI). Each comes with its 0\"'11
combination of temporal and spatial properties, making one m ethod sui t-
able for researching some ques tions and other methods more suitable for
o ther questions.

CHAPTER 2

The First Steps in Vision:
From Light to Neural Signals
WITH THE NAKED EYE WE CAN SEE STARS tha t are up to about 20)() light-years
(al most 6 trilli on miles) away, and in a dark winter sky far frorn ci ty lights the
neighboring galaxy, Andromeda, is visible at over 2 million light-years away!
This chapter describes the first s teps in seeing. To understand h ow we see,
we must fi rst consi der a little physics and optics, and then we' ll look at how
the eye is built to cap ture and begin to process light.
In C hapters 3-8 we' ll see how light in forma ti on g lea ned by the eyes
travels bac k throug h the hea d to the brain, and h ow the brain transforms
this information into a meaningful inte rpretatio n of the ou tside world. Fo r a
preview o f the e ntire visua l process, ""·ork throu gh Web Activity 2.1: Visual
System overview.
A Little Light Physics

Light is a fo m1 of radiation--en,e rP!' produced by v1'brations
of eJectrically charged materiaL There are twb way.s 'f o conceptualize lig ht:
as a wave or as a strea m of photons, tiny particles that ead1 consist of one
quantw11 of energy. This dual nature of light can be confusing to physics and
psychology studen ts a like. ln this discussion we11 try to avoid confusion as
1nuch as possible by treating light as being made up of wava w hen it m oves
ar ound the world, ,1nd being made up of pho tons when it is absorbed.
Althou gh the foll spectrum of electromagn etic radiation is very wide, light
makes up only a tiny portion of this spectn.un. Figure 2.1 illustrates the electro-
magnetic spectrum, from gamma rays (which have ve.iy shorh1hweleng ths) to
rad io a nd te levision waves (which have verv long wnvelengthsJ . Visible light
waves have wa velengths between 400 and 7oo nanometers (nm; 1 iun = ] Q-9
meter), as illustrated on the bottom of the figure. Note tha t as the wavelen g th
v,uies in the visible spectrnn"I, the color \.\'e observe changes, from violet at wave An oocUlatlon that travels
about400 run through the wh ole spectrum of the rainbow up to red a t about thrrugila medium bytransferrln;i
650 run.. (As v1,.-e' ll discuss in C hapter Sr the light waves themselves energy 1rom c.ne particle or point to
another without causing arr/ perma-
are n ot colored; it is only after our visual systen1 interprets an incoming wave nent dlsplaoement of the medium.
that we perceive the light as a specific color.)
photon A Q'"1ntum of visible light cr
Let's consi der l-vl"la thappens to li ght on its \.•.:ay from a s tar to an eye. (See Web other form of electromagnetic radiation
Activity 2.2: From Sun to Eye.) In empty space the electromagnetic radiation demonstrating both partic le and wave
from a star travels in a s traight line at the speed of light (about 186,000 miles propertloo.
per second). O nce it readi.es the atmosphere, som e of the starlight's photons absorb To take up something -
will be absorbed by en counters with dust, vaporized 'va ter, and so on; and such as light. noise. or energy-and
some of the light '"'·ill be scattered (sometimes ca lled diffracte d ) by these not transmit It at all.
particles. Most of the photons, however, w ill make it through the atrnosphere scatter To dlsperse sornethlng-
and wi ll eventually hit the s urface of an object such as light-In an Irregular fashion.

32 CHAPTER 2
1Navdmgth (nm)
FIGURE 2.1 Sp9ctrum of
slectromagnetic ent1rgy (specified in 1012 "10 11 10LO l(f lcf-1 ID' -io' 105 10' 10' lO' 10 I 10- 1 10-2 10....)
spect1um
nan om etQ'"s), with the visi
.,,
(400-700 nm) expanded at right. Note
that 1 nrn:::: 10-9 m. ..
§ 0
:;
1 :;
if j ]
!l
"
,);
••
70) 675 650 575
••••••
550 525
(nm)
500 475 450 425 400
If the ray of s tarlig ht were to strike a li ght-colored s u rface, m ost of the

light would be reflected. Indeed, the fact that most of the light bounces off
the surfuce accounts for that surface's "light" appearance. Howeverr mos t of
the light striking a dark surfa ce is absorbed. Light that is n either reflected nor
absorbed by the s urfa ce is transmitted through the surface. If \<Ve are gazin g
at our s tar through .1 \'\indow as the light travels from air into the glass, some
of the lig ht rays \1,.i ll be bent, or refracted, as it is transmitted.
Re fraction also occurs when lig ht passes from air into water or into the
eyeball. ln foct, the part of an eye exa m in which the eye d octor checks the
s prescription is often called a "refraction" because the doctor determines
how much the light must be bent by eyeglasses for it to be properly focused
on the retina. In the next section we1l see how the optic sys tem of our eyes
performs this same kind of focusing.
Eyes That Capture Light

ln order to see sta rs or a nything else, \•ve n eed some type of physiological
mech anism for sensing light. Even single-celled organisms s uch as £lmoebas
respond to light, changing thei r d irection of m otion to avoid bright light when
it is But eyes go well beyond m ere light detection. An eye can fom1
reflect To redirect somett>ng that an Image of the outside world, enabling animals that possess eyes to use light
strikes a surtace-especlaly IQht, to recognize objects, not just to de termine w hether lig ht is present and wha t
samd, or heat- usualy back toward direction it's coming from.
Its p:jnt of origin. Before explainin g how eyes form images, let's take a tour through the
transmit To corNflY somethlrg hum an eye to become fa miliar with its important parts. Ftgure 22 shows a
(e.g., frorn cne place or thlr>J to front-to-back s lice through a hmnan eye, \<Vi.th the m ost imp ortant struch1res
an::ither.
labeled . (S<!e Web Activity 2.3: Eye Structure.)
refract 1. To alter the course or a 1l1e first tissue that light from the star wi ll encmuiter is the cornea. Contact
W@Je o f energy that passes Into some-
lenses sit on a thin film o f tears in front o f the cornea. The cornea provides a
thing from another medium. as water
does to llgl1t entering It frorn the air. window to the world because it is transparent (tha t is, 1no.st light photons are
2. To measure the degree of refraction transmitted throu gh it, rather than being reflected or absorbed). It is trans par-
In a lens cr eye. ent because it is made of a highly ordered arran gement of fibers :cmd because
image A picture cr likeness. it contains no blood vessels or blood, \vhich would absorb light. The cornea
cornea The transparent •wlrdc:w" d oes, h0\>1.1ever, have a rich s upply of transpa rent sensory nerve endings,
Into the eyeball. which are there to force the eyes to close and produce tears if the cornea is
scratched 1 pteserving its trans parency. lf you have ever scratched your cornea
transparent AJ lowlng light to pass
thrcugh w ith no lntefruptlon, so that or ""·om contact lenses too lon g1 you know exactly how painful this can be!
objects on the other side can be Forhmatel)·i the external layers of the cornea regenera te very quickly, so even
clearly seen. when a cornea is scratched, it usually heals wi thin 24. hours.

THE FIRST STEPS IN \.1SION 33
FIG URE 2.2 The human right fil'/9

in c ross sectbn Mewed from above).
Note that the R on the retina B
revgsed right to l.eft, and it ls upside
down . The in the retina wherg
the optic neirve leaves tM 91eball is
the optic disc {where the absenc e o f
photoreceptors results In a blind spot}.
(After Breedlove and Watson. 2013.)
Blood vei;sel.s
The aqueous hlll1or,a fluid derived from blood, fills the space immediately
behind the cornea, supply ing oxygen and nutrients to, and removing waste
from, the cornea and the lens. Like the cornea, the lens has no blood supply,
so it can be comple tely transparent; a nd, as we' ll see later, the shape of the
lens is controlled by the ciliary muscle.
To ge t to the lens, the Light from our st.ar must pass through the pupil, which
is simply a hole in a muscular s tructure called the Iris. The iris gives th e eye
its distinctive color, and controls the size of the pupil, and thus the amount
of light that reaches the retina, via the pupillary light reflex. \'\!hen the level
of light increases or decreases, the iris '1Utomatically expa nds or contrai..."'ts to acp.Jeous humor The watery fluid In
allow m ore or less light into the eye. the anter1or chamber of the f1'10.
After p assin g thro ugh the lens, our starlight '"rill ente.r the vitreous dlmn- lens Tue lens Inside the eye that
ber (the space be hveen the lens and th e retina), where it will be refracted for enables the c hanging of focus.
the fourth and final time by the vitreous humor. Th.is is the longest part of pupil The dark. circular cpenlng at
the jo urney throu gh the eyeball; th.is ch..1mber comprises 803 o f the intemal the cent"' of the Ins In the fft9, where
volume of the eye. TI1e vitreous is gel-like and viscous (a bit like egg w hite), light enters the fft9.
and generally transparent While s taring up a t the bright blue s ky on a lazy Ins The colored part of the eye,
sunn y day, however, you may have noti ced " floa te rs," s mall bits of d ebris consisting of a muscular diaphragm
(biodebris) that drift around in the vitreous. Floaters are quite common, and surrrundlng the pupil and re>gulatlng
the Ilg ht enten ng the eye by expardlng
they are us ually n ot a cause for concern . and contracting the pupil.
Finally, after traveling through the vitreous chambe.r, the light emitted by
our favorite s tar will (hopefo lly) be brought into foc us at the retina. To be a vitreous humor TI'e transparent
fiuld that Ills the Wreom chamber In
bit mo re precise, only some bf d>e fighi J i11 actually-\,,.1ch the retina. Much the posterior part of t11e eye.
of the light energy will have been lost in space or the atmosphere, becc1 use of
retina A light-sensitive membrane In
absorption and scatterin g. as d escribed already. In addition, a good deal of the bac k of the "fe that =italns rods
light becomes lost in the eyeball, so only about half of the that arrives and cones, which recetve an Image
at the cornea actually reach es the retin..1 . The role of the reti na is to detect frcm the lens ard sand It to the brain
light and "te ll the brain about aspects of lig ht that are rela ted to objects in the thrrugh the optb nerve.
world" (Oys ter, 1999). [n othe r words, the retina is w here seeing really begins. ernmetropla The condition In which
there Is no refractive error. because the
Focusing Ught onto the Retina refractive pow... of the eye Is perfectly
matched to the length of the eyeball.
To foc us a dis tant star on the retina, the refractive power of the four optic
components of the eye mus t be perfectly matched to the length of the eyeball. diopter (D} A unit of measurement
This perfect matd1, known as emmetropia, i s illus trated in Figure 2.3a . The of the cptlc P'.>W<ll of a lens. It Is equal
to the reciprocal of the focal length. In
average eyeba ll is abo ut 24 millimeters (mm) lon g a nd h..1s a power, when meters. A 2-dlopter lens Wiii bring par-
unaccommo dated (focu sed on a dis tant object like our star), of about +60 allel rays of light into focus at 16 meter
diopters. (We'll explain what accommodation means in a bit.) (50cm).

34 CHAPTER 2
0
.
G>l FIG URE 2.3 Optics o1 the human gye. See the text for details. {After
Oyster. 1999.)
Refractive errors occur when the eyeball is too long or too short relative
to the power of the optic components. If the eyeba ll is too long for the optics
(b) M yopi.:l (Ftgure 2.3b), the image of our star will be focused in front of the retina, and
the star will th us be seen asa blur rather than a spot of li ght. Th is condition is
called myopia (or "nearsightedness"). Myopia can be corrected with negative
(mi nus) lenses, wh ich di verge the rays of starlight before they enter the eye
(Figure 2.3c). ll the eyeball is too s h or t for the op tics (Figure 2.3d), the image
of our star will be focused belrfnd the retina-a condition rnlled hyperopia
(or "fars ig htedness"). If the hyperopia is not too severe, a youn g hyperope
ca n compensate by accommoda ting, the reby increasing the power of the eye.
If accommodation foils to correct the hyperopia, the star's image ,,,,.ill again
be blurred. Hyperopia can be corrected vdth positive (plus) le nses, w h ich
con verge the rays of s ta rli ght before they en ter the eye.
On average, the adult human eye is 24 mm long, abo ut the diameter of
a quarter. However, eyeballs can be quite a bit longer or s horter and s till be
ernmetropk because eyes generally grow to ma tch the pmver of the optic
components we're born with. (Most newborns .u'e' hyperopk because the optic
components of their eyes are relatively well developed at birth compa red with
the len gth of tl1eir eyeballs.)
TI1e m ost powerful refracting surface in the eye is the cornea, w hich con-
lTibu tes about two-thirds of the eye' s focusing \Vhen the come.a is no t
sphericat th e result is astigmatism. With astig matis m, verti ca l lines might
be focused sli ghtly in front of the retirn::i1 w hil e horizontal l.ines are focu sed
myopia Nearsightedness, a com· slightly behind it (or vice versa). If you h.:1 ve a reasonable degree of uncorrected
rnon cordltlon In which light entering astigrnatis m, one or rnore of the lines in Figure 2.4 might appea r to be o ut of
the f!!Ye Is focused ln frcnt of the retina foc us whil e other lines appear sharp. Len ses that have h-vo focal poin ts (that
and distant objects cannot be soon
sharply. is, lenses that provide different amounts of focusing power in the horizontal
and vertical planes) can correct astigmatism.
hyperopla Farsightedness. a com-
mon cordltlon In which light entering So far, we've considered the image of a distant object. But wha t happens
the eye Is focused behlrd the retina when \Ve wa nt to focus on some thing dose by (like the words on this page)?
and accornrnodatlon Is required In Remember that refraction (light bending) is n ecessary to foc us light rays.
crder to see near objects clearly. Because the cornea is highly curved and has a higher refractive index tha n air
astl9'Tlatism A visual defect caused (L376 versus 1)1 it for ms the most p owerful refractive s urface in the eye. The
by the unequal of ore or more aqueous and "itreous humors also help refract light. However, the refractive
of the refractive surtacoo of the eye, power of each of these three structures is fi.xed, so they cannot be used to bring
usl.Blly the cornea. close objects into focus. This job is performed by the lens, which can alter the
accommodation The process by refracti ve power by cha nging its s hape-a process called accommodation.
which the eye changoo Its focus (In Accommodation {change in foc us) is accomplished through contraction
w111c11 the lens gets falter as gaze Is
directed toward nearer objects). of the cil.iary muscle. ll1e lens is attached to the ciliaiy muscle th.rough tiny
fibers (knm'\-T\ as the zonules of Zinn) (see Figure 2.2). When the ciliary muscle
is relaxed, the zonules c'.\re stretched and the lens is relatively flat. ln this s tate,
the eye w ill be focused on very distant objects (like our s tar). But to focus on
something a \-Vristwatch-the ciliary muscle mus t contract. This
contracti on red uces the tension on th e zonules and enables the lens to bulge.
The fatter the lens is, the m ore p ovver it has.
FIG URE 2. 4 Fan chart for astigmaJ:ism . Tai<Q off your glasses Of you wear glasses)
and IJl.ew this If you have a significant degree of astigmatism. on9 or more of th9
lines w ill appear to lower oontrast.

THE FIRST STEPS IN VISION 35
FIG URE 2 .5 The precipitous drop in amplitude with age. Each

sym bol represents data 'from a different study. ThoB gray dashed Ung indlcafos the
amplitud€.I of accommodation required t q focu,S at a cistance 40 cm. Each colored
s;irnbol repres9nts data 'from a different cias:::Jcal" study.
4
Accommodation enab les the power of the lens to vary by as much as 15

diopters. Note tl10t lens power (P) 11f, where f is the focal in meters.
So., if your unacconunoda ted eyes were perfectly corrected for distant vision,
15 diopters o f accommod a ti on would enable you to read your watch at a
dis tance of about 0.067 meters (1/ 15) or 6.7 centimeters (crn.; to con vert me ters
to centimeters, simply multiply by 100). lf you can read your watch at 6.7 cm
(while wearing your d istance correction ), you are either very lucky or ve:ry
young. Our ability to acconmux l.ate declines 'vi.th age, starting from about 8
years old, and we lose about l diopter of accommodation every 5 years up to
age 30 (and even more after age 30). By the time most people are be h.veen 40
and 50 years old, they find that their arms are too sh ort because they can no
longer easily accommodate the 2.5 diopters or so need ed to see dearly at 40
c n1 (1 / 0.4 = 2.5). Titls co nditi on is call ed presbyopia (meaning "old sight''),
and it is, like dea th and taxes 1 inevit.1ble! Figure 2.5 illustrates the precip itous
drop in accommodation with age.
Why d o we all h ave presbyopia to look fonva rd to? The main reason is
that the lens becomes sclero tic (h arder) and the capsule that encircles the
lens (enabling it to change s hape) loses its e las ticity. Lu cky for us, Benjamin
Franklin (170&-17'l0) invented bifocals-lenses that have one power a t the
top (permitting us to see distant objects) and a different power a t the bottom
(a llo\."ling us to be in focus a t a comfortable rea ding di.stance).
Like the other optic components of the eye, the lens is n om1ally transpar-
ent. It is hanspcu ent because the crystallins (a class of proteins that make up
the lens) are pac ke d together very densely and therefore are very regular.
Anything that interferes \-vith the regularity of the crystaUins will result in
loss of trans parency (areas are opaque-that is, "opacities''). Opacities
of the lens are known as cataracts. can occm a t different ages and
take man y different forn1.S. Con genit.11 ca taracts (present at birth) are relati vely
rare, but if they are dense (and the refore interfere 'vi th retirui.I image quality),
they can have d evastating effects on normal visua l d evelopn1ent if n ot treated.
promptly. Most cataracts are discovered after age 50, and the prevalence of
cataracts increases vvith age, so by 70 almost e"·eryone has some loss of trans-
parency. Cataracts can interfere \vi th vision because they absorb and scatter
more light than the n ormal lens d oes. trea bnent of cataracts (in
whid\ the opacified lens i s extracted and replal"'ed Y\-'i.th a plastic or silicone
irnplant) h as become qulte routine.
The Retina
The preceding discussion covered h ow the hwnan visual system delivers a
focused ima ge of our favorite s tar onto the retina, w hich is spread across the
back of the eyeba ll. The optics involved (see Figure 2.2) have been likened
to those in mos t cameras, which a lso include a mecha nism for regulating the
amoun t of lig ht (the "stop," analogous to the iris in human eyes) and a lens
for adjusting focal len gth so that both n ear and far objects can be focused on presbyopia "old sight. " The
the fihn s pread across the back of the cam era. Hmvever, this is as far as we age-related la>s of accommcdatlon,
can take this analogy, because the purpose of a cmnera is s irnply to record the which makes It to focus on
image projected onto the film, w hile the purp ose of the hmnan visual system near objects.
is to interpret this image. This is the difference between taking a picture and cataract An opacity of the crystal·
seeing a pictrne. And the proress of seeing begins \-vith the retina, where the lln:> lens.

36 CHAPTER 2
FIG URE 2..6 Fundus of the right eye

o f a human. {From Rodi.::ock. 1998.)
li ght energy from o tu star is transduced into neural energy that can be inter-
preted by the brain.
What the Doctor Saw

transck.lce To convert from one Eye doctors use an instrument cal led <m ophthalmoscope to look a t the back
form of enerw to anothEf (e.g., from s urface of their patients' eyes, w hid1 is the fund us (p lural fund;). (You
"1)ht to neural electncal erergy, o r fran probably remember all too well having tha t bright light s hining into you r
rnechanlcal movement to neural elec-
trical energy). eye while the d octor exam ined your fund us .) Agure 2.6 shows a n orma l fon-
d us. 1l1e white circle is k.nmvn as the optic d isc. This is the point \vhere the
fundus The back layer of the retina:
what tha aye doctor sees through an
arte.ries and veins tha t feed the retina enter the eye, and w here the a.xons o f
cphtl'JalmCSO(lle. ganglion cells (w hich we ""ill get to sh ortly) leave the eye via the optic ne rve.
Th.is portion of the retina contains n o p h otoreceptors, and consequentl y it is
blind . You can experie nce your m<\<·n blind spotr correspond ing to the optic
d.iscr by dosing yo ur left eye, fixatins ,on the fin Figure 2.7a with you r right
(a)
•
FIG URE 2.7 To experlerce your blind
F
•
•
spot. dose your left Qfil, fixating on the
Fin part (a) w ith your right eye. Hold the
book about 15 cm from your eye,
and a:::tJust the distance of the book
fro m your QYQS until the red c irc le dis-
appears. This is your blind spot. Ordi-
narily you are not aware of it, because
the visual system .. fills in" the blind spot (b)
w ith information from the surrounding
arQa If you focate on thei Fin part (/;))
w ith you r tight eye and again adjust the F
distance. w hen the gap in th9 line falls
in your blind spot. it will fill in and you
w HJ sea a continuous red llne.

eye, and adjusting the d istance of the book from your eyes until the red circle
d isappears. The reason you d on 't normally n otice thi s large blind spot in your
visual field is that the ·dsual syste m "fill s it in'' with inform ation from the
surrounding area (Figure 2.7b ).
The hmdus is the only pla ce in the body "vhere one can see the arte ries and
vein s so it provides doctors Vlri th an import..' lnt \vi.n dow on the •,;.ve ll-
being of the bod y's vascular syste1n. The vascula r "tree" sp reads out across
the retina in a characteristic way but stops short of the fo vea (the center of the
bro\l\rn ish sp ot near the center of th e fundus in Fig ure 2.6).
You can see your o""'Tl vascular tree by using a simple trick that requires only
a penlig ht. In a dark room, d ose your eyes and place the penlight against the
outsid e corner of one eye. Holding the p enlight against the eye, gently move
the light around (up and d m\11\, and back and forth). \'Vi thin a few seconds you
should see the shad ows cas t by your blood vessels looking like the
of a tree . We do11 .,.t n ottJU'1 ly beta\. :the b lood vessels move with
our eyes, so their s h..1dows are s tabilized retina l images a nd , as with the blind
sp ot, the visual system fills in behind the m. The m otion o f the penlight makes
the shadows mo,·e, enabling us to see them.
Even when v1evv·ed through an ophth almoscope \•.ri th a lot of magnifica-
tion, the ftmdu s does not provide a detailed view of the retina. To ge t a good
look at the structure of the re tina, we need a photomicrograph (Figu re 2.S),
w hi ch reveals th at the retina is a layered s heet of dear neu rons, about half the
thickn.ess of a credit card (Rodieck, 1998}, with another of darker cells,
the pigm ent epithelimn, lying behind the final layer.
f
Sdera {
P\sm'nt ep>thelicun
Photorecepto r layer l "'] }°'""''S"-'

} Inner &2,Sments
Externcil limiting m embrane =
Outer nuclear layer { } Photorecepto r nuclei
• *
Photo receptor axons
(Henle's fiber layer)
°'"". plexiforn>lay"· { }'
}
U 7
l j)
} Outer synaptic layer
Horizont.11 o::ll
} Horizontal ce lls
Bipol.u
lnnernudm Jar" <ell
} Amac1lne cells
Inn" p1'xifo•= Jar" { ' Q crlls •
Ganglion cell
"'ll lay"· •• Light
.!
{
, aXon
FIGURE 2.8 Photomic rograph of the
retina The section shovvn is near thQ
fovea. Note that the axons in this region
te'"ld to run diagonalty to get them out
of the center o f the fovea. (From Oys-
ter, 1900: micrograph from Boycott
Inner limiting membrane c: and Dowling, 196Q.)

38 CHAPTER 2 8230336 Amria Acea
Rod Cone As we'll see in the next section, together these neurons constitute a mini-
computer that begins the process o f interpreting the information contained in
Outer \o·isu al images. The transduction of light energy into neural energy begins in the
SO!gment back.most layer of the retina, which is made up of cells called photorec:eptors
(see Figure 2$). When photoreceptors sense light, they can s timulate neurons in
the intermediate layers, including bipolar cells, horizontal cells, and am acrine
Inner
segment cells. These neu rons then connect \o'.'l th the frontrnost layer of the retina, m ade
up of gan glion cells, whose axons pass th rough the optic nerve to the brain.
Before we describe the function of these layers, we sho uld a ddress an
obvious question regarding the structure of the retina (see Figure 2.8): Why
are the photoreceptors at the back-that is, in the last layer? This ar ran gement
requires ligh t to pass throu gh the gang li on, horizon tal, and amacrin e cells
before making contact with the photoreceptors. Hov1i'ever, these n eu rons are
Syn'l'tic mostly transparent, whereas cel ls in the pigment epithelitun, \vhich p rovide
te rminal vi ta I nutrients to the photo receptors, are opa que. O n ce we see th at the photo--
receptors mus t be next to both the pig1r1ent epithelium and the o ther neumns,
the layering order makes much m ore sense.
FIG URE 2.9 Aod and conR (From
Rxli9Ck 1008.) Retinal Geography and Function
The retin a con tains roughly 100 million photoreceptors. These are the neu-
rons that capture lig ht and initiate the ac t o f seeing by producing chemical
sign als. The hmn an retina contains at least two types of photoreceptors: rods
and cones. l11e.se two types not only h(1ve diffe rent shapes ('"'·h ich is how
they earned. their namesi see Figure 2.9}, but they have different distributions
across the retina and serve different hmctions. Because retinas have
both rods and cones, they are cons idered to be duplex retinas. Some animals,
su ch as rats and owls, have n10s tl y rod retinas; others (e.g., certain lizard s)
have mostly cone retinas.
Huma ns have many m ore rods (about 90 million ) than cones (about 4-5
million), a nd the two types of cell s ha ve very different geographic distribu-
tions on the retina (Figure 2.10 ). Rods a re completely absent from the center
of the fovea, and their d en sity increases to a peak a t about 20 d egreES and then
declines again. The cones a re most con centrated in the center of the fovea,
an d their dens ity drops off drama ticaJly wi th retin al eccentricity (d i.s tance
fr om the fovea). The fo vea is the 11pit" in the inne r retina that is specialized
for seeing fine detail.
photoreceptor A llght-sensltlve As the photographs of photoreceptors a t different eccentricities in Figure
receptor ln the retina. 2.10 illustrate, cones are also smaller a nd more ti ghtly packed in the fovea l
rod A photoreceptOf specialized for center (0.0 in Figure 2 10). This "rod-free" area (about 300 square microme ters
nlgl1t vision. [µm] on the retina) s ubtends a visual an gle of about 1 degree, and it is directly
cone A photoreoepta specialized behind the center of the pupiL So if we look directly at an object whose image
for daylight vision. fine 'Asual acuity,
and color.
duplex In reference to the retina, TABLE 2. 1
consisting of two parts: the rods and Properties of the fovea and periphery in human viskm
cones. which operate under different
conditions. Property Fovea Periphery
fovea A small pit, near the center Photoreceptor Mostly cones Mostly rods
o f the macula. that contains the high- Bipolar cell type Midget Diffuse
est concentration o f cones, and no
rods. It Is the portk:n of the retina tl1at Convergence Low High
produces the hig hest >Asual ac uity and Receptive-field size Small Large
seNeS as the point o f flx.9tbn.
Acuity (detail) High Low
eccentricity The distance between
the retinal Image and the fwea. Light sensitivity Low Hi gh

THE FIRST STEPS IN "1SION 39
16D 8.0 5.0 1.35 0.0 1.35 5.0 SD 16D
10
TcrnpornJ OUtan.:e from foveal (mm)
0£
20
Visual .m gle
FIGUR E 2.10 Photoreceptor density across the retina The top pane.ls show sMceis
through the photoreceptor innq segmns at different eccentriciti9S (dlstane&s from
the fovea). The grar::fi shows the density of rods and oones plottoo as a function of
from f0\'9a. Note that In the PElfipheral slices, the cones are always the
larger OOls. i;Aft9r Oyster, 1000; rnk::rographs from Curcio et al., 1000.)
is s malle r than 1 degree, the image w ill land on a region of the retina that has
only cones. (How big is 1 degree? rule of thumb, illus tra ted in Figure
2.11 : your thmnb, when viewed a t am1's length, subtends an angle of about
2 degrees on the retina, assuming your thumb is about 2 cm across and your
outstretched arm extends about 57 011 from yo ur eye). Table 2.1 illustra tes
sorne of the funda mental differences in the properties of the fovea compared
FIG U RE 2.11 The o f thumb":

w hen v igWQd at arm 's lgngth, your
thumb subt.cnds an anglg o f about 2
dggrQQs on the r&tina

40 CHAPTER 2
l6Am
wi th the peripheral retina. Most important for us, the fovea has hi gh acuity
an d we use it to identi fy objects, to read, and to inspect fine d e tail. O n the
o ther hand , we use the periphery w hen d etecting and localizing stimuli tha t
we aren't lookfrlg at directly (e.g., seeing a m oving truck out of the "corner
of the eye" ).
TI1e cones become larger and more sparse away from the foveal center, and
the s mall ce lls that appem outside the fovea (e.g., 1.35 in Figure 2.10) are rods,
which a re about the same size as the foveal cones. ln a11 of the micrographs
excep t for 0.01 the large cells are always the cones.
Rod.sand cones operate bes t under different lighting conditi ons: Ro ds
hmction relatively well under conditions of din.1 (scotopic) illumination (whid1
is w hy anitnal.s wi th all-rod retinas are nocturnal), but cones require brighter
(pho topic) illumination (e.g., s unlight or room Ligh ts) to opera te efficiently.
Having an area at the center of the fovea with n o rods means th at under dim
illumina tion the central 1 degree or so around the fovea is e ffectively blind!
Ind eed , practiced. s targazers know tha t it is often easier to s po t a dim s ta r by
looking o ut o f the comer o f one's eye than by looking directly at it. We '"rill
revisit ph otopic and scotopk vision again in 01ap te r 5.
Rods and cones d iffer fun ctio n.a.Uy in another important way. Because all
rods h ave the same type of photo pig ment, they ca nno t signal differences in
color. Ead1 con e, on th e o ther hand , has one of three differentpho topig ments
that differ in the wavele ngths at w hich they absorb light m ost efficientl y.
Therefore, cones can signal information about wavelen gth, and thus they
provide the basis for o ur co lor vision.
Retinal Information Processing

The retina contains fi ve major classes of ne urons: photoreceptors, h orizontal
cells, bipolar cells, amacrine cells, and ganglion cells mentioned in the prev i-
ous section (see Figure 2.8). Let's take a cl oser look at the fonctions of each of
these cell types. (See Web Activity 2.4: Retinal Structure.}
Light Transduction by Rod and Cone Photoreceptors

Both types of photoreceptors consist of an outer segTient (lvhich is adjacentto
the pig1nentepitheli\.m1)1 an lnnersegrnent,and a synaptic terminal. Molecules
called visual pigments are m ade in the inner segment (whid1 is like a little
factory, filled with mitochondria) and stored in the outer segment, w here they
are incorporated into the membran e. Each visual pigment mo lecule consists
outer se"11ent The part of a ph:>- of a protein (an opsin ), the s tructure of v; hich determines w hich \Vavelengths
toreceptor that contains photcplgment
molecules. o f light the pigment molecule absorbs, and a chromophore, w hlch captures
light photons. The chromophore is the part of the molecule respons ible for its
inner segment The part of a pho-
toreceptor that lies between the outer color, and it selectively absorbs specific vvavelengths of light. The chromophme,
segment and the cell nucleus. known as retin.:1t is de rived from vitamin A, \vhich is in hun m anufactured
from beta-carotene, w hich is why your m other told you to ea t your carrots!
synaptic terminal The locat bn
where axons tennlnate at the synapse The opsin and ch romophore are co nnected . Each photoreceptor has only one
for transmission o1infa'matlon by the of the four types of vis ual pigments fol.ind in the human retina. ll1e p igment
release of a chemical transmltt€f, rhodopsln is fo und in the rods, concentrated rnainly in the stack of men'\bra-
chromophore The light-catching n ous discs i..n the outer segment. Each cone has one of the other th ree pig-
part of the >Asual pigments of the ments-which respond to lon g, med.i LUn,and short waveleng ths, respectively.
retina. Recen t evidence su gges ts that there may be a third type of pho torecep-
rtlodopsln The visual pigment found tor--one that ''lives'' a mong the ganglion cells and that is involved in adjust-
In rods. ing o ur bi ological rhytlu11s to m.a td1 the day and ni ght of the externa l world
melanopsin A photcplgment that Is {Ba:ringa, 2002). These photorecep tors are sensi ti ve to the ambient light level
sensitive to ambient light. and con tain the photopigment melanopsln, and they send their signals to the

THE FIRST STEPS IN 41
suprachiasmatic nucle us (SCN), the home of the brain's circadian clock, \<1,r hich pholoactlvation Act"1atlon by light.
regulates 24-hour patterns of behavior and physiology. hyperpolarlzatlon M Increase
When a photon from our fuvoritestarmakes its way into the outer segment In membrane potential such that the
of a rod an d is absorbed by a 1no lecu.le of rhodopsin, it transfers its energy to Inner membrare surface becomes
the duomophore p orti on of the "isual pigment molecule. This proress, kno\>vn more than the outer mem-
brane surface.
as photoacttvatlon, initiates a biochemical cascad e of eventsevenhtally resul t-
ing in the closin g of channels in the cell m embrane that normally allow ions graded potential An electrical
potential that c an vary contlnuou'1y In
to flow into the rod outer segrn.ent. Closing these channels alters the balance arnpll!IJ:je.
of e lectrical current between the inside and ou tside of the rod o uter segment,
making the inside of the cell more n egatively d'lc-u ged. Tilis prcx:es.s is k.nmvn as
hyperpolarlzatlon. Hyperp obrization closes calcium channels at the .synaptic
terminal, thereby reducing the concentration of free calci um inside the rells.
The lowering of the calcium. concentrati on, in him, reduces the concentration
of neu rotransmitter (glutamate) m olecules at the synaptic terminals, and this
change signal s to the bipolar cell that the rod has captured ph oton. The
entire .sequen ce of even.ts takes only a matter of rnilliseconds. Con es act in a
qualitatively similar fashion. You can lea rn more about pho totransduction at
si tes.sinauer.com / neuroscience5e/ cmimations 11 .02.hhnL
The runotmt of glutam ate present in the photorea::ptor-bipolar cell synapse
at a ny one time is inverse ly propo rti ona l to the numbeT of photons being
absorbed by th e pho toreceptor. TI·ms, tmlik.e most o the r types of neu rons,
photoreceptors d o n ot respond in an all-or-nothing fashion. They pass their
information on to bipola r cells via graded potentials, which va ry in size,
instead of alJ-or-no ne action potentials or spikes, w hich are fmmd tluoughout
the nervou s system (see 1}.
The three cone photopig m ents are n ot dis tribu ted equa lly a m ong the
cones (Figure 2.12). Short wavelength-sensi tive cones (S-cones) constitute
only about S-1 0% of the total cone population, and they are essentially m iss-
ing from the center of the fovea . Tints, the foveal center is diduoma ti c (i t
h as only two color-sensitive cone types). \ Ve also know that the.re are more
long V\ravelen gth-sens itive cones (.L-conesJ than m editun waveleng th-sens i-
FIGURE 2.1 2 Blue, gr8Eln, ard red representth9 S-. M -. and

L-cones. o1 a li'Ving human bQlng ln a patch ofr9tlna at
1 dsgree from the fovea The pseudooolor Wnage was made by the
use of adapttve cptics to ITIQasure and b)'Pass the atMwratlons of 1h9
eye. and o f to isdate the different photoplgments.
(From Roorda. and Williams, 1QQQ: courtesy of Austin Roorda.)

42 CHAPTER 2
TA BLE 2.2
Properties of human photopic and scotopic vision
Property Photop;c system Scotop;c system

Photoreceptors 4-5 milli on cones 90 million rods
Location in retina Throughout retina, with Outside of fovea
highest concentration
close t o fovea
Acuity (detail) High Low
Sensitivity Low High
tive cone.s (M-cones)i it has been estimated that there are, on average, about
twice as many L-cones as M-cones, altho ug h the ratio of L- to M-cones va ries
enormously am ong indiv iduals. Table 2 .2 illustrates some of the fund..1n1ental
differences in the properties of the photopic (h.igh-illurnination) a nd scotopic
{low-illumination ) visu al systems.
\Vhen ph otoreceptors ca ptme Hght, they produce chemical ch anges tha t
st.·ut a cascade of n e ural events ending in a visual sensa tion. Pho toreceptors
send theh by way of the synaptic terminals, specia li zed s tn1ctures
for contacting other reti nal neurons. Figure 2.8 s hows examples of rod and
cone synaptic terminals. The syn aptic tenninals cont.a.i n connections from the
1
n eurons that photoreceptors " talk to' the horizontal an d bipolar cells.
:
FURTHER DISa.JSSION of cones and color detection can be found in

Oiapter 5 on page 124.
Lateral Inhibition through Horizontal and Amacrine Cells

As the name implies, horizontal cells run pe rpendicub.r to the photorecep-
tors_, making contacts behveen nearby photoreceptors. Th ese latera l connec-
ti ons play an unpQl'tan t funqioruil role in the form of lateral inhibition, w hid1
enables the s ignals that reach retina l gan g li on cells to be based on differen ces
in activa tion between nearby photoreceptors. La teral inhibition p lays an irn-
p ortant role in vis ual perception, and in several visual illusio ns (e.g., Mach
bands and the Hermann grid-see Web Essay 2.1: Seeing Illusory Stripes
and Spots). We w ill have more to s..,y abo ut late ral inhibition in the 11Cen ter-
Surround Receptive Fields" section be.low.
Amacrine cells are also part of the lateral pathway. Like horizo ntal cells,
mnacrine cells run perpendicul ar to the photorecep tors in the inner layers of
the retina, where they receive inputs from bipolar cells and other amacrine
cells and send signals to bipolar_, a macrine_, and retin a l ganglion cells. Amacrine
cells come in many flavors.. by smn e estimates as many as 40 (Rodieck, 1Q98).
horizontal cell A specialized retinal Altho ugh an'!acrine cells have been implica ted in bo th contras t enhancement
c ell that ocntacts both ph:itoreceptor and temporal sensitivi ty (the detection o f chan ges in light pa tterns over ti1ne),
and bipolar oells.
the ir precise fun ction remains unclear.
lateral Inhibition Antagonistic
neural Interaction between adJooent Convergence and Divergence of Information via Bipolar Cells
regbns of the retina
If h o rizontal a n d amacrin e cells fo rm a la te ral pathway in the retina, then
amacr1ne cell A retln1' oell fourid photoreceptors, bip olar cells, a nd gang.lion cells can be cons idered to fo m1 a
In the lnrier synaptic layer that makes
synaptlo cmtacts With blpoi'lr cells, vertical pathway (see Figure 2.8). Bipolar cells are the intermediaries. The re
ganglion cells1 and other amacrlne are va rious types of bipolar cells_, and their wiring d etenTtines the information
cells. that is pas.seti from the photoreceptors to the ga ngUon cells. For example.. in

peripheral vision a bipolar cell recei\·es input from as many as 50photorecep- bipolar cell A retinal cell that syn-
tors, pools this information, and passes it on to a ga nglion cell. Titls convel'- apses "1th either rods cr cones (not
gen ce of infom1ation from many photoreceptors to a sing le bipobr cell (knmvn both) am with horizontal cells. and
then passes the signals on to ganglion
as a diffuse bipolar cell ) is a characteristic of the rod and the sa:me cells.
sort of convergence also occurs in th e cone pathway in the peripheral retina.
diffuse bipolar cell A bipolar retinal
Pooling of information from many photoreceptors is a very important cell wt'Dse prooesoos are spread a.rt
mechanism for increasing visual sensitivity. Indeed, the fact that m ost rods to re::::etv.e Input frcm multlple cx:::nes.
communicate 'vi th ganglion cells through diffuse bipolar cells largely aci:ow1ts sensttMty 1. The ablllty to PffOOMl
for the ability of the rod system to function \·veil in dim lighting conditi ons. via the sense organs. 2. Extreme
A diffuse bipolar cell nl.:1y fire at the same rate in response to a single point rooponsMiness to radlatbn, especially
o f bright light or several s pots of dim light, so a ganglion cell listening to the to Ilg ht of a spedl c wavelength. 3.
diffuse bipolar cell will be unable to tell w hich pattern of light is present. The ability to respond to transmitted
elgnals.
The high degree of neural convergence in peripheral 1,.-ision has important
consequences for visual acuity, whid1 falls off rapidly with eccentricity (see visual acuity A measure of the fln-
Table 2-1), wh id1 is discussed further in Chapter 3. est detail that can be resorJed by the
eyes.
In contra st, in the fovea, midget bipolar eels receive input from sin gle rones
and pass this information on to sin gle ganglion cells. The fact tha t one-to-one midget bipolar cell A small bipolar
pathways behveen cones and ga nglions exist only in the fovea accounts for cell In the central retina that rec:elves
lnpLrt frcrn a elfXie cone.
w hy images are seen most dearly when they fall o n this part of the retina. The
ON bipolar cell A bipolar c ell that
high degree of con vergence in the retinal periphery ensures high sensitivity to
responds to an lnorease In light c ap-
light but poor acuity. The low degree of convergence in the fovea ens ures high tured by the cones.
acuity but poor sensitivity to light. You can ex-p lore this trade-off of sensitivity
OFF bipolar cell A bipolar cell that
and acuity in Web ActlVlty 2.5: Acuity versus senslUvtty.
responds to a decrease In light cap-
Each foveal cone actua lly contacts two bipolar cells (representing a diver- tured by the cones.
gence o f information): one respond s to an increase in light captured by the
cell A retinal cell that
cone and is ca lled an ON bipolar cell; the other respond s to a decrease and is receives visual lnfamatlon from ph:::ito·
called an OFF bipolar cell . TI1e fad that tl1ere ore both ON and OFF bipoku receptors via 11..o lntet"medlate neurcn
cells provides infomlation about whether the retinal illumination increased types (bipolar cells and amacrlne cells)
or d ecreased, and as we \Ylll see, the ON / OFF distinction built into the ana- and transmits lnformatlcn to the brain
tomical s tructure of the retina is present at many levels of the ";sual pathway. and mldbrain.
P ganglion cell A small ganglion
Communicating to the Brain via Gangfo Cells cell that recewes excitatory Input
elngle midget bipolar cells In the cen-
By the time signals arrive at the 6nal lnyer of the retina (the ganglion cells), tral retina am feeds the paivocellular
th ere has already been a lo t of information processi ng. Some infor mati on 1(:('fer of the lateral genlculate nu:::leus.
has been pooled through convergence; some has been enhanced by lateral
M ganglion cell A ganglion cell
pathways. Ga nglion cells receive their input from bipolar and amacrine cells, resembling a little umbrella that
process this input further, and send messages off to the brain through their reOOvoo excitatory Input from diffuse
axons, which gather in the back of the eyeball and emerge together as the bipolar cells and feeds the magno-
optic nerve (see Figl.Ue 22). cellular layer of the lateral genlculate
nu:::leus.
By now you are probably getting the idea that ead' cell type comes in many
varieties, and ganglion cells are n o exception. The human retina contains about
t250,000 ganglion cells, about one-htmdred th the number ofph otoreceptors.
Midget bipolar cells send their s ignals to small ganglion cells, which are widely
referred to as P ganglion cells beca use they feed the parvocellular ("small
cell") layer of the lateral geniculate nudeus (LGN) (discussed in Ch.,,pter3). P
ganglion cells cons titute abou t 70% of the ganglion cells in the htunan retina.
Diffuse bipolar cells project to ganglion cells that are known as M ganglion
cells (Figure 2.13) because they feed the magnocellular {''large cell") layer of
the LGN. TI1e dendrites of these ganglion cells spread out much more than
those of the P ganglion cells, givi ng them an tunbrella-like appearnn ce. About
&-10% of ga nglion cells in the htm1an retina are of the M variety. The dendrites
of both P and M ganglion cells incre.3se in size with retinal eccentricity, but at
all eccentricities the P ganglion cells h ave much smaller dendritic trees than
d o tl>e M gan glion cells.

44 CHAPTER 2
FIG URE 2. 13 Different types o f retinal P and M ganglion OOls . ShO"Ml are ganglion
calls in sQCtion. (After Oyst er, 1009.)
1l1e astute reader may have no ticed that M and P ganglion cells togethe r
cons titute about 8()Q/u of a ll ga ng lio n cells . Other gang lio n cell types, known as
konlocellularcells, project to koniocellular layers in the LG N. Some of these,
wi th input from S-cones, may be part of a ''primordial '' blue-yellow p athwc1y
(seeChn pter 5), w lUle yet other gan glion cells that p roject to the konlocellular
layers a re thou ght to corresp o nd to "n onblu e 11 ko n.iocellular cells.
CENTER-SURROUND RECEPTIVE FIELDS Much o f w ha t w e kn ow about

h O\v retinal gan g lion cells work comes from p ain staking physiological s tud-
ies in wh ich tiny electrod es are used to s tudy the electrical ch anges in in-
dividual ga ng lion cells. Gangli on cells fire acti on p o ten tia ls sp ontaneo usly,
at il bout on e spike p er second, e ven in the absen ce o f vis uaJ s timul ation.
H owever, each gang lion cell has a sm a ll w indow on the world kno\-vTl as
its receptive field . The recep tive fie ld is the region on the retina in which
vis ual s timuli influence the neu ron' s fir ing rate. Th is influence can be eithe r
exci ta tory, increasi n g the gan g lion's firing ra te, o r inhibitory, d ecreasin g the
gang lio n's firing rate.
FURTHER DISCUSSION of receptiw fields can be found in Chapter 3 on

pages 70-74.
\Vork on horseshoe crabs and frogs provid ed some of our earliest infom1ation
on the receptive fields of retintil neurons (Ha rtline, 1940). But it w as Stephen
Kuffl er vd10 first m apped out the receptive fields of individual retinal gan glion
cells in the cat, using s mall spo ts of light (Kuffler, 1953). Flgure 2.14 illustrates
Kuffier 1s main find in gs, which also apply to primate retina, and p rovides so me
impo rtant insig h ts in to how the retina p rocesses ";stial info m1ati on. Kuffler 1s
expe rimen ts a re sllnulated in Web Ac tivity 2.6: Ganglion Receptive Fields.
Let's conside r Fig ure 2.14b. Kuffler 's visual s timulus was a srnall spot of
light, w hich he m o ved about on the retina, tuming it on and off while recording
impulses from a s ingle retinal gang lion cell \ Vhen the spot was placed on a
specific s ma ll region of the retina, the ganglion cell increfl'Sed its firin g rate when
the li ght W4lS turned on (this response is indicated by a plus s ign in the fi glU'e).
cellular layers of the lateral genlculate This area o f the retina is called the /(center" of the gar1g lion celrs receptive
nucleus. This layer Is knoW11 as the field. \.Yhen the spot '""·as m oved to an a djacent area of the retina, the ganglion
konlocellular layer. cell decreased its firing ra te w hen the light was turned on (indica ted by a minus
receptive field The region on the sign). It is in teres ting tha t tuming the light off in th.is area s urrounding the
retina In whic h \/lsual stlmUll n fluence a recepti ve-fie ld renter led to a brief s mge in the cell' s firin g rate, after whid1
neuron's firing rate. the cell settled d own to its spontan eous rate.

(a) Reoord signal

from optic nen•e FIGURE 2.14 Retinal ganglion 09!1 r909ptiv€1 fi8lds. (a}
Mapping reoeptivg fields. fP) ON -center field. In
each image on the left, the small white circle illustrates
thEi region o n thQ r9tina whera thQ rc;itinal ganglion CQll
increased its firin g rate whoo the spot (small yellow
circle) was turn9d on. The largg gray c irc le illus trates
the region on the retina where. the reitlnal ganglion ct:iH
decreased its firing rate when the spot was turned on
and incrQaSlQd its rate whQO th9 spot was turmtd o ff.
The pbts on the right illustrate the spikes fired by the
retlpal ganglion C911. tc) OFF-center field. In
each lmag4':) on the left, the large Wille c irc le Illustrates
thQ region on th9 retina v.tiere th9 retinal ganglion cell
ilcreasQCi Its firing rate wt1'Wl the spot V\laS turned en
(b) ON-center ganglion cell The small g<Bf c ircle illus trates the regbn on the retina
Light on .....tiere the retinal ganglion eel decreased its firing rate
y,,tien the spot was turned on and hcr93Sed its rate
Vvf'len thQ spot was turned o ff. The pots on the right
Illustrate the spik9s fir9d by the associated gan·
Spot in
center R.•poru• 11 11111111111111 I I I gllon cell. (d) The efft1et of varying the spot slzQ. Note
that firing is fastest wh0n the spot just fills the RF cen -
ter. Increasing tt-R spot size further results in reduc9d
firing due to latgr-al inhibition from the surround (Part
d aftg Kuffi9r, 1053.)
Spot in
surround l-+111-11-lll!+j!-111111+++-l
(c) ganglion a! ll
Llghtru'
R.oporu• II I 111111111111
(d)

46 CHAPTER 2
ON-center cell A cell that depolar- 11'1.e cell jus t d escribed is known ns an ON-center cell. It increases its firing
izes In response to an Increase In light rate wh en a light is h.tmed on in the center of its recepti ve field , and it decreases
Intensity in Its receptive· field oonter. its firing rate when the light is turned on in the s urro und. However, nearly as
OFF-center cell A cell that d epolar- rnan y ganglion cells d o exactly the opposite: their firing ra tes decrease w hen
tzes in response to a decrease In light a light is turned on in a spot in the cen ter of the recep tive fie ld, a nd increase
Intensity In Its receptive-field center. wh en a light is turned on in a .spot in the s ur round . These are knmvn as
filter An acoustic, electrical, elec- center cells (Fig ure 2.14c). Mo.st retinal gan glion cells h ave one of these two
tronlc, cr optic d0\lice, Instrument. types of concentric center-surrm md organization .
computer program . or neuron that
allows the passage or some freq uen-
.4.n im.porta.nt finding of Kuffler's was that the spatial la)r"Ollt of the gan g lion
c ies cr dl;Jltal elements and blocks the cell's receptive field is essentially concentric; that is, a small circular area in the
passage o f others. center responds to an in crease in ilhunination, and a surrotmding ring resp onds
contrast The difference In lurnl· to a d ecrease in illumina tion , and size rnatters! The ganglion cell fires fastest
nance between an object and the when the size o f the spot of light 1natches the s ize of the excita tory center,
background, or between lighter and and it reduces its firing rate when the spot of Ught begins to encroad1 on its
darker parts of the same object. inhibitory surround (Figure 2.14d). This interaction behveen the antagonistic
center and surrolUld is k.novm as lateral inhibition.
The center-sur ro und organization has two important fon ctiona l conse-
quences. First, as n oted above, each ganglion cell vl'i ll respond best to spots
of a par ti cu lar size (and will respond less to spots that are eithe r bigger or
smaller). Jn this way. . retin.aJ ganglio n cells act filter by respondJng best to
stimuli tha t are just the right size, a nd le-ss to stimuli th'1t are larger or s maller.
Second , ganglion cells are most sensitive to i.n the intensity of the
light in the center and in thes urrotmd, and they are less affected by the aver-
age In tensity of the lig ht 111is isa useful quality because the average intensity
of the li ght faLLing on the retina varies a lot1 depending on w he ther you are
indoors or o utdoors, wh ether it is daytime or nighttime, how far away you and
the objects you' re looking nt .ire from the source of ilhunination, and so on.
But the contrast-the difference in lum.inan ceor brighh1ess behNeen adjacent
bits of the scene-will be roug hly the s..<tm e regardless of lighting conditions.
The cente r-s urround an tagon ism, or lateral inhibit.ion, also h as oth er
important perceptual consequences-resulting in the illusions of stripes and
spots (Figure 2.15 ). (See Web Essay 2.1: seeing Illusory Stripes and Spots. )
Phenomen ologicall)r; \Ve have the impression tha t our eyes work like video
cam eras, capturing faithful snapsh ots of the world around us . Note, thou gh,
tha t the rest of the visual system sees only w hat the retinal gan glion cells show
8230336 AmMa Appa
FIGURE 2. 15 •eartoon" showing a dasstc n€·Uronal explanation for Mach bands.

See the text for dQtails.

it, a nd the gan glion cells are not content si mply to pass along the raw images
encoded by the ph otoreceptors. Instead, the g anglion cells, together with the
bipolar, arnacri:ne, and horizontal ce lls, net as an iinage filter, transfom1iilg
the rmv image into a new representation. This new representation hig hJights
certain imporklnt information, su ch as contras t, and largely d isco unts other
types of less useful infor mati on, su ch as ambient light intensity. In fact, the
whole visual system can be consid ered a long series of filters, with each s tage
in the system. res pons ible for extractin g a particular aspect of the visual wodd
and passing this aspect on to the next s tage.
P AND M GANGLION CELLS REVISITED As al ready me ntioned, retinal gan-

glion cells come in several types; for htunans, the most important of these are
the P M cells. The recep tive fields of these two types of gan glion cells
d iffer in some important ways. First, a t all eccentricities P cells have s maller
receptive fields than M cel ls have. This isn ' t too surpris ing, because the size
of the receptive field is probably determined by the size of its d e ndritic field
(see Figure 2.13): sin ce M cells lis ten to more ph otoreceptors (via bipoku,
horizontal, and amacrine ce lls) thnn P cell s d o, M cells respond to a larger
portion of the visual field.
An conseq uence of the differin g sizes o f M a nd Preceptive
fie ld s is that M cells are much more sensitive-better a ble to detect visu a l
stimuU-thanare P cells t.mder Jm,,·-light conditions (e.g., a t night). However,
the s ma ller recep tive fields o f P cells enable them to providt!iiner reso.l uti on
(greate r acuity) than M if th ere is en ough light for tli'e per- App a
ate. See Web Activity 25: Acuity versus Sensitivity for m ore on th.is trade-off.
P and M ganglion cells also differ in their temporal responses. Pcells tend
to resp ond '"';th s ustained firin g w hile li ght shines on their excita tory regions.
M 1."'e lls tend to respond mo.re transiently: an M cell will respond with a brief
burs t of impulse.s when the spot is turned onrand then it wiJl quickly return to
its sponta neous rate, even if the s p ot 1-em ains lit. llnLS, Mand P ga ng lion cells
s ignal different information to the brain. P cells provide information mainly
about the contrast in the retinal image, and M cells signa l information about
how the image changes over time.
Finally, P and M cells differ in what they say to the brain about the color
of tl1e li ght they detect (see Chapter 5).
Dark and Light Adaptation

When you em erge fr om a dark room into bright light (e.g., coming out o f
a movie theater), your pupil cons tricts to reduce the amount of lig ht arriv-
ing at your retina. Tllis automatic reflex is n othing to s neeze at, but there's a
good ch ance that you will! Sneezing in response to being exposed to a bright
light-the 1' photic sneeze reflex"- is n ot yet unders tood, even though it has
intrigued some o f history's g reatest minds. Aristotle (384-322 BCE) thought
the hea t of the s un on the nose might be responsible. However, Francis Ba-
con (1561-1626) sh owed that Aris totle was w ron g. Bacon s tepp ed into the
sun with his eyes closed and did no t sneeze; the heat was s till there, but the
sn eeze was not. Bacon g uessed that the s un 's light makes the eyes water1 and
that moisture (''braine hum our/' ill his words) then seeps into and irritates the
nose. We n ow kn ow th at the s neeze takes place too soon after light exposure
to be the resul t of the comparatively slow formation of tears. C 1rrrent thinking
suggests that the photk sneeze reflex is a re.suit of crossed '"; res in the brain.

48 CHAPTER 2 8230336 Amna Acea
FIGURE 2.16 Dark adaptation curw. The purpl9 curve shows th9 threshold llght
intensity required to detect a peripheral spot following sev9ral minutes of adaptatb n
tp a bright light. The r9d c urve illustrates the rapid adaptation o f the COMS. Th9 bU9
curve shows the slower reoovery of the rods to much low9r threshold inti9f"'ISiti9S (that
Cones only is, greater sensitMty). The purpe represn s the most sensitive of the two at
any gtven time.
\Vhen you ente r a d ark room from bright s unlight, the numbe r of photons
of light entering your eye might be reduced by a factor of several billion.
Initially you 'vill have trouble seeing anything, but afte r abo ut 30 minutes in
the dark you will be able to detect even just a few photons. The purple curve
10 15 20 25 30 in Figure 2.1 6 illustra tes the cha nge in th e threshold light intensity needed to
Tm"te in the d<uk (min) d etect a peripheral s p o t. Initially the threshold is very high, indicating low
sensitivity. But over 20 minutes or so the thresh old is greatly reduced (m ea n-
in g sensi tivity is incre..1sed ). And w hen you em erge from the dark and retum
to the sunlight, you w ill be able to see almost instantly. How does the vis ua. 1
.system alter its sensitivity over such a large opera ting r.mge?
1llere are four main ways in vdUch the visual syste m adjusts to ch anges in
illumi.nation: pupil size, photopigment regeneration, the duplex retina, and
neural ci rcuitry.
Pupil Size
\Vh e n a fl ashligh t is shone in someone's eye in a d imly Ht room, the pupil
l1uickly cons tri cts. The diameter o f the pupil ca n vary by about a factor of 4,
from about 2 nun in bright ill mni nation to about 8 mm in the d ark lF1gure 2.17 ).
Because the arnount of light entering the eye i.s proportional to the area of the
pupil, the 4-fold increase in diameter accoun ts fo r a 16-fold Unprovement in
sen.siti vi ty. In o ther words, 16 times <1S many quanta can enter the eye w hen the
pupil is completely dila ted, compa red \Vith whe.n it is cons tricted . Although
this adaptive ability certainly he lps, pupil dila tion accounts for only a smal l
paii of the visua l syste m 'soverall ability to ad ap t to l.ig ht and dark conditions.
Photopigment Regeneration
A second mech an ism fo r adUeving a large sensitivity range is provided by
the way photopigments are used up and replaced in recepto r cells. [n diin
(a) Bright illumination
2-mmpupil 8-mmpupil
FIGURE 2. 17 The black spots In thtli middle o f thoo.e two Irises show thtli possible
range o f pupil slz9s as W9 go from bright illumination (a) into th9 dark fP).

lighting conditions, plenty of photopigm ent is available, and r ods and cones
absorb and resp ond to as man.y photons as they can . As a lrea dy n oted, ro ds
provid e better sens itivity in s uch situa tions than do cones. Ind eed, the rod
system is capable of de tecting a sing le qu:anttun of light! (See Web Essay 2.2:
How Many Quanta Does It Take?) After a photopigment molecule is bleached
(used to d etect a ph oton), the molecule must be regenerated before it can be
used again to absorb another photon.
As the overall light level increases, the nwnber of photons s tarts to over-
whelm the system ; photopigment mo lecules ca nnot be regenerated fast eno ugh
to d etect a ll the photons hitting the photoreceptors. This s low regeneration
is a good thing for increasin g our sensitiv ity range. H photons m e scarce, we
use them a ll to see; if we have an overabundance, we s imply throw some of
them away and use the leftovers.
The Duplex Retina

This light com pensa tion mech anism is enha nced by humans' duplex retina s.
Rods p rovid e exquisi te sensithity at Im¥ light levels, but they becom e over-
whelmed when the bac kground lig ht becomes m ode rately bright, leading to
a loss in informa ti on quality. Cones ai·e rn uch less sensiti ve than rods (they
hmcti on poorly under very dim light), but thei r ope rating range is much
larger, stretching from about ten photons per second (jus t enough lig ht to see
color ) to hundred s of thousands o f photons per second (e.g., a snowca pped
m OLm tain in bright s unlight). So we use rods to see w hen the light is low, and
the cones take over when there is too much light for the rods to hmcti on well.
After ad aptin g to a brig ht light, cones recover se ns itivity quickly (red curve in
Figure 2..16) and then sa hmite. They are n ot very sensitive to very d im light.
Rod s recover m ore slm·vly (blu e curve in Figure 2.16), but after 20 minutes o r
so they a re very sensitive to dim light.
Neural Circuitry
Altho ugh pupil size, ph o topigm ent regene ra tion rates, and the rod / cone
d ichotomy all play a role in d a rk and light ad ap tation, the m o.s t important
reason we me not bo thered by varia tions in overall light levels has to do wi th
the neura l d rcuihy of th e retina. As we lemned earlier, ganglion ce lls fire at
their m a.xi mum rate \·v hen the centers of their receptive fields are brightly lit
whil e the surrounds are completely d ark (or v ice versa ) (see Figu re 2.14). But
the cells w ill s till fi re at an abo\o·e-spontaneous rate w hen lig ht fa lls on th e
entire receptive field, as long as the light is brighter in the ON p ortion than in
the OFF portion of the recepti ve field. Therefore, as long as the photoreceptors
feeding the ga ng lion cells are n ot completely s.a ttuated , the ga nglion cells \.Vill
encode the pa ttem of re lati vely light and rela tively d a rk areas in th e retinal
image. And the pa ttern of illurnit1L1tion, not the overall lig ht level, is the pri-
mary concem of the rest of the visua l system .
To s um up, the a nswer to the questi on of how the visual system deals wi th
s uch large va riati ons it1 overall Light levels has two parts. Firs t, we reduce the
scale of the problem by regula ting the a mo unt o f light en tering the eyeball,
by using differen t types of pho torecep tors in diffe rent s ituati ons, and b y ef-
fectively th rowing away photons we d on ' t need . Second, by responding to
the contras t between adjacent retinal regions, the gan glion cell s d o their best
to ignore w hatever variation in overn ll light level is left over.
336 Amma Appa

50 CHAPTER 2
Sensation & Perception in Everyday life
When Good Retina Goes Bad

Millions of pe:::>ple around the wortd suffEr" from blinding diseases in
v.tiich the rods and/or cones degenerate. These Include age-related
macular degeneration (AMO) and retlnltls plgmentosa (RP). (See
Web Essay 2.3: Clinical case: The Man Who Couldn't Read and
Web Activity 2.7: Simulated Scotoma.) At present, there are no effec-
tive cures top-event the progressive degeneration of the photoreceptors
that ocour in these diseases. For patients with AMD this may lead to an
lnablllty to read or recognize faces. Fcr patients with long-standing RR
this leads nevltably to lrreversltle blindness.
Fortunately, there are several exclt<ig new technological develop -
ments that provide hope for these patients. These are all based on the
notion that v.tilie the photoreceptors are dead cr dyng, p;>st-receptor
reurons and their connections are largely Intact. One such approach is
to substitute an electronic prosthesis (an artlficlBI devloo to repace or
augment a missing or Impaired part of the bodYl Into the retina. Typi -
cally the prosthesis uses a camera to convert light Into energy; an array
of electrodes Implanted in the retina generates an e-.ctr1cal stimula-
tion pattern based on the light pattern on the camera and delivers this
stimUlatlcn pattern to the Intact p;>st-receptor neurons (Figure 2 .18).
age-related macular degeneration Unfortunately, v.tille these retinal prostheses can restcre some sight,
(AMO) A disease associated with there are technical challenges to Implanting them, and they suffer low
aging that affects the macula AMD spatial resolution (Weiland, Che. and Huma'y1.Jn, 2011), allowing only
gradually destroys sharp central vision, perooptlon of spots of light end very high-contrast edges.
making It dlffk:ult to read , drlw, and Another approach that has had some earty success In animal mod-
reocgnlze facoo. There are two forms els Is to use gere therapy to express Nght-actlvated channels <i survlvng
of AMD: wet end dry. photoreooptors us<ig adeno-assoclated viral (AAV) vectors. This ap-
retinitis plgmentosa (RP) f!. pro- proach has been successfully used In several clnlcal tl1als In patl01ts.
gr""5lve degeneration of the retina Ii A third strategy Is to chemically modify endogenous ohannels in
that affects night \llslcn and peripheral retinal ganglion cells to make them light-sensitive. This approach es-
\llslcn. RP commonly runs In familee sentially adds a synthetic small molecule "photoswitch" to confer light
and can be caused by defects In a sensltMty onto retinal ganglion ceRs. end It has been shoWn to reinstate
number of different genes that have
reoently been Identified. light sensitivity In bHnd mloe (Tochltsky et al., 2014).
Each of these approaches provides promise for new treatments
for patients with t:lln<llng retinal dlscrders.
Video Laser
o•RF
Subretinal
implant
FIGURE 2.18 Retinal Prostheises. The ltlsert sho\NS sensors Implant-
ed In the retina. (Mer Welland et al .. 2011.) Photore<Epton;

THE FIRST STEPS IN \/lSION 51
Summary
1. This chapter p rovided some insight into the complex joumer_. thati 3
required for us to see stars and other spots of light. The pa th ofthe light Refer to 1he
was traced from a dis tant star through the eyeball and to its absorption by Sensation and Perception
photoreceptors and its transduction into neural signals. In subsequen t chap- Companion Website
ters we'll leam how those signals are transmitted to the brain and translated sttes.slnauer.COfll/Wotte4e
into the experience of percep tion .
fer actM1ies. essays. stt.dy
2. Light, on its w ay to becoming a sensation (a visu al sensation1 tha t is), can be
absorbed, scattered, reflected, trans m.itted, or refracted . It can become a sen- questbns, and other study aids.
sation o nly \vhen it's absorbed. by a pho torecep to r in the re tina.
3. Vision begins in the retina, when light is absorbed by rods or cones. The ret-
ina is like a minicomputer th.."1.t transduces light energy into ne wal energy.
4. The retina sends information to the brain via g.m.g lion cells, neurons ' "'' hose
axons make up the o ptic nerves. Retinal ganglion cells ha ve center-sur-
round receptive fields and are concem ed with changes in contrast (the dif-
ference in intensity between adjacent b its of the scene).
5. The visual system deals w ith large va ria tions in overall light intensity by (a)
regulating the amount of light ente ring the eyeba ll1 (b) u sing different types
of photoreceptors in d ifferent s itua tions, and (c) e ffectively thrm•d ng away
photons .....e don't need.
6. Age-related macular degeneration (AMD) is a d isease associated v.i th aging
tha t affects the rnac ula TI1e leading muse of visua l loss a mong the elderly
in the United States, AMD gradually d estroys sharp cen tral vision, making
it difficult to read, drive, and recognize faces.
7. Retinitis p igmentosa (RP) is a family of here:litary diseases characterized by
the p rogressive d eath of photoreceptors and degeneration of the pigment
epithe lium. In the most common fo rm of the disease, pa tients first notice
vision problems in their peripheral vision and under low-light conditions--
situat ions in w h.id1 rods play the dominant role in collec ting light.
8. A n um ber of exciting new d evelopments are aimed at restoring sigh t in
indi vidua ls with blinding re tin al disease.

CHAPTER 3
I Ed Churohill, Swrd w.ivss in 30 . 2011

Spatial Vision:
From Spots to Stripes
IN CHAP"TER 2 LEARNED that the macroscopic structures of the human
eye fun ction essentially as a biological camera: the iris regulates the a1nount
of light entering the eyeball; and the cornea, le ns, and aqueous and vitreous
humors focus these rays so that a dear image is formed on the retina. The rod
and cone photoreceptors capture this image in a way that is roughly analogous
to the way the fihn in a ca1nera captu res photographic images.
lt is here, however, tha t the analogy beh\.·een visual syste m and ca mera
ends. Ca meras take pkht res. Visual syste ms see. How do we ge t fro m an
image of the world in front of u s to an interpreh'ttion of that wor ld- what is
out there, where it is, and what we can do to it? This p rocess shuts in the eye-
ball itself, w here the pos trecept or layers of the retina translate the raw light
array captured by the photoreceptors into the patterns o f spots s urrounded
by darkness, or vice versa, detected by the ganglion cells (see Figure 2.14 in
Chapter 2). As we discussed in C hapte r 2, this retinal tra nsbtion help.s us
perceive the pattern of light and dark areas in the visu.:i.l field, regardless o f
the ove·rall light level (e.g., it en ab les us to see almost as well at dus k as we
can at n oon ).
In this chapter we follow the path of image processing from the eye ball
to the brain (Fig ure 3.1). Ga nglion cells in the retina respond preferentially
to s p ots of light. As we ,..,,,ill see, ne u ron s in the cerebral cortex prefer lines,
edges, and stripes. Fu1'them1ore, thi s portion of visual cortex is organized into
thousands of tiny computers, each responsible for determining the orientation,
w idth, color, and othe r characteris tics of the stri pes in one s mall portion o f
the visual fie ld . ln C hapter 4 we \vill continue this story by exa mining how
other part.s of the brain assemble the outputs from these minicomputers to
produce a coherent rep resenta tion of the objects whose reflected light sta rted
the photoreceptors firin g in the first p lace.
Visual Acuity: Oh Say, Can You See?

Tire King said, ''] sent the two Me::.-sengers1 either. They're lx>th gone
to the fawn . Just look along the road1and tell me 1:f you can see either of
tliem."
"'I see nobody on the road," said Alice.
•'J only ·wish Iliad such eyes," the King remarked h1 a fretful tone. "To be able
to see Nobody! And at that distance, too!"
- Lewis Carroll, Through the Looking Glass

54 CHAPTER 3
(a) L•ft R;ght
(c) Retinal Tha.Jamus Primary Visual Higher-order

ganglion cells LGN vi.suaJ cortex oll!!lsociation visual alil8ociation
cortex cortex
JU
Motion/spatial Dorsol.ateral
parietoteniporal
ilnalysis
"'"""
ILay.. 2,3 in.. <b-1 l
Fonn layer.s
__j_ interstripe
.lnfe"'"
ocap1tot,emporal
"'"'
Color
FIGURE 3.. 1 Cortical visual pathways. (a) Th9 basic organization ofth9 primary
visual pathway from eyeball to striate ccftex, In transverse section. (b} A lateral view
of th€1 brain, Hlustratlng 8raf'ly visual areas and showing th& putattva whsre (dorsaO
and what streams. (c) The flow of information for motbn, form, and oolor
analysis from retinal ganglion cells to hlgher"()rder visual association cortex. (Part a
after Purvgs et al., 2012: b aft9r Briegdlove and Watson 2013; and c after BlumenfGold,
2002.)
Since we1.l be talking in this chapter about how the visual system codes im-
ages in terms of oriented stripes, let's start by determining just how well we
contrast The difference In lumi-
nance between an object and the see stripes whe_n they are very close together and / or when the difference in
background, or between Ughter and illumination between the stripes and the background (the contrast) is very low.
darker parts of the same object. In addition to setting the boundary conditions for how well we should expect

SPATIAL "1SION 55
FIG URE 3.2 A visual acuity t9st. S99

the text for details.
the visual sys tem to be able to perform, we w ill use this sectio n to introduce acuity ltte smallest spatial detail
some importa nt jargon that we' ll n eed in the rest of the d iapter. that can be resolved at 100% contrast.
Get a tape measure, prop your-textbook up, and, while lookin g a t the X in cycle Fcr a grating , a pair consi>tlng
the midd le of Figure 3 .2, back up tmtil you cannot tell the orientatio n of the of one dark bar ard one bright bar.
black and white s tripes. Measure how far your eye is from the page. Now walk visual angle ltte angle subtended
forward a bit until you 're s u re you can see which grating includes vertical by an object at the retina
stripes i.lnd which ho rizonta l s tripes, and again m easu re your d ista nce fro m
the page. Cong ratula tions! You just complete..1 a fast {but not terribly accurnte)
m easurement of your own visual resolutio n acuity.
Eye d octors specify acu ity in terms like 20/20 (m ore about this in a mo-
me nt), but vision scientists prefer to talk about the sma lles t visual angle of a
cycle of the grating that we can perceive (Figure 3 .3). A cycle is simply one
repetition of iJ black and a white s tripe (both of the gratings in Figure 3.2 h(1ve
25 total cycles). Visual angle, which we mentioned briefl y in Chapter 2,is the
iJngle tha t would be formed by lines going from the to p iJnd bottom (or le ft
and right, depending on the orientation of the stripes) o f a cycle on the page,
passing through the cen ter of the lens, and ending on the retina. You can learn
more about this co ncep t in Web Ac tivity 3.1: Visual Angle.
More precisely, to calculate the visu al angle of your resolution acuity, di\.i de
the size of the cycle in Figme3.2 (which is 2 millimeters, or 1/ 16 inch) by the
viewi ng dis tance a t w hich you could jus t barely make out the orientatio n
o f the gratings (average your firs t and second m easurem ents to get a rough
estimate of this distance), and then take the arctangent of this ratio. Under
ideal conditions, htm1ans with very good vision can resolve gratings like those
in Figure 3.2 w hen on e cycle s ubtends a n angle of approximately 1 minute o f
arc (1 arc minute or 0.017 d egree).
111..is resolution acui ty represents on e of the fund am ental limits of spatial
vision: it is the fines t high-contrast detail that can be resolved.. The limit is
d etermin ed primarily by the s pacin g of photoreceptors in the retina. To see
.': •-······· · • • •.·.::::;;;z·· · ···· · ·· Y F'IG URE 3.3 Visual angl9. ShOIMI
h.:.:fe is the angle sizei of o ne cycle of a
grating at the retina..

56 CHAPTER 3
FIGURE 3.4 A squarewwe grating (a) (b)

(a) and a s ine w'fNe grating (b). (c} The
1111 1111
stripgs of thg sine w.:we grating arg
wider than the pho toreceptors !):>ink
circles in th9 top panel), and the grating
c an be recons tructed vertically. (d) The
s tripes of the sine wave grating are nar-
ro wsr than the photoreceptors, so both
black and whittio bars w ill fall inside a
si nglei r909ptor (top panel"), r&sulting in
a uniform gray tield (bottom panel).
(\(\(\
jv v v v
(d)
sine wave grating A grating with a why, imagine that we1re projecting the sine wave gratings s hown in Figure
sinusddal lumlnance proflle as shown 3.4a o nto the retina. The light intensity in s u ch gra tings varies sm oothly an d
In Figure 3.48. con tinuously across each cycle (unli ke the gratings in Figures3. 2, 3.3, and 3.4b,
in w hich inte ns ity cha nges abmptly from black to white and back to black).
H owever, the visual system "samplesu the grating through the ar-
ray of receptors at the back of the re tina (in this respect the eye is rnore like a
digital camera tha n like a traditional ca mera that uses fihn). lf the receptors are
spared s uch tha t the whitest a nd bla.ckest parts of the gra ting fall on separate
8230336 A cone (Figure 3.4c ), we should be able to make out the grating. But if the entire
cycle falls on a single cone (Flgure 3.4d), \Ve will see n othing but a gray fie ld.
Cones in the fovea have a cen ter-to--center separntion of about 0.5 minute of
arc (0.008 desree), w hich fits nicely with the observed acuity limit of l minute
o f arc (remember th at we need two cones per cycle to be able to perceive the
grating ttccurtt te ly). Rods a nd cones in the periphery are packed togethe r less
tightly (rec,1U that in the periphery, rods are physic,tll y more tightly packed

SPATIAL 1'1SION 57
(denser} than cones, as shown in Figu re 2.10 in Chapte r 2), and he re m any
receptors con verge on eac h cell. As a result, visual acuity is much
poorer in the periphery than in the fove a . For a demonstration of the d ifference
I Stroh
between foveal and periphe ral vision, see Web Activity 3.2: Foveal Ac uity.
(See also Web Essay 3.1: Hyperaculty.)
A Visit to the Eye Doctor

Eye d octors don ' t in te rms of visual angles a nd cycles. The last
titne you visited your eye d octor, sh may h ave as d you to read le tte rs, de-
creasing the size of the le tters tmtil you made several errors. Then she m ay have
told you th at your visual acuHy was 20/ 20 ii your vision was good, or 20/30 if FIGURE 3.5 A Snellen E. The letter
you need ed gla sses, or p ossibly 20/ 10 if you could read the s ma llest letters on size is fiv.9 tim9s the stroke size.
the eye d'\art. Tllis method for designating visual acuity WclS invented in 1862
by a Dutd1 eye doctor, Hem1'1n Snellen (1834--1908). Snellen constmcted a set
of hlock le tters for which the le tter as cl w hole was five times as large as the
strokes that formed the letter (Figure 3 .5). Note tha t the resul ting pa ttern s are
re.minisa:!nt of the gratings in Figure 3.2 He then defined visual acuity as follows:
(d ista nce a t which a p erson can jus t id entify the lette rs)
(dis tance at w hich a person with "n ormal'' vision can jus t id entify the letters)
In later adap ta tions o f the Sn elle n te.s t, the vie wer w as positioned a t a
constant d istance of 20 feet, and the s ize of the le tters, ra ther th an the position
o f the vie'\ver, w as altered . So normal vision ca me to be defined as 20/ 20. To
rela te this m easure bac k to visual angle, a 20 / 20 le tter is designed to s ubtend
an angle of 5 a rc minutes (0.083 degree) a t the eye, and each stroke of a 20 / 20
le tter subtends an angle of 1 a rc minu te (the fa rnili<'lr 0.01 7 degree). Thus, if
you can read a 20/ 20 le tter, you ca n d iscern de tai l that s ubtends 1 minute o f
ilrc. If you have to be at 20 feet to read a le tter that som eone with n om'\al vi-
si on ca n read a t40 feet, yo u have 20 / 40 vision {worse than normal) . Althou gh
20 / 20 is often con sid ered the gold s ta nd ilJ'ci, most healthy young adul ts h.:w e
an acu ity level closer to 20/ 15.
Types of Visual Acuity

So far '"'-e' ve d iscu ssed h vo fom'\s o f v isual acuity: the fin est s tripes that can
be reso lved (some times referred to as the minimum reso lvab le acuity) and
the smaUest lette r that can be recogni zed (the mininnun recognizable acuity).
H ow shou ld we d efine the keetu'\ess of sight? Visua.i acuity is used to specify a
sp a tial limit. O ver the centuries there have e mel'ged a large number of differ-
ent ideas about how to d efine, measure, and s pecify visual acu ity (Table 3 .1 }.
MINIMUM VISIBLE ACUITY Minimum v isible acuity refe rs to the s mal lest
object that on e can de tect. As early as the seve nteenth ce ntury, Benito Da,.;;a
d e Va ld ez (1 591-1 634 ) meas ured the dis tance at w hich a row of mus tard
seeds could no longer be cotmted , and enrly as trono me rs like Robe rt Hoo ke
(1635-1 703) were inte res ted in the size of s ta rs that could be d etecte d and their
relation to re tinal anatomy. In th is context, the minimum visible acuity refers
to the sm alles t targe t that can be de tected . Under ideal conditions, htunans
can de tect a lon g, dark wi re (like a cable of the Golden Ga te Bridge) agains t
a very brigh t background (like the s ky on a bri ght s unny day) w he n they
subte nd an angle of just 0.5 arc second (about0.00014 d egree). It is \'li dely ac-
cepted that the minim tun vis ible acuity is so s rnaU for hvo reason s: firs t, the
optics o f the eye (d escr ibed later in the chapter) the image of the thin
line, ma king it much wider on the retina; second, and the fu zzy re tinal image

58 CHAPTER 3
TABLE3.1
Summary of the different forms of acuity and their I imits
Type of acuity Measured Acuity (degree)

Minimum visibte Detection of a feature 0.00014
Minimum resolvable Resolution of two features O.Q17
Minimum recognizable Identifi cation of a feature 0.017
Minimum discriminable Discrimination of a change in 0 .00024
a feature
of the line cas ts a sha d ow th at reduces the light on a row of cones to a level
that is just de tectably less than the light on the row of con es on either si de. In
oth er words, although we specify the minimum visible acuity in terms of the
Mtgular size of the target a t the retin a, it is actua lly limited by o ur abi li ty to
discriminate the intensity of the target relative to its backgro und . Although
the minimum visible acuity represents one limit to spatia l vision, it is in fact
a lim it in the ability to discern s mall changes in contrast, rather than a s pa tia l
limit per se, and minimum visibfe acuity is not used clinica lly.
MINIMUM RESOLVABLE ACUITY Minim tun resolvable acuity refers to the

snrnliest angular separation beti.-.ieen neighboring objects that one cm resolve.
More tlum 5(00 years ago, the Egyptians assessed visual acuity by the ability
of a n observer to resolve d o uble s tars. There is currentl y still debate about
h ow bes t to de fine and measure resolution. H owever, today, the minimum
resolvable acuity ij. much more likely to be assessed by determining the fin-
est black a nd white s tripes that can be resolved . Under ideal conditions (e.g.,
high contrast and luminance), humans wi th very good \.; s.ion can resolve
black and white stripes when one cy·de subtends an angle of approxin-...1. tely 1
minute of arc (0.017 degree). This minimum resolvable acuity represen ts one
of th e fundamental limits of s patial vision: it is the finest high-contrast d etail
that can be resolved. ln foveal vision the limit is detemlined primarily by the
s pacing of photorecep tors in th e retina.
MINIMUM RECOGNIZABLE ACUITY Minimum recognizable acuity refers

to the a ng ular s ize of tl1e s ma llest fen ture that one can recognize or identify.
AltllOugh this method h ns been used s ince tl1e seventeenth century; tl1e ap-
8230336 Amma Ap pro;ich still used by eye d octors to day was introduced m ore than a century
ago by Hem1an Snellen a nd h is colleagues, and is discussed in the main text.
MINIMUM DISCRIMINABLE ACUITY Minimum discriminable acuity refers

to the angular size o f the s ma llest change in a fea ture (e.g., a cha nge in size,
position, or orientation) that one can discriminate. Perhaps the m ost s tudied
examp le of mlnimum d iscriminable acuity is our abili ty to discen1 a differ-
ence in the re lative positions of two features . O ur vis ual sys te ms are very
good at tell ing where things are relative to each o the r. Consider two ab uttin g
h orizontal lines, o ne s lightly higher than the other (see Web Essay 3 .1: Hy-
perac uity). TI1e smalles t m isa lignment that we can reliably discern is knmvn
as Vernie r acuity-narned afte r the Frendunan Pierre Vernier (1580-1637t
whose scale, developed in the seventeenth centu ry, was widely used to aid
s hips' navigators. The s uccess of the Vernier scale b..1.Sed on the fact that
humans are very adept at judging whether nearby Lines are lined up or n ot.
Tirns_, Vernier alignment is s till wide ly used in precis ion machin es, and even
in tl1e dia l switches in mode m ovens. Under ideal conditions,, Ventler acuity

SPATIAL 1'1SION 59
may be just 3 arc seconds (about 0.0008 degree)! TIUs performance is even m ore spaUa frequency The number of
remarkable w hen you con.sider that it is abou t ten tiir1e.s s maller than even grating cycles (I.e .. dark and bright
the srn.allest foveal cones. Note that the optics of the eye the image of bars) In a given unit of space.
a thin line ove r a number o f retina l con es, and that the eyes are in cons tant cycles per degree The number
m.o tion, and this performance appears even more remark..'lb le. of grating cydes pe< deg ree of visual
angle.
Vernie r acuity is not the m ost remarkable form of hyperacu.ity. Guiuness
V\'orld Records 2005 describes the "highest hyperacuity'' as follows: ''In 1jril
1984, Dr. Denrus M. Levi" (yes-that's one of the authors of tJ-J..,_booli , ,.pa
repeatedly identified the relati ve posi tion of a thin, bright green line within
0.8 seconds of arc (0.00024 degree). This is equivalent to a displaceme nt of
-s ome 0.25 inches (6 mm) a t a distance of 1 mile (1 .6 krn) ." This "remarkable"
position acuity was accomplished with a bisection task, but it can actually be
understood based on a straightforward model of human spa tial vision.
Acuity for Low-Contrast Stripes

Up to now we' ve been d iscussing the tiniest high-contrast d etails that we can
resol\re. We learn ed that h igh-contrast sin e wave gratings a m be distinguished
from a uni fo rm g ray field, as long as adjacent pairs of light or dark s tripes
are separated by at least 1 arc minute of visu..i.l angle. But what happens if the
contras t of the s tripes is reduced-that is, if the light s tripes are made darker
and the dark strip es lighter?
1ltis was the ques tion asked by Otto Scha de in 1956, when he \Vas work-
ing for the RCA Corporation. Schade s howed people s in e \.Vave \.vi th
different spatial frequencies and ha d thern adjus t the co ntrast of the gratings
until they could just be detected. Spatial frequency refers to the nttmber o f
times a pattern, such as a si ne wave grating, repeats in a given t.mit of space.
For example, if you v iew your book from abou t 120 centimeters (about 47
inches) away, the visual angle between each pair of white s tripes in Figure
3.6a shows a grating with a relatively low spa tial frequen cy (about 2 cycles
per degree). Figure 3.Gb is about 025 degree, so the spatial frequency of this
grating is 1/ 0.25 = 4 cycles per degree, and Figure 3.6c illus trates a rebtively
higher spatial frequency (about 8 cycles per degree). Web Activity 3.3: Gabor
Patches provides additional illus trations o f sine wave gratings at different
spatial frequencies.
you might think that the wider th e stripes (that is, the lower
th e spatial freque ncy), the easier it would be to distinguish the light s tripes
from the dark stripes. But this is not exactly w hat Schade fotmd . He, and FIGURE 3.6 Sine wave gratings illus ·
la ter Ferg us Campbell and Dan Green (1965), demonstrated that the human trating low '3), medium (b), and high (c)
spatial frequencies,
(a) (b) (c)

60 CHAPTER 3
FIGURE 3. 7 Thir;:i oontrast s.&nsiti\lity function {rOO line): the

window of v isibility. Any objects whosir;:i spatial frequenc ieis
and contrasts fall within the yGllOVI/ regio n will b9 \lisible. Thosir;:i
cutside the yellow rir;:igion are outside the window of visibility.
ThQ red linQ delimits th€! seeing and not
seeing.
contrast sensitivity function (CSF) is s haped like an

10
upside-dov"n ll,asshovvn in Figure 3.7 . 11le y-axis o n the
right is the observer 's contras t thresho ld. We o btain the
units for the left s ide of they- axis by taking the reciproca l
of the contrast threshold. For example, for a 1-<ycle/
100 degree gra ting to be jus t d isting uis hable from unifoml
u w 100 gray, the s tripes must have a contrast of about 1.0% (that
Spatitl frequency (cycles/ degree') is, if the m ea n luminance is 1000 ph o tons , than a tiny
patch of a U.ght s tripe re flects 1010 pho tons and a patd1
of a dark s tripe should refl e ct 9QO pho tons). The contras t
of a grating is generally specified a.coording to the Michelson definition, C =
+!,,,,,). example, C (1010 - 990) / (1010 + 990) 0.01
or 1% (0.01 x 100). The reciprocal of this thres hold is so tl1is is
the p oi nt plotted on the red CSF line in Figure 3.7 for this s p iltia.I frequency.
Note tha t a contras t of 100% corresponds to a contrast sens iti vity value o f
l. The CSF reaches this value on the far right sid e of the curve ln Figm e 3.7, at
about 60 cycles / d eg ree. Sixty cydes/ degree corresponds to a cycle \vid th o f 1
minute o f arc, the resolution limit we measured previou sly for high-contras t
contrast sensnMtv function s tripes, which, recall, is d etermined prima rily by cone spaci ng. The falloff in
(CSF) A !Unction doocntJng how tlie the CSF o n the o ther sid e of the curve cannot be explained by cone s pacing or
sensitivity to contrast (defined as the
reciprocal of the contrast threshdd) by limitations in the optics of the eye. lnstead, this part of the hm ction mus t
depends on the spatial trequency (size) be due to neura l factors, w h ich we will discuss later in the chapter.
of the stknulus. You can visu.:-i.lize your ovvn CSF by using F1gure 3.8. Here we see a sintts0idal
contrast threshold The smalloot grating whose contrast increases continuously from the top of the figure to
amount of contrast required to detect the bottom, an d '"'hose spa tial frequency increases continuously fro m the left
a pattern . s ide of the graph to the right. If you view the fig ure from a distance of about
FIGURE 3.8 A grating modulatir;:id b y contrast (:vMic.ally) and by

spatial frequency (horizon tally). (From Robson and Carni::bell. 1 007;
courtesy of Izumi Ohzawa.)

SPATIAL VISION 61
(o) Adapt.1tion level (b) Temporal modulation

500
,..Photopk
200
;,- Mesopic ;,-
100
50
10 ;\'.)
b
Ci 6
u
10
·.
100 2
10 60 100 0.2 d.5 1 5 10 20 50
Spatial frequency (cycles/degree) Sp<1tial frequency (cycles/degree)
(<) As•
1000 FIG URE 3. 9 The shape and height of
the contrast sensitl\'tty function is influ-
enced by a wid9 variety o f factors such
as adapt..'ltion l9Vel ternporal m odula -
tion (b) wh9rQ 90.ch c urve r9pl'&9Qflts th9
CSF at different temporal frequ9nci9s,
and age (c).
0.1 10 100
Spatial frequency (cydes/de,gee)
2 m eters, you wi U notice th e in ver ted ti s hape wh ere the gra ting fodes from
visibility to in\oisibility. lf you bring the book closer to your eye, you should be
able to see the stripes on the right side of the figure going farther up, whereas
the tops of the s tripes on the left side will become less di stinct_
There are ma ny factors tha t influence the exact fo rm of the CSF. These
include the adaptation level of the eye( R gure 3.9a), the temporal mo dulation
o f the targets (Figure 3.9b}, and the age (Figure 3.9c) and refractive state of
the subject.
Why Sine Wave Gratings?

O ne :answer to this question is that, although " pure" sine wave gra tings rn.a y
be rare in the real world, patten1S of stripes with m ore or less fuzzy bo undaries
are quite common: think of trees in a fores t, books on a bookshelf, and a map
of Manhattan (the latter includes a patten1 of horizontal s tripes superimposed
on a pattern of vertica l s tripes}. Furthennore, the edge of any object produces
230336 Amma Appa
a sin gle stripe, often blurred by a s hadow, in the retinal image.
On a larger scale, the visua l system appears to break d own real-world
images into a vast number of componen ts, each of whid1 is, essentially, a si ne
wave g rating with a particular spatial frequen cy. Titls method of processing is
analogous to the way in which the auditory system deals with sound and is
called Fo urier analysis (Figure 3.1 0; also see Chapter s 1and12). \¥e'll retun1
to this idea later in the chapter. For n ow, rest assured that scientists d on' t use
sine gra tings just because they're convenient to manipulate in experi-
ments (although they d o m ake veJy n ice sthnu1i).

62 CHAPTER 3
FIGURE 3.10 Fouri?r :;nalysls: the

left column shows sine WCN9S of dlff9r-
9flt spatial fr91qu9rlcies and .:vnplitudes.
From top to bottom: first row. the
.. fundamenta f' spatial frequency Vi with
amplitude = a: second row, 31, a = 1/3;
third row, 5f, a = 115: fourth row, ?f,
a = 1fl; fifth row, 91, a ""1/Q.
+l
-1
+l r +l
-1 \j\Jtf\Jl
WWJ_[
10
harmonics
All
10
Retinal Ganglion Cells and Stripes

Jn Chapter 2 we learned that retinal ga nglion cells respond vigorously to spots
of light. As it tums out, each gan glion cell also respon ds well to certain types
of .stripes or g ratings. Figure 3.11 shows how an ON-center retinal ganglion
cell responds to gra tin gs of diffe rent spatial freq uencies. \Vhen the spatia l
frequency of the grating is too low (Figure 3.1111), the ganglion cell responds
weakly because part of the fat, bright bar of the grating lands in the inhibi-
tory surround, damping the cell 's response. Similarly, when the s patial fre-
quency is too high (Figtue 3.lk), the ganglion cell resp onds wea kly because
both d ark a nd bright s tripes fa ll \·vithin the receptive-fie ld center, \Vashing
out the response. But when the spa tial frequen cy is just righ t (Figi.ue 3.11 b),
with a bright bar filling the center, a nd dark bars filling the su rround , the cell
responds vigorously. Thus, these retinal gang lion cell s are "tuned" to s patial
frequency: each cell responds best to Lt specific spatial frequency that matches
its receptive-field size, and it resp onds less to both higher and lower s patial
&equendes.
C hris tina Enroth-Cugell and John Robson (1984) were the first to record
the responses of retina l gang li on cells to si ne wave g ratings. In addition to
phase The relative poeltlon of a s howing that these cel ls respond vigorous ly to gratings of jus t the right size,
gretklg. th ese investigators d iscovered that responses depend on the phase of the

SPATIAL "1SION 63
{a) Low frequency yields weak re1>pOnse FIGURE3. 11 The r9Sp0ns& (right) of an ON-center retinal
ganglion cell to gratings of diff«ent spatial fr€quenvies Oeft):
l\Af\ 19) low, 'Pl medium, and (c) high.
I I I I
(b) Medium frequency yields strong response
1111111111111
--
(c) High frequency yields weak response
Amma Appd
llllllllfillllllll I II I
grating-its position within the receptive field . Agure 3.12 illustrates how
an ON-center retinal ganglion cell might respond to a grating of just the right
spatial frequency (a bar width about the size of the receptive-field center) in
four different phases.
\'Vhen the grating has '3 light bar filling the receptive-field center and dark
bars filling the surround (Figure 3.12"), this ON-center ganglion cell responds
vigorously, increasing its firing rate. If the grating phase is shifted by 00 degrees
(Figure 3.12b), hali the receptive-field center will be filled by a light bar and
half by a dark bar, and si milarly for the surround. In other words, there will
be no net difference between the light intensity in the receptive fie ld's center
and its surround. In this case the cell's response rate does not change from
its resting rate when the grating is tWTI.ed on-just what we would predict if
the ganglion cell were averaging the amount of light falling on its center and
its surround. A second 90-degree shift puts the dark bar in the center and the FIGURE 3.12 TM resp:>nS8 of a
light bars in the surround, producing a negative response (Figure3.12c); and a gangllon cell to a grating depgnds on
third phase shift returns us to the situation after the first s hift, with the overall the phase of the grating. This fgure
1Uustrat9S the responS8 of an ON-cen-
intensities in the center and surround equivalent and the cell therefore blind ter retinal ganglion cell to four dlfferMt
phasQs of an optimally sized grating.
(a) 0°- Positive response (b) 9\Y - No n?sparu;e (c) 1800-Negative response (d) 270° - No respanse

64 CHAPTER 3
to the grating (Figm e 3.12d). (Note that other ga n glio n cells loca ted to the
left or right of our target cell might respond to the 90-d egree and 270-d egree
p11ases but not to the 0-degree and 180-degree p h ases, w hich is w hy the visual
system as a w hole is able to see all four phases equally well .)
The Lateral Gen iculate Nucleus

l11e a..xons of retinal ganglion 1.."ells synapse in the two lateral genlculate nu-
clei (LGNs), on e in each cerebral h emisphere. These act as relay stations on
the 'VRY from tl re re_}ina t ecortex (see-Figure3.1). Flgure 3.13 s hows tha t
the LGN o f primates is a s ix-layered struch1re, a bit Like a stack of pancakes
that has been bent in the middle (ge11k11late means ''bent"'). The neurons in
the bottom two layers are p hysica!Jy larger than those in the top four layers;
for th.is reason, the bo ttom h,yo are called magnocellular layers, and the top
FIG URE 3.13 ThG primate lateral four parvocellular layers (mngua- and pnnJO- are Latin for 111.irge" a nd "small,"
ggniculate nuclGus. (From Breedlove respectively). The two types o f layers also cliller in another, more important,
and Watson, 20 13.)
way; the magnocellular layers receive input from M ganglion cells in the retina,
and the parvocellular layers receive input from P ganglion cells. TI1e layers
differ in m ore tha n the s ize of the cells. Stud ies in which m agnocellular and
parvocellular layers are chemically lesioned indicate tha t the magnocellular
pat11wdy responds to large, fast-moving objects, and the parvocellular path\,:ay
is responsible fo r processing details of stationary targets. This distinction is
interesting because it s hows th.:,t the vis ual system spli ts input fr om the image
into differen t types of information.
Even m ore spli ttin g takes place between the layers. TI1e[e we find the layers
consisting of koniocellular cells (konio- is G reek for "duse'; t11ese Httle cells
lateral genlculate nucleus (LGN) were ignored for man y years). Each layel' seems to be involved in different
A structure in the thalamus, part of aspects of processin g. For exa mpl e 1 one lnyer is specialized for relayin g s ign..lls
the mldbraln, that rooelves Input frcm from the 5-cones and may be part of a " primordi.:tl " blue-yellow path way (see
the retinal ganglion oells and has Input Chapter 5 on color vision) (Hendry and Reid, 2000).
and output oonnectlons to the visual
TI1e organization of the retint'l l inputs to the LGN, diagrammed in Flgure
cortex.
3.14, provides some importru1t insights into how our visu al world is mapped to
magnocellular layer Either of the
OOttcm two neuron-containing Byers
the brain. First, the left LGN receives projections from the left s ide of the retina
of the lateral genlculate nudeus. the in both eyes, and the right LGN receives projections from the right sid e of both
c ells of Vl11k:h are larger than retinas. Second, ead1 layer of the LGN receives input from on e or the othe r eye.
those In the top frur layefs. From botto m to top, layers 1, 41 and 6 o f the right LGN receive input from the
pavocellular layer Any of the top left (contralateral) eye, w hile layers 21 3, and 5 get their input from the right
four neuron-containing layers of the (lpsilateral) eye. TI1e koniocellular layers appear to fall in the s paces beh\'een
lateral genlc ulate nucleus, t he cells of each of the magno and parvo layers dearly labeled in Figure 3.14 (Cas.:'lgrande
which are phy'Scally smaller than those et al., 2007; Nossi and Callaway, 2009; Szmajda, Griinert, and Martin, 2008).
In the bottom two ler,rers.
koniocellular cell A neurm locatEK:I
between the magro::ellular and parvo- FURTHER DISCUSSION of koniooollular cells can be found in Chapter 2
cellular layers of the lateral genlculate c:npage44.
nucleus. This layer Is known as the
korJooellul9r layer.
contralateral Referring to the oppo- Each LGN layer contains a highly organ ized inap of a complete half of the
stte side of the txxJy (or brain). visua l field. Figure 3.14 s hows schematica ll y h ow objects in the right visual
ipsilatera Referring to the same field (objects to the right of where o ur gaze is fixated) a re mapp ed onto the
slJe of the body (or bralrJ. difierent layers oi the left LGN (the right side of the world falls on the left
side of the retina, whose ganglion cells project to the left LGN). ll1is ordered
topog-aphical mapping The
orderty mapping of the wortd 111 the lat- mapping of the world onto the visu al n ervous system, known as topographi-
eral genlc ulate nucleus and the visual cal mapping, provides us with a nemal basis fo r knowing w here things me
cortex. in s pace (we v.;U return to this point a little later).

SPATIAL \.1SION 65
Left visual field Right \rlliual field FIG URE 3.14 Input (in this case the letters ABCDEF) from the right visual fi9d
is rnapp9d in an orderly fashicn onto the diff.went lay91S o f the left LGN , and input
A B F fr:om the tett visual fleld is m apped to the right LGN . Information from the two eyes
is. S9gr99at9d into s9para!Q laYQrs. Layers 1 and 2 or9 th9 magnoCGllular
layers 3-6 are the parvoc91u1ar layers.
C •. E
LG N n euro ns ha ve co n cen-
tric recepti ve fie lds that are very
Left ey e: Right ey e s imilar to those of retina l ganglion
cells: they resp ond well to sp ots
and gratings. Given tha t the LGN
cells respond to the same patterns
a s the gang lion cells that provid e
their input, you might \o\'onder why
the visu al system bothers wi th the
LGN. Why don' t the ganglion cell
axons s llnpl y travel directly back to
the cerebral cortex? One im portant
reason is that the LGN is n ot m erely
a s top on the line fro m retina to
cortex. There are many connections
beru:een other parts of the brain and
1-"fiLGN the LGN (Babadi et aL, 2010; Dubin
o, and Cleland, there
F E
are morefeedb!chonn onsfrd ppa
F E o, the visu al cortex to the LGN than
F E o, from the LG N to the cortex.
o, ft seem s tha tthe LCN isa loca-
tion. where various parts of the brain
F E D2 c1n modulate in p ut from the eyes.
o, For exampl e, the LGN is pa r t of
a larger brain s tructu re called the
th a lam us (the media l genicu la te
nucleus, part of th e auditory path way, is a nother p ortion of the thalam us; see
Chapter 9). \ Vhen you g o to sleep, the enti re thalamu s is inhibited by circuitry
else\vhere in the brain that works to keep you asleep. Thus, even if your eye lids
were open while you were s leeping at night1 you would n ot see any thing in
your dimly lit r oom. Input w ould travel from your retinas to your LGNs, but
the ne ural signals would stop there before reaching the cortex, so they would
neve r be registered . The tha lami c inhibition is n ot co mple te, which is w hy
loud n oises (e.g. , the alarm clock) or bright ligh ts w ill be perceived , caus ing
you to wake up.
The Striate Cortex

If yo u place one hand at the back o f your head, about an indl or tvo.•o above
the top of your neck, you sho uld be able to feel a s mall bump known as the
inion . The receiv ing area for LG N inputs in the cerebral cortex lies below the primary visual cortex {V1), aea
inion Th.is area has several nmnes: primary visual cortex (V1 ), area 17, or 17. or striate cortex The area of
thl oerebral cortex of tl1e brain that
striate cortex (stria te means "striped ,'' for the striped patte rn Vl takes on, recavoo direct Inputs from the lateral
foll o\-ving a certain type of staining procedure). By now you' re prob a.bly ge t- nucleus, as well as
ting the idea that la yers are an important property of ne ural structures in the back trcm othlr brain areas.

66 CHAPTER 3
8230336 Am111a Appa
FIG URE 3.1 5 Striate ccrtax. (From HuOOI, 1988.)
vi sual pa thway. The striate cortex consists of six major layers, some of wh ich
have subl ayers (Flgure 3 .15). Fibers from the LGN project mainly (but n o t
exclusively) to laye.r 4, with mag noi...--ellular axons coming in to the upper part
of layer 4C (ktio\¥11 as4Ca) a nd parvocellula r axons projecting to the lower
part of layer 4C (known as (Yabuta and Callaway, 1998). (See Web Essay
3.2: The Whole Brain Atlas.)
Like the LGN, the s triate cortex ha s a syste matic topograp hica l mapping
of the visual field . But the s triate cortex is n o t simply a bigger version of the
LGN. A major a nd complex tran sformation of visua l i nforma tio n takes p lace
in the s triate cortex. For s ta rters, s triate cortex contains on the order of 200
million cells-more than 100 times as many as the LGN hasl
Fagure 3.16 illustrates hvo important fea tures of the vi.su.."'ll co1iex: topog-
raphy and rnagnification. First, the fact that the image of the woman's right
eyebrow (her right; it appears on the left in Figure 3.16) is mapped onto regions
corresp onding to the numbers 3 and 4 in the striate cortex tells the vi.sua l
system that the eyebrow mLLst be in posi tions 3 and 4 o f the visual field . Th.is
is topographical m apping. Second, information is dramatically scaled fro m
differen t parts of the visu al field. ln Figure 3.16, the fovea is represen ted by
number 5 on the retina. Objects im aged on or near the fovea are processed by
n eurons in a large part of the s triate cortex, but objects imaged in the far ri ght
or left periphery are a ll ocated only a tiny portion of the s tria te cortex. This
dis tortion of the visual-field map o n the cortex is kno\o\>T\ as cortical magni-
fication the cortical represen tation of the fovea is grea tly magnified
compared with the cortical representation of peripheral vision. (See Web Essay
3.3: Seeing Images on the Cortex.)
To gain a sense of the extent of this cortical magnifica tion factor, h old your
h-vo arms out in front of you, put up your index fingers, hold them abou t 10
centimeters (4 inches) apa rt, close your left eye, a nd fi xdte your right finger. In
th.is position, your right fingernrut w h..id1 is taking up about l d egree of visua l
angle on the is being processed by neurons in about 20 millimeters (mm}
cortical magnification The amount
of rortlc al area specmed In mil - of str iate cor tex. Yom left fingernail, which is covering th e s..1.me arn ount of
limeters) devoted to a speclnc region visual an gle but is falling 10 d egrees to the left of th e fovea, is being processed
(e.g .. 1 degree) In the visual field_ by only ] .5 nrn1 of cortex..

SPATIAL \/ISION 67
FIGURE 3. 16 The mapping of objects In spac€1 onto the

visual cortex. This figure lllustratgs both the tOJX)graphlcal
mapPng and the dramatic magnification of the foveal repre-
s9nta.tion In th9 cortex. (After Frisby, 1QSO.)
maAppa
Striat-eoortex
The Topography of the Human Cortex

Much of what we know aOOut cortical topography and magnification comes
from anatomkal and physiological sh1dies in animals. The earliest s h1dies in
humans were based on correlating visual-field defects with cortical lesions.
That is, someone with damage to this part of visual cortex would be blind in
that part of the 'isual field. Figure 3.17a illustrates how eccentricity (distance

68 CHAPTER 3
(a) (b)
L_____J
FIG URE 3. 17 Mapping thei visual 2.55102040 I an
field onto the cortex. as d9duoed from Eccentricily (degrees)
s tudying lasions (8) and from functional
MRI (b). diffQl'"Gnt parts o f thie
visual fie kJ are m a ppgd mto the cort1;1X
frorn the fo vea) 1n.:1.ps onto \o; sual corte.x, as deduced from s tudyi ng lesi ons.
according to the color c:009 pro vid9d . H owever, in the las t 20 year s or so there h:::we been major advances in our
The red area is just the central fgw knowledge of lumrn n visual cortex-in large pmt through developmen ts in
degrees. whereas the blue areas COVE( brain-imaging techniques. If you 'veever injured your knee or back, you may
substantially rrore o f the JXiripheral be familiar with rna.gnetic resonance imagin g (MRI ). MRI is very useful for
field . from 20 to 40 d egregs, (Part a
anato m ical imag ing of soft tissues, indud.ing the brain .
from Horton and Hoyt . 199 1: b from
Wan<1g1 andWlnawer. 20 11 .) lets us see the structure of th e bra in. Frmc tional m agnetic reson an ce
imagin g (frv1RI) is a n onin vasive technique for m easuring and localizing br ain
ac ti vity. As d iscu.s sed in 0 1a pte r 1, fMRJ d oes no t measure neural acti vity
directly. Rather, it mel1sures cha n ges in blood oxygen level that reflect neura l
acti vity. Blood oxygen le vel-depe ndent (BO LD) signa ls reflect a r an ge o f
metabolically demanding ne uml sign..1ls (\o\7.;mdell and Win awer1 2011 ). If you
compare blood flow w hen visua l s timuli a re p resented in o ne portion of the
visua l field to blood flow w hen the field is blank, you ca n find p ortions of
the brain tha t respond specilically to tha t s timuL.'l tion of th:1t p orti on of the
field. In th is way it is p ossi ble to map the top ography of Vl in the living lrn-
man brain (Figure 3.17b). Different p arts of the vis ual field are mapped o nto
Figure 3.1 7b \'\'ith a color code. Notice th.:1t the red area is just the central few
degrees. The blue areas cove r vast parts of the mo re periphe ral fie ld, fr om
20 to 40 degrees. Beca use of cortical magniHcatio n of the central these
red and blue regions are similar in size. In d1apte r 4, we w il l see that ffvlRI
can be used to localize o the r visual functions in stri ate cortex a nd beyondf in
''extrastria te" cortex, the cortex s urrounding s triate cortex. Recent advances
in fMRI m a ke it p ossible to recons truct the visual images vi ewed by humans
(Miyawaki et a l., 2008; N ishimo to eta! ., 2011 ) (Figure 3.18). ln the futu re, will
we be ab le to use fMRI to tell w hat yo u are thinking? Sta y tuned t
Some Perceptual Consequences of Cortical Magnification

One i. r nporta nt consequen ce of cortical rnagni firn tion is that vis ual acui ty d e-
clines in an orderly fashi on with eccentricity (Le vi, Klein, and A itsebaomo,
1985)-a phenomenon d em ons tra ted by Hermrum Rud olf Aubert well over
a century ago (Aubert, 1886). Web Ac tMty 3.2: Fove al Acuity allows you to
d em on stra te this phen.omen on yo urself, as d oes Figure 3.19, in which the
le tte rs are sca led in size s uch that each o ne covers an a pproxima tely equal
cortical area.

SPATIAL "1SION 69
(a) (b) (c) (d)

Presented D MAP reconstmction ---- Chance level
AHP reconstruction (p = 0.01)
mo\•ies
Highest posterio r
movie.s (1Yl4.P)
.-r
3rd highe:s.I .!i 0.2
" 0.1
51 S2
Reconstructed
m ovi es {AHP) Subject
FIGURE 3. 18 Using fMRI to n=icon -

struct Images from the brain ellcited by
natural movieis. (From Nishimoto et al .,
Why is the fovea l represenh1tio n in the cortex so highly magnified ? 1l1e 2011 .)
visua l system must ma_ke a trade-off. High resolution req uires a great number
o f resources: a d e n se array o f pho to recep tors, o ne-to -one lines from p h o to--
receptors to retinal gan g lion cells, a nd a large chunk o f s triate corte.x (n o t to
mentio n the real estate in o the.r are..1.s of cortex necessary to d o som ething '¥ith
the dsual infor ma tio n coming out o f Vl ). To see the entire vis ual field with
s uch h igh resolution, we might need eyes and brains too l1rge to fit in our
headsl Thus, we have evolved a visual system that provides high resolution
in the center and lmver resoluti on in the periphery. lf you need to process the
d etails of a n object in the com er of your eye, you c(1n s imply tum your eye or
head so tha t the object falls on the fovea ins tea d (see Chapter 8for a discussion
of eye m ovemen ts).
Interestingly, although visual acuity frills off rapid ly in the vis ual periphery,
it is not the major obstacle to readin g or obJect recogniti on. The real problem
in the pe riphery is kn own as visual crowding . Vis ua l crowdin g refers to
the d eleterious e ffect of clutter on peripheral object recognition {Levi, 2008;
Whitn.ey and Levi1 201 1). Objects that can be easily identified in isolation seern
indis tinct and jumbled in clutter (Figure 3.20).
Crowding is a n essential bottl e eek; tti.ng limi on ol!ject perception 1
eye and hand movements, visual searCh, reading.. and pe rhaps other fw1ct ions
in peripheral vision. Crowding impairs not only the discrimination of object visual c rowding The deleterious
features and contours but the ability to recognize and respond appropriately effect of clutt€f on p€flpheraJ object
to objects in clutter. Interestingly, crowd ed objects d on' t simply disappear. recognition.
w m
3 W
FIGURE 3. 19 A letter c hart in which the lettef size Increases with eccentricity in
proportion to the inVEfSQ cortic al magnification factor. 11}'OU 1ixatQ your gaze on the
far left side of the figure, all lettsrs should be equally easf t o sea because thc:ise
on the right. which are in the p.9rij:hery, are so muc h larg'1f.

70 CHAPTER 3
(a) (b)
= p
+ + +
C>
= --- c
'
::: w
p
FIGURE 3. 20 VBual c roV\l'dlng. (a) Vtsual crmvdlng occurs in natural \l\ih9n

fixating the bull 's-eye near the oonsttuctbn zone, note that it is d ifficu lt or
to recognize that tho8fe is a c hltd on the left-hard side of the road . sl mpt>; because o f
the prgsenoe of the nearby sig ns. Hawf:Ner, it is relattvely OOS'f to recognize the child
o n the light-hand While fixating each cross. it is easy to ldrentlfy the
Qeft cross), li ne o ri9""itation (middle cross), o r l9tto9r (right cross) abow It, t::ut it is dif-
ficult or imposslbl9 to identify the sarne shape, line orientation , or letter w hen it is In
the midcUe of a g rcup the cross. However. it is easy to do \/Yhen looking at the
patterns d ir9Ctt:/. Crowding In pgripheral \oision Impairs the ability to recognize objects,
but it cbBS not make them disappear. (From Whitney and Le\1, 20 11 .)
R.,ther,crowded objects appear "jumbled" (the target and neighborin g objects

are combined into a texture). crowding may be a process tha t simplifies
the a ppearan ce of the peripheral array by promoting consistent appearan ce
among adjacent objects, at the expense of an abili ty to p ick out individua l
objects. Lucki ly, \>Ve are a ble to make eye movements in order to fovea te nnd
scrutinize ind iv;d ual objects in clutter.
Receptive Fields in Striate Cortex

In 1958, David Hubel and Torsten Wiesel began work as postd octoral s tudents
in Stephen Kuffle.r 1s laboratory. TI1ei r goa l vi,•as to extend Kuffler's grmllld-
breaking work on retinal gan glion cells and to apply it to the cortex. So they
began trying to map the receptive fields of ne u rons ln striate cortex of cats,
using spots of light, much as Ku&1er (1953) had done earlier (see Ch apter 2).
Recall that the receptive field of a n eu ron is the region in space in whi ch the
presence of a s timulus alters the neuron 's firing rate (see Chapter 2). To Hube l
and Wiesel' s di smay. they found tha t a cat's cortical cells h ardly responded a t
all to the same sp ots that ma d e its gang lion cells fi re like crazy. To prnjecttheir
s timuli onto the retina, Hube l and Wiesel inserted a glass s lide with a black
spot in to a slot in a special op htha lmoscope (that's the i.n.stnunent the d octor
uses w hen s he sh in es a bright light into your eye in order to see your retina).
One day, they h ad been recording from a nem on without mud1 luck, w hen
s uddenly the cell e mitted a s trong burst of firing as they inserted the g lass
s lide into the slot. Eventually they realized that the resp onse had nothing to
d o with the s,pot itself; instea d, the cell had been :responding to the shadow
823 mst by the a.-lge of the glass slide as it swept across the ophtha hnoscope' s
light path . And the res t, us they 5.lly, is his tory. Hubel related this s tory w hen

SPATIAL \.1SION 71
FIGURE 3.21 Orientation tuning 1unc tion of a cortical

cell. The mR1ron fires vigorously when the line is oriented
but it fires hardly at all when the line orlffitation
is c han99d by 30 degrc;,es.
- ' I
01i1mt.1tion of line (degrees)
/ -
he Wiesel received the 1981 Nobel Prize in Physiology or Medicine for

uncoverin g many of the remarkable properties of the visual cor tex.
Hubel and \Viesel's most fundamental discovery was that the receptive
fields o f striate cortex n eurons are not ci.rctUar, as they are in the retina and
LGN. Rather:. they '1re elongated. As a res ult, they 1espond much more v igor-
ously to bars, lines, edges, and than to round spots of tight.
Orientation Selectivity
Ftu'ther investigation by Hubel and\ Viesel 0962) w1rovered a number of other orientation tuning The terdency
important properties of the receptive fields of neurons in str-iate cortex. First, of neurons in striate cortex to respond
an individual neuron will n ot respond equivalen tly to just :any old s tripe in to certain orentatkms and
less to others.
its receptive field . It responds best when the line or edge is at just the right
orientation, and hardly at all when the lli1e is tilted more than 30 degrees away
from the opti mal orientation (a ch an ge eciuivalent to movem ent of the min-
ute hand of a clock from 12 to 1). Scientists call this selective responsiveness
orientation tuning: the cell is hmed to de tect lines in a specific orient.:ition.
A typical orientation tuning function looks ILke the pl ot in R gure 321. The
neuron featured here fires vigorous ly when the line is o riented vertically, but
the response tapers off rapidly ns the line is tilted on e way or
ishlng to close to the cel l' s restin g rate when the line is tilted. about 30 degrees
in either direction. Other cells in striate cortex are selective for horizontal lines
and lines at 4-5 d egrees, 20 degrees, 62 degrees, and so on, so the population of
n eurons as a whole detects all possible orien ta tions. However, more cell s are
responsive to horizontal and vertical orientations than to obliqu es (De Valois,
Yund, and Hepler, 1982; B. Li, Peterson, and Freeman,2003). This physiologi-
cal finding meshes \·vell with the psychophysical finding that humans have
somewhat lower visual acuity and con trast sensitivity for oblique targets than
for horizontal and vertical targets.
Hm.,,.· are the circular recep ti ve fields in the LGN transformed into the
elongated receptive fields in striate cortex? Hubel and \Viesel s uggested a
very simple schem e to explain this transformation (Figure 3.22). Simply put,
their idea was that the concentric LGN cells that feed into a cortical cell are
FIGURE 3.22 Hubel and Wiesel's m ode:4 of how striate cortex cells gl?t their orien-
tation tuning: they hypothesized that LGN cells WEl("to lined up in a rCNV, feeding into
the ebngat9d arrang""1lQnt of ths. s triat€1 cortex reoeptive lields

72 CHAPTER 3
filter An aooustlc, eloctrlcal, ele:::- all in a row. Later studies (e.g., J. S. Anderson e t al.,2000) ha\re shown that the
tronlc, er optic de\lice, Instrument. arrangement of LGN inputs is indeed crucial for establishing the orientation
computer program, or neuron that selectivity of striate cortex cells. However, other evidence s uggests th at ne ura l
allows the passage of some range
o f parameters (e.g., orlentatlons, fre- interactions (e.g., lateral inh.ibition-seeChapter 2) w ithin the cortex also play
quencies) and blocKs the passage of an itnportant role in the d ynamics of orienta tion tuning (Pugh et aL, 20CO).
others.
ocular dominance The property
Other Receptive-Reid Properties
of the re:::eptlve ftelds of striate oortex Cortica l cells respond n ot just to ba rs, lines, and edges. Like retinal gan glion
neurons by which they demonstrate cells, they also respond well to gratings (which are, after all, collections of
a preference. respcndlng somewhal lines). And like gan g li on cells, they respond bes t to gratin gs that have jus t
more rapidly when a stimulus Is pre-
the right spa tial frequency to fill the receptive-field center. That is, ead1 s tri-
sented In cne "10 than when It Is pre-
sented In the other. ate cortex cell is hmed to a particular spatial frequency, which corresp onds to
a particular line width. lndeed, cortica l cells are much rn ore n arrowly tuned
simple cell A cortical neuron whose
receptive field has cleany defined exclt- (they resp ond to a s malle r range o f s patial freq uencies} than retinal ganglion
atay and Inhibitory regions. cells (De Va lois, Albrecht, and l11orell, 1982). These n arrow h.ming functi ons
mean that each striate cortex ne uron functions as a filter fo r the portion of
the image that exci tes the cell . We will re.h1rn to the idea of stria te cortex as a
collection of filters la ter in the chapter.
Another important d iscovery m ade by Hube l and Wiesel was that rrumy
cortical ce11s respond especially well to moi1h1g lines, bars, and gm.t-
ings. Moreover, many ne urons resp ond stron gly w hen a line moves in on e
direction---say, from left to right-but not a t all w he n the 5'1 me line moves,
say, from right to left.
As n oted earlie r, information from the n.. . . o eyes is kept separate in the
LGN: each LGN cell responds to one eye or the o the r, but n ever to both eyes.
1l1is arrangement changes dra1natically in s triate cortex, w here a majOJity
of cells can be influenced by inpu t from both the left eye and the right eye.
(a) Edge detector In other words, if a s triate cortex neu ron resp onds best to a 5-cycle/ degree
grating oriented a t45 degrees, it will resp ond to s uch a stllnulus whethe r that
s timulus is p resented in the right eye or the eye. However, striate cortex
neurons often have a preference, resp onding somew hat more st rongly w h en a
s timulus is presented in one eye than when it is p resen ted in the other. Hubel
and Wiesel ca lled thi s property of striate receptive fie ld s ocular dominance.
Given that we see a single, unified ,...,orld, in tuiti vely it makes sense that
infonnation fro m the h vo eyes sho uld be brought together at some point.
Until Hubel and Wiese l's di scovery; however, there were heated arguments
about w hether the infom1ation converged a t all and, if so, whether it was in a
specialized "fusion center" in the brain-a no tion that dates back to Descartes
(1664) (see I. P. Hovv-a rd and Rogers, 2001). \i\fe' ll describe some o f these issues
-1-
in Chapter 6, when we discnss binocular vision.
(b) Stripe detector
Simple and Complex Cells
Like p recortica l neurons, cortical neurons come in a wid e variety of types.
- - Hube l and Wiesel d 1aracte:rized n eurons as simple cells. Simple cells are
cortical neu rons '"'hose !'ecep tive fields have clearly defined exci tatory and
- - inhibitory regions. Figure 3 .23 sh ows tv.'o varieties of s imple-cell receptive
•
- - fields and thei r p referred s timuli. An edge detector (Figure3.23n) is rnost highly
exci te d when there is light on one s ide of its reC"q)tive field and d arkness on
the other side. A s tripe detector (Figure 3.23b) responds best to a line of li ght
tha t h as a particular v.-id th,strrrounded on both sides by da rknes.s. lf a gra ting
\vi th the appropriate s patial freq uen cy drifts across the receptive field of this
FIG u RE 3.23 Two flavcrs of simple cell, the cell's response w ill be m odulated as d ark and bright bars drift across
c ells : {a) an edge detector and (p) a the receptive-fie ld center, in exactly the same way the response of the reti nal
s tripe detector, ganglion cell shO\·Vn in Figure 3.12 is rnodulated .

SPATIAL \.1SION 73
FIGURE 3.24 call and a oompl9X c9ll might

both be tuned to the sarne orientation and striPQ width
(spatial frequency), but the complex: ci911 will respond to
Stim u.lui;; that stripe presented anywhere within its receptive 'fiQkj ,
whereas the simple cell might t o the s tripe in
only one !X>Sition.
Simple-a ll resi-"'Clnse -ttttttttHttt- +++-+

Complex-a ll response -ttttttttHttt- +++-+
Other n e urons s how responses that cannot be simply predicted fro m their complex cell A cortical neurcn
responses to s tationary bars of light. Hubel _a n Wiesel C:'alled these complex whose receptM> leld dooo not have
cells. Complex cells are cortical ne urons \"l1hose receptive fie lds d o not have clearly delned excitatory and Inhibitory
regkJns.
dearly defined exci tatory and inhibitory regions. Like simple cells, each com-
plex cel l is htned to a p articu lar orientation and spatinJ frequency and shows end stopping The process by
which a cell In the catex lrst lncreasoo
an ocular preference. However, whereas a stmple cell might respond only if a Its Iring rate as the bar length
stripe is presented in its receptive field, a complex cell w ill respond Increases to Ill up Its receptive field,
regard less of where the s tripe is p resente d, as long as it is somewhere within and then decreases Its Mng rate as
the cell's receptive field (Figure 3.24). When tested w ith a drifting gra ting, the bar ls lengthened further .
the complex cell gives a robust response1 \.'1.-ith little or n one of the modulation
sho\>\'ll by si mple cells (as well as retinal gan glion and LGN cells). Another
way of stat ing this difference is to say th:lt si mple cells are 11ph...1Se-sensitive,"
<'lnd complex cells a re " phase-insensi tive."
As with all other n e tuo ns in the visual system , \vi th the excep tion of retinal
photoreceptors, the receptive fields of complex cells represent a pooling of the
responses of several subunits. The s ubunits give the complex cell its spatial
freq uency and orientation tuning, but the complex p ooling operation makes
the co mplex cell insensitive to the precise positi on of the stimulus within its
receptive field _ Hubel and \Viesel hypothesized a h ierarchy in which LG N
cells fed into simple ce lls, w hich in tum p rodd ed excita tory inputs to complex
cells. However, substantial evidence now suggests that complex cells represent
a separate path way (that is, that both simpl e and complex cells get
direct inpu t from LGN neurons).
Further Complications
Hubel and Wiesel described another property of some cells in striate cortex
that they called end stopping. When they tested an end -stopped cell with bars
of increasing lengths, the response rate first increased as the bar filled up the
cell 's receptive field, and then decreased markedly as the bar was lengthened
further (Figure 3.25). Hubel and Wiesel ca lled. these cells "h ypercomplex"
cells, al thou g h they n ow appear to be subclasses of the simple and complex
cells already discussed h ere (that is, there are s imple end-stopped cells and
FIGU RE 3.25 When the stimulus

i):)ar) d09s no t nmc h the outside edgi9
o f the field or It extends
beyond the receptrve field o f an end-
stoppOO cortical neuron, the neuron
fire s less than whEn the stimulus is just
+++-+ +++++++ +++-+ tha right length.

74 CHAPTER 3
complex end-s topped cells). End stopping is thou ght to play an important role
in our ability to de tect luminan ce boundaries and discontinuities.
R ecent researd1 h.'ls
. reveal ed additional idiosyncrasies in the receptive fields
of striate cortex neurons. For example, the size of a particular cell's receptive
field a ppears to vaiy \·vi th target contrast; for instance, the cell might respond to
a s maller portion of the visu al field '"'hen the gratin g stimulus has a high con-
trast than it wi ll when the differen1..--e behveen light and dark bars is m ore s ubtle
(Sceniak et a l., 1999). And ne urons can be influen ced by stimuli that fall outside
the classic recepti\'e field, via sh o rt- o r lon g -range lateral connections and/or
via feedback from neurons in other layers (Zipser1 Lamrne,and Schiller, 1996).
As is the ca.se for mo.st of the visual system, w hat \\-"e d on't know about
the workin gs of s triate cortex neurons almost certainly dwarfs what we d o
know. But to review what ""·e do know, spend some time Web AcUvtty
3 .4: Striate Receptive Fields, w here you can try your hand at detern,ining
the receptive field of an tm.known vi rtua l neuron.
Columns and Hypercolumns

As. we've discussed, ead1 of the approximate y 200 mi"Hion neurons m s iate
cortex responds to a d istinctive set of stimulus properties: stripes, edges, and/
or grntings tha t a.re oriented a t a particular angle, \vith a particular width or
spatia l freque ncy, possibly moving in a particula r direction. Some neurons
are si mple ce ll s and some Me complex cells, and each one is end-s topped o r
n ot. Most ne u rons also respond preferentially to stimuli presented in one eye
or another. And each n e uron responds only w hen its p refe rred stimulus is
presented in one particular part of the visual field .
Hubel and Wiese] noticed very early on tha t these va rious recep tive-fi eld
properties are not .sca ttered haphazardly around. stria te cortex. Once they h ad
figured o ut w ha t the cells were looking for (stripes ra ther thi:ln spots), they
d iscovere d that if they pushed the recordin g electro de down through the
layers o f the cortex in a direction perpend icular to the cortical s urface, all the
cells they e ncountered s howed si milar orienta tion preferen ces. lf they shi fted
the e lectrod e position over a tiny distance and m.ade an other perpendicular
penetration, a ll the cel ls then !'esponded best to a slightly d ifferent orientation,
perhaps 10 or 15 degrees from the original orientation. On the basis of these
observations, Hubel and Wiesel concluded that neurons w;th similar orientation
preferences are arranged in columns tha t extend vertically throu gh the cortex.
\Vhen Hubel and \.Yiesel m ade tan gential penetrations into striate cortex
(inserting an electrode in a d irection parallel to the cortical surface, rather than
perpendicular), they found a sys te ma tic and progressive change in preferred
orientation and en cmmtered essentially aU the orientations in a distance of
about 05 mm. TI-US finding has been confinned via alternative physio logical
techniques. Figure 3.26a shows a sm a ll portion of a mo nkey's s triate cortex
prepared so tha t neurons responding to vertically oriented lines are s tained
black, w hile o ther neurons rem ain V•thite. The between the vertica l
orienta tion co1mnns revealed by this technique is, s ure enough 1 just about 0.5
mm (LeVay, Hubel, and Wiesel, 1975).
Orient(1tion is not the on1y prope1ty armnged in columns in the visual cortex.
Neurons tha t sh.:1.re the sa rn e eye preference (exhibiting ocular dominance)
also have a coh.mmar arran gem en t (Figure 3.26b). Furthennore, single-cell
column A vertical arran-;iement reco rding experi me n ts, as well as s taining exp e rirne nts, indicate tha t eye
of neurons. Neurons within a single
column 1€fld to have similar recep- preference swi tches {you g uessed it) every 0.5 mm or so.
tive fields and slrnltar orientation Roughly 25 years ago, it becam e p ossible to make detailed m aps of the
preferencee. orien tation ti.ming of cells in the cortex, using optical imaging. Figure 3.26c

SPATIAL \.1SION 75
(a) Orientation columns (b) Ocubrdominaru:ecolumns
(c) Orient<1ti on mc1ps

FIGURE 3.26 Orientation (a) and ocular dominanc .oa Ip) columns of
the striate cortex. revealed by staining . (c) Optical imaging o f the ori-
entation maps In m::mkey cort9X. (Parts a, b from Hubel , WiesEll, and
Stryker. 1078: c from Nauhaus et al ., 2008.)
sh ows a n optical imaging study in w hid-1 light reflected from the s urface of
the exposed brain reflects the activity o f underlying neur ons. In th.is figure,
each color s hows a set of neurons, activated by one orientation. You can see the
complex, yet orderly organization of tuning in the cortex (Blasdel
and Sa.lama, 1986). Recent work (Paik and Ri..ngadt, 2011) suggests tha t this
bea utiful arrangement arises during ear ly development as a consequence o f
statistical wiring mech..1.n isms combined wi th evenly s paced of ON-
and OFF-center retinal ganglion cells.
Th.rough their s tudies, Hube l and Wiesel a rri ved at the model of striate hypercolumn A 1·rnllllrneter block
cortical architech.ue illustrated in Figure 3.27. Hley proposed that a 1-mm block of striate cortex contalnlt>;J two sets of
columns. each covering fN"Y possible
of striate cor tex contains 11all the machinery tq I 00 after "''CWth..ing
Ollentatlon (0-1BO degrees), with one
the dsual cortex is responsible for, in a certa in s mall part o f the visual world 11 set Pfeferrlng Input frc:m the Jett &;e
(Hubel, 1982). Each o f these sections of cortex is caUecl a hypercolumn. It and one set prefenlrg input from the
contains at least P.vo sets of colurnns, each coverin g every possib le orientation right "f9.

76 CHAPTER 3
.
:: f i
Color code fo r orien tation colum ns
! ; : :
:
lllZJDJll
FIGU RE 3 .27 This model of a h)'Pef- Right-eye
l....eft-:e)'i!
oolumn ocular dominance ocuLu dominance slab ocular dominance sk1b
columns (one for each eye) and many
orlentaticn columns, and Illustrates the
locations of the cytochroroo oxidas9
(00) tjobs. (from E<eedk>ve and Wat- (0--180 degrees), \-vith one se t prefe rring input from the left eye and one set
son, 20 13.) preferring input fro m the right eye.
Hyperco lunms are rou ghly 1 mm across throughou t thes tri.atecortex, but
because of the cortical magnification fuctor discussed ear lie r, not all hyper-
co lumns see the world a t the s.."'me level of d etail. A hypercolumn in the part
of the cortex tha t represen ts the fO\•ea m ay "see" a p orti on af the vis uaJ field
that is 0.05 degree of visual an gle acrossj a hypercolumn responding to input
10 d egrees to the right of the fove;.:i s hould cover about 14 times as large an
area (0.7 d egree across).
O rien tation and oc ular dominance are probably not the onl y s timulus
dirnens ions that have a system.a tic colunmar arrangement in the visua l cortex.
For example, another staining teclmique, which takes advantage of an enzyme
called cytochrome oxldase (CO}, has revealed a regular array of "CO blobs "
(sh own in section in R gure 3.28), spaced that magical distance of about05 rnm
apart {see Figure 3.27). Tl'le functional role of these blobs remains unclear, but
CO blob columns have been implicated in processing color, with the interblob
regions (note the elegant scientific jargon that has de veloped around this field
o f stud y) processi ng m otion and spa tial structure (Livingstone and Hubel,
1988). This view is probably too si mplistic, but the blob array d oes sugges t
som e kind of additi onal organizational layer on top of the o rientation and
oc ular d o1nin.1.nce arrays.
FURTHER DISCUSSION of CO blobs. which are also implicated in pro-

cessing cobr, can be found in Chapter 5 on pages 13Q- 140.
ln sum, the current sta teof understa nding is tha t striate cortex is concem.ed
with ana lyzin g the on enthtiOn, ze}Shape; S'peed1nnd direction of motion of
objects in the world, and that it does so using modular groups of nei.uons-hy-
cytochrome oxldase (CO) "'1 percolunrn.s--ead1 of w hidl receives input from and processes a s mall piece of
enzyme used to reveal the regular
array of ·co t1obs," 'lltllch are spaoed the visual world. (You can explore these organizational principles interactively
about 0.5 m lllimeter apart In the pri- in Web Activity 3.5: Hypercolumns.) We can think of this arrangemen t as
mary visual conex. a big bank of filters. Combinin5 information fro m multiple hypercolumns

SPATIAL VISION 77
FIG URE 3.28 Cytochrome oxldase (00) t]obs. (From Hubie!, 1988.)
is p restm1a bly the job of o ther p or a ns of cortex farther downsheam m the

vis ual system. We ,....;11 consid er some o f these portions in Cha.pter 4, w hen we
d iscuss the representa tion a nd recognition of wh ole objects.
Selective Adaptation:
The Psychologist's Electrode
Most of the physiologicrd research reported up to this p oi nt in the c hapter adaptation A reduction In response
was d one using cats, monkeys, or other a ni mals as subjects. Does the human c aused by prior or continuing
stimulation.
visuril system also include n eu rons selecti ve for orienta tion, Line '"·id th, direc-
tion of m otion, a nd so on? The difficult thing about a nswering this questi on
is tha t we can ' t nom1ally poke electrodes into a huma n 's brain (which is w hy
Hubet Wiesel, and their peers had to u se rntsand m on keys in the first place),
so ind irect me thods of learning about brain functi on lrnd to be d evised. O n e
su ch me thod is m lled adaptation, a technique that gives psych ologists a non-
invasive "electrode" they can u se to probe the huma n bra in. Alth ough o ften
attributed to John Frisby (1980), the te rm ''the psychologist's electrod e'' was
act ually coined by Colin Blakem ore in the 1970s..
Selecti ve ad aptatio n can p rovide insig hts into the properties of cortical
n eurons, as illustra ted in Figure 3.29. 1l1e green bars in Fig ure 3.29a illus tra te
the n om1al firing rates of cells hmed to vario us orie ntation s (0, 10, -10, 20, and
- 20 degrees, and so on) relative to a ve rtical gra tin g. By d efinition, grating.s
oriented a t 0 degrees (vertical) e licit the strongest respon se fr om the 0-degree
selective cel ls, followed closely by tl1e JO-degree a nd -J O-degree selective cel ls,
fo llow ed by the 20-degree and -20-degree selective cells, a nd so on (see Figure
3.29a ). Now suppose we expose the visual syste m tha t contains these cells to a
20-degree gra tin g fox an extended p e riod of time. TI1is ada pting stimulus wi ll
cause the 20-degree selective cells to be most active, and the extend ed acti vity
w ill fati gue these cells (that is, thei r maxi mum firing ra te w ill be reduced fo r a
short pe riod follow in g ad aptation ). 1l1e adaptation procedme w ill a lso affect
the o ther cells to some extent: th e 10-degree and 30-d egree cells v..dll be the
next most fati gued , foll owed by the 0-degree and 40-degree cells, and so on .
Figu re3.29bshows lvha t should ha ppen w hen we present the vertical grating
again afte!' adapta tion to the 20-degree gra ting, assuming that our orientation
perception is really due to p opula tions of orien tation-selective ce!Js like- those

78 CHAPTER 3
FIGURE 3.29 The psyctologisf s (a) Before ad.1pt.1tion

electrod flo, This scheimatic diaQram
High
shows how Si9lec tfv& adaptafon may
alt.er th9 distribution of nrural respcns-
es and therefore perception . See the
text for details.
Relative orient,1tion of line (degree&.)
•
Adapt to 21Y
(b) After adaptation

High
90
Relative olientation of line (degrees)
tha t Hubel and \ Viesel fm.md in the cat cortex. As the dark g reen bars s how,
because the 0-d egree cells have been fa tigued ni.ore tha n the -10-degreecells,
the -10-degree cells a re now firin g fas teS:t.. As a z;esult, we s h ould perceive the
vertical test s timulus as being oriente d 10 to the le.ft.
You can tes t the validi ty of this technique yourself using th estim uli in Rgure
3.30. Start by carefully fi xating the black line between the two sets o f til ted
s tri pes in Figure 3.30a for a fe\<\' seconds. This exercise w ill result in adaptation
to left-tilted strip es in the upper visual field, and right-tilted stripes in the
lower field. Now, look quickly a t the fixa tion point between the two sets of
vertical stripes in Figi..u e 3.30b. M ove back and fo rth betvveen a. few seconds of
the adap ting stimuli (Figi..1re 3.3011) and a quick look at the test .stimuli (Figure
3.30b). You s hould n otice th.:'1 t Fig m e 3.30a looks like a chevron 1 p ointing right
and Fig ure 3..30b has come to look like it p oints a little to the lefl That is the
tilt alteref1ect, ju.st as predicted by the m od el o f the human visuaJ system
based on the cat rese.u ch and d iagrammed in Figure 329. The tilt aftereffect

SPATIAL 1'1SION 79
(a) (b) FIG URE 3.30 Stimuli for demo nstrating

s91ective adaptation. text for details.
(M er Frisby, 198J.)
Ill II
r. .............
8230336 Arnn a Appa
111
strongly supports the idea that the htm\an visua l sys te m conta in s individual
neuro ns selective for different o rientations.
Selectivea daph1ti on also provides evidence that the human visuil1 sys tem
contains ne urons selecti ve for spa tial frequency. You can check thls w ith the
gratin gs shown in Ftgura 3.31. First, look at Figtue 3.31b and make a me ntal
no te of yo ur contrast sensitiv ity functio n (the inverted U-shaped area vvhere
th e gratin gs fad e into the grn y background ). Next, ad ap t for about 10-20
seconds to the grating in Figure 3.31n, and then quickly s hift your gaze back
to Figure 3.3l band make a rnental note of your CSF now. After you repeat this
proced ure a few tim es, the Ol.1tline of your CSP in Figure 3.31b s hould look
som.ething like the red curve in Figure 3.31c. lt should have a notch {indicating
reduced contras t sensitivity) for spatial frequencies that are d ose to the adapt-
ing spatial frequency (Figure 331n). This demons tration shows that adaptation
to the high-contras t top panel is selective- resulting in a loss of sensi ti vity
for spa tial frequencies dose to the ad ap ting frequency, but n ot for s patial
frequencies tha t are much hig her or lower than the adapting frequency. Note
that adaptation to the bottom panel (Figu:re3.31d) d oes not result in a sin1Uar
notch, since spatial-frequency adaptation is o rientation -selective. Thus, there
is little or n o effect on sens itivi ty to vertica l gratings following adaptation to
a horizontal grating.
As noted earlie r, selective adaptation causes the ne m ons m ost sensitive
to the a dapting stimulus to become fatigued . h1 thi s d emo nstra tion , ne urons
sensi tive to the spatial frequency of the adapting sti mulus ha\·e thei r contras t
sensi tivity reduced. That is, hig he r contrast is need ed, afte r adaptation fo r a
test grating, to s tllnulate the.se neuro n.s_ Neurons responsive to much highe r or tilt atterellect The perceptual lllu-
mud1 lower s pa tial frequen cies are n ot fatigued by the adaptation procedure, sicn of tilt , produced by adaptation to
so contrast sensitivity for these s patial freq uencies is not affected . a pattern of a glveri orlentatlcn.

80 CHAPTER 3
(a) FIGURE 3 .3 1 A dem:instratbn of adaptation that is spgc11ic to ·spatial fre-

quency. (a) The adapting grating. rp) A grating modulated In contrast (vertically)
and spatial frgquency Oiorizonto.ll)'). This pattBm 19ts you visualiza your O"Wn
contrast sensitMty function (CSF) (see also Rgure 3.8}. Before adaptation, your
CSF should have the appearanc.9 of an inverted U. After adaption to (a), your
CSF should look something IIke the red c uive in ii';}. The n;id curve illustrates the
effect o f adaptation. The n otch indicates reduced contrast sensitivity for spatial
fr9quencies that aro9 dose to the adapting spatial frgquency. Adapting to thB
horizontal grating ir1 {d) results ir1 little or no reduc tion in S9nsiti\'ity. (Part b from
Robson and C ampbGll, 1QQ7; courtQSY of Izumi Ohzawa.)
Fig ure 3.32a shows m ore precise ly h ow selecti ve adaptation to cl

7-cyd e / d egree gra ting produces a selective loss of contrast sens itivity at
(b)
spatial freq uencies of about 7 cycles/ degree, with little or no loss at, for
example, 1 cycle/ degree or 15 cycles/ degree. Alter adaptation, the CSFhas
a "n otch," as if the detectors sensi ti ve to spatial freq uencies near 7 cycles/
degree were selectively desensitized (luckily the effects of
adaptation are reversiblel). From these measureme nts of contrast sensitiv ity
before a nd after adaptntion, we can construct the s patial-frequency tunin g
function sh own in Figure 3 .32b. TI1is fun ction represents the change in
contrast thresh old (contras t threshold after adaptati on, divided by contrast
threshold before a daptation ) pl otted against the spatia l frequency of the
test grating. 111is curve represents the spatial-frequency tun ing functi on
fo r a "channel" tha t is most sens itive to a grating of 7 "-" )·des / d egree.
The s h ape and selecti vity of this channe l are very similar to the spatial-
(c) frequency tuning functions for striate cortex neurons of cats a nd rnonkeys.
Figure 3.32c illustra tes that the CSF (gray curve) represents the "upper
envelope'' of the sensiti vi ties of m any spa tial-freciuency channels, ead1
hmed to a d iffe re nt spa tial freq ue ncy. The key idea h ere is that you can
see the pattern when at least one mechanism (dumnel) detects it.
The Site of Selective Adaptation Effects

The adaptation experiments rep lica ted here p rovide stron g ev;d ence tha t
orientation and spatial frequency are coded by n eu rons somewhere in the
hwnan visual system . In cats and m onkeys, we know that these neurons
are located in striate cortex, not in the retina or LGN. Can we locali ze the
orientation --selec tive and s patial frequency-selective neurons in humans?
As it turns out, we can do just that w ith a clever variati on on the
(d) adapt.a ti on experimen ts. Repeat the orien tation (Figure 3.30) and spatia !-
frequency (Figure 3.31) adaptation demonstrations, but th is tim e view
the ad apting stinnUi \vith your left eye only (keep your right eye cl osed
dtuing the adaptation period), and then view the test s timuli wi th yom
right eye (d ose your left eye and open your right eye as you shift your
gaze to the tes t s timuli). You s h ou ld find tha t the tilt afte reffect and
the decreased contras t sensi tivity transfer from one eye to the other,
although the effect may be somewhat less pronounced th{m w hen you
did the de mon stra tions with bo th eyes open. This transfer of adaptation
from the ada pted to the n onadapted eye is known as interocular transfer
(C . Blakemore.md Campbell, 1%9).
NO\.J?.ecaU thttt mfom1ation from the t\.'/oeyes is kept complete ly sepa-
rate in the retinas a nd in the two LGNs; n o s ingle neuron receives input
from both eyes tmtil the striate cortex. The transfer of adaptation effects
from one eye to the other thus implies tha t selective adaptation O...""C"ltrs in
cortical neurons, just as we would predict from animal physiology s tudies.

SPATIAL VISION 81
(a) (b) 4 (c)
..
100 Jo> 100
."
·Oi
&
E
lO
-ii
] 1'
j
10
Cl
1 1
1 10 100 1 rn 100 10 100
5pati.ctl freq uency (cycles/ degree) Spatial fr.?q uency (crcl.e&/ degi"'e) Spatial frequency (cyd.:-s/degree)
FIGURE 3.32 Spatial-fr9quElncy adaptation. {a) S&l9Ctiw adaptation to a 7 -cycJQ/

degreg grating produces a selectivie loss o f contrast siensltMty at spatial fr9quencies
of about 7 cyclWdegreg, lea\1ng a notch Qndicated by red line) In the CSF (gray line).
f/J) Threshold 1918\18.tion (th9 change in contrast threshold) fo llowing adaptation.
{c) Physlologicalt'f rneasured tuning functions for str1at9 cortID.": n9u-
rons In (cola-eel c urves represent d iffer9nt neurons). Th9 gray c urve. the
CS F. represents the •upper envelope" of the sensitMties of the underlying channels.
(Part b atte< c. Blakemore and campbsll. 1Q69; c aftE< Maffei and FbrMtinl, 197:3.)
Spatial Frequency- Tuned Pattern Analyzers in Human Vision

Selective a daptatio n to spatial frequency, as well as other eviden ce, provides
strong s upport fo r the notion, first s uggested by Fergus Campbell and Jolm
Robson (1968), that the hum,m CSFactually reflects the sensitivity of multiple
individual pattem an alyzers. l11esepattem analyzers are implemented by en-
sembles o f cortica l neurons, with each set of ce lls tuned to a limited of
spatial hell uencies and and they are often referred to as spatlal-
frequency channels. Remember the initially unexp lained falloff in the CSF
at very low spatial freq uencies? A lth ough a number of different explanations
have been su ggested (e.g., Lateral inhibition), the most likely expla nation is
that we simply have fewer neurons tuned to low spatia l frequencies (De \'alois,
Albrecht, and Thorell, 1982).
The multip le-spatial-frequency m odel of vision implies that s patial frequen-
cies that s timulate different pattern analyzers \'\.·ill be detected independently,
even if the different frequertcies are combined in the srun e image. Consider the
compound grating pattem in Figure 3.33, m ade by adding a sin e \vave with
frequency J to a sine w •.we \.Vith frequency 3f Graham a nd Nachmias (1971)
fotmd tha t the contrast sensi tivity for this compou nd pattern was a lmost the
same as the contrast sensi tivity for detecting the individ ual comp onents o f the
pattemsepar.ately. If the two component sine waves had stimulated a conunon
pattern analyzer, then their effects o n the a nalyzer s hould have been added ,
so con trast sensitivity should have been grea tly improved. Th.is may sow1d
a bit like the Fourier analysis idea discussed early in th.is ch apter. However,
it's now pretty dear that the visual system doesn't carry out a n actual Fomier
an alysis, ana lyzing the world into very narrow bands of spatial frequencies.
Rather, it filters the image into s patially localized receptive fields that have a
limi ted range of spatial frequencies. (Som ething like a Fourier analysis ma y
be importa nt in quickly apprecia ting the gis t of a scene; see C hapter 7.) spaUal-frequency channel A
pattern analyzer, implemented by an
Wh y would the visual system use spatial-frequency filters to an.."l..lyze. im- ensemble of cortical na irons, In which
ages? One important reason may be that different spatial frequencies emphasize each set of neurons Is tuned to a lim·
different types of information. Fig ure 3.34b and c sh ow only high-frequency lted range of spatial rrequercles.

82 CHAPTER 3
f
l=IG URE 3.33 A compound grating
pattern (right), made by the additbn of
a sine W'iNe of frequency f (top !&ft) to
ong of frequency 31 (bottom left). (After
Graham and Nachmias, 1971.)
f+ 3f
!WW
FIGURE 3.34 A complete image o r low-frequency components, respectively, of the face in Figure 3.34a. These
and reconstructions: with the low images show that low frequencies (Figure 3.32c) emphasize the broad outlines
spatial frequ9f"'lci9s r9m0Ved, 19raYlng o f the face, and high frequencies (Figure 3.34b) carry infonuation about fine
only the high-frequency oomponents o f
details. If we want to know how many people are in a scene, it is most efficient
that Image f/:>), and with the hlgh spa-
tial fnlJquenclgs removed, leavlng only to consult our low-frequency channels. But if we want to hi.ow about the fine
the low-frequency oomponsnts of that details--for example, whether the people are frowning or smiling-we must
lmag9 (c). For another 9Xa.mple, SQQ rely on our high-frequency channels.
Figure 1 . 1 8.
(•) (b) (c)

SPATIAL \.1SION 83
FIG URE 3.35 Who is hidden behind the high-spatial·frequen·

c:y mask in this image?
It is ""·idely agreed th at at nea r- thres h old contrasts1 pa tte rn a nal yzers

operatin g at differen t scales of analysis are independent. At high contrasts,
however, pattern .:malyzers d o internet. You can experience this fo r yourself
in Figu re 3.35, w here the hig h .sp,1tial frequencies introduced by the s rnC1U
blocks mask the low spa tial frequencies that ron ve}' an und erlying p ortrait
of a famous Atnerican. Squinting your eyes will blur the blocks, minimizing
the e ffect of the mask to a point where the face you've seen countles.s tirnes
on the 5-do1lar'blll will prob!: bly s'how tl1 rough.
The Development of Spatial Vision

William James (1890) described the infa nt's world as "3 blooming, buzzing
confos ion ." However, s h 1dies over the past severa l decad es ha ve s hown that
the visual system is much mo re d eveloped a t birth than we used to think.
One of the difficulties in assessi ng vision in infants is that we can 1 t simply
ask them what they see. Ra the r, we have to think of tricky wa ys to coax that
informati on from them . The m ost widely used method for studying infant
vision is base d on an observation that Robert Fantz made in the early 1960s.
What Fantz n oticed is th at if infants a re shov..rn two scenes, thev
s tare a t the m ore complex scene (the scene with the m ost So,
infant is sh ovvn two patches, one containing s tripes and the other uniform
gray, the in fant will prefer to look at the s tripes. Of cotuse, an infant \vho
couldn' t see the stripes would be equally likely to s tare at the gray patch
as at the s triped pntd1. Thus, p referential looking is on e important method
used by infant resea rchers (grown psyd1ologistsshld yi ng infant vi sion, not
babies in lab coats) to lea rn what infants can see and respond to behav ior-
ally (Figure 3.36a).
The su ccess of preferen tial lookin g depends on the willingne..ss of babies
to s tare a t sti muli near thres hold level. An a lterna tive approach , used with
considerab le s uccess in m ore recent is to m easure visu ally evoked

84 CHAPTER 3
(a) Stimuluscan:I
(b) (c) Sweep VEP (grating .mlity)
0 10 15202530
Sr atial frequ.:>n cy
(cydes / degree) Thresho ld= 27.6
FIG URE 3.36 Assoo.slrg vision In

u- mrn
Stimuli cycle>/ degree
infants . (a) Forced -choic9 preferential-

stlmuli Oeft) and the experi- electrical p otentials (V EPs)- that is, electrical signals from the brain that are
rnental setup l)'ight). (/)) Visually evokOO evoked by visual s timuli-by a ttadtln g electrodes to the scalp and measur-
f:'.>OtMtial {VEP) SQtup. (c) R1;1sults of a
ing the ch an ges in electrical acti vity tha t are elici ted by the changin g visual
sweep VEP experiment in which the
tr.equB'"lcy of the stimulus is
s timulus (Figures 3.36b and c). Using this technique, we can meas ure an entire
swept (continuousty variOO from k:iw contrast sens itivity hmctio n in as littl e as 10 seconds in a n onverbal infant.
to high spatial frequency), ii lust rating
the @<.trapolated acuity (thrg,shold-in Development of the Contrast Sensitivity Function
this case, 27 .6 This 11'1e emerging pi chue su gges ts that sens itivity to low spa tia l frequ encies d e--
partia.Jlar 9XpQrim""'1t w as cbne at 80%
contrast. (Part c after Norcia. T)'1er , and velops much mo re rnpicily than sen:siti ...i ty to high s patial frequencies. Thus,
Ham9r, 1QQO.) at low spatial frequencies, contrast sens itivity may reach nearly adult levels as
early as about 9 weeks of age, whereas sens iti vity at higher spatial frequencies
continues to develop dramatically (Figure 3 .37). TI1ere remains a subs tantia l
difference in the con tras t sensitiv ity of adults and 33--week-o lds at high spatial
frequencies (N orcia, Ty ler, and Hamer, 1990).
\Vhat limits the d evelopment of acu ity and contrast sens itivity? TI1e pri-
mary p osb1 ata l changes in the retina concern differentiation of the m acul ar
region (Boothe, Dobson, and Teller, 1985). After birth, receptor d ensi ty
and cone outer segment len gth both increase, as fovea l.con es become thinne r
and more e longated. There is a dra matic m igration of ganglion celJs and inner
nuclear la ye rs from the foveal region as the fovea l pit d evelops d ming th e firs t

SPATIAL \.1SION 85
FIG URE 3. 37 The dw91o pment of contrast SQt"lsitivity. Note t hat the shaPQ of
CSF is "lowpass"" (that is, there is no drop in sensltMty at low spati3l frequencies)
because the gratings were t emporally m odulated. (After Norcia, Ty ler, and Hamer,
1000.)
4 m onths o f life, and n ot u n ti l about 4 years of age is th e fovea fully ad ultl ike
(Yuodelis and Hen drickson, 1986).
From birth to beyond 4 year.s of age, cone d ensity increases in th e central
region, beca use of bo th the nligration o f recep tors an d decreases in their di-
1nens ions. Both of these factors resu lt in finer cone sampling (by d ecreasing
the d istance ben.veen ne igh boring cones). Al terations in con e sp acing and the
light-gathering p roperties of the cones during early d evelopment p robably
conh·ibute a g reat d eal toward the improvem en ts in acu ity and con trast sen-
si tiv ity durin g th e first m on ths of life. The massive migration of retinal cells, Sp,llial frequency (cyd es / de@\"e)
.m d the alterations in the size of retina a nd eyeba ll (along w ith changes in
inte rpu p ill a ry d is ta nce), may necessi tate the p L'lstici ty of corti cal coru1ections
early in life. [n teres tit1gl); the peripheral retina appears to de velop mud 1 m ore
rapidly th an th e fo vea (Yuoclelis and Hendrickson, 1986).
Sensation & Perception in Everyday Life
The Girl Who Almost Couldn't See Stripes

Nonna! visual develop-nent r9::1 uires nonnal \'1SU9J Abno nnal
early visual experience can have sa1ous and often conse-
quenc es for seeing patterns. as illustrated by the story of a gh1 named
Jane. Jane was b orn with a dense cataract (an opaplty of t he lens), In her
left gye, which prevented clear patterns from tOm\irg o n h<t left rerlna. In
ad dition to causing form deprivation In the lett gye, the cataract p-event-
00 J ane's two eyes from seein;J the same Images at the same time.
Stud ies in cats and m onkeys dating back to Hubel and Wiesel In
the early 19 60s have sho>'m that monocular form depr1vation can cause
massr,-e changes in cortical physiology that result In a d evastating and
permanent loss of spatial vision (Wiesel. 1982). Hubel and Wiesel. and
many other workers dem onstrated that there is a critic al
period of early visual development when no nnal binocular visual stimu-
lat ion is rEquired for normal oortical d evelopT1ent . In cats and monkeys
this critical pericd covers t11e l rst 3-4 m onths of life: in humans it is ex-
tended to something on the order of the first 3-B years. During the critl·
ral i::eri od , corti cal neurons are st ill b eing wired up to their inputs from
the two eyes . This is a pericd of neural plastlclty, wt e n abnormal visual
experience can alter the normal neural wiring process. If one eye is not
receiVlng ncrmal stimulation. the neurons that sho uld t:e destined to
respond to that eye do not be::::ome properly connected. In fact. sane
evidence suggests that these neurons are actual ty oo-o pte::t by inp..Jts
from the other, no nnally functio ning eye.
If cataracts are lett untreat ed d uring the Clitic al perio d , the mis-
placed oortic.al connect ions can never be repaired. The result is often c rlllcal period A phase In the life
amblyopia (re::1uced visual acuity In one eye b e::::ause of abno nnal early span durln;i which abrumal
visual experience- commonly known as "lazy eye") and an inabllity to experience can alter normaJ neuronal
perceive sterecpsis (a lack of blnowlar dept h perception : see Chapter development.
6). correcting Iha condition later In life will thus have little effec t . t:e - a mblyopia A developmental disor·
cause the lnfo nnatio n fro m the no·w-functioning eye can never be prop- der charactenzed by reduced spatial
erly conveyed to or processed by the cortex. vision In an otherwise healthy eye,
even with proper ccn ection for refrac-
tlve errcr. AJso known as/azy eye.

86 CHAPTER 3
strct>lsmus A mlsaJlgnment of the LLckily for Jane. her pediatrician found the cataract early, and the
two eyee such that a single object In cataractous lens was surgically replaced by an artificlal lens l/'lhen she
space Is Imaged on the fovea of one was 3 months old. The visual acuity in Jane's left eye just after the re-
eye and on a nonfoveal area of the placement lens was inserted was 20/ 1200. aboLrt four times worse than
other aye,
the no rmal value for a 3-month-old . But when tested ag ain 1 month
anlsometropia A oondttlm In which later, acuity in her left eye had already begun to catch up vvith the acuity
the two"""" have d ifferent refractive in her right "fe. In fact. a recent study of 28 infants (Maurer et al., 1999)
errcrs (e.g., me aye Is farsighted and found significant acuity improvements only an t1our after correcttve mea-
the other sures had been taken.
Not all individuals ·with o::>rQenltal cataracts are as lucky as Jane.
For example, in much of the third world , because of poverty, children
b orn w tth congenital catarac ts (often in both "f9S) go untreated and
grow up essentially blind. According t o the Wortd Health Organization ,
India Is home to the largest population of blird children in the wor1d.
to track how these "blind" chllcten learn to see. These studies are Just
be;Jlnning to provide important new Insights Into brain pl asticity.
Congenital cataracts are not the mly cause of amb lyopla. Eal1y
in life, t'wo other di sorders-strablsmus On vJhlch one f1'/0 Is turned so
that it is receiving a view of the wortd from an abnonmal angle) and an-
lsometropla un w hich the two eyes have very different refractive errors;
e. g. , one eye is farsighted and the other not)- may also cause amblyo-
pla. These fonns of amblycpla are typically less severe , and often they
have a later onset than congenital cataracts. The standard clinical treat-
m ent fo r aml:Jyop a, for over 250 years, 11as been to patch the good aye
and ''l crce" the ambtyopic eye to work. This treatment is ordinarily per-
formed only in children (typically younger than 8 years). However,
several recent studies suggest that there may be hope for recovery of
visio n in okJer children. and even in adults tliro ug h "perceptual learn-
ing" - repeatoct practice of a d emanding visual task (Levi and Li. 20091
or p laying action video games (Li et al. , 20 11).

SPATIAL VISION 87
Summary
1. In this cha pter we followed the path of image p rocessing from the eyeball
to the brain. Neurons in the cerebral cortex translate the array of activity Refer to the
signaled by retinal ganglio n cells into the beginnings of forms and patterns. Sensation and Perception
The primary visual cortex is organized into thousands of tiny comput- Companirn Website
ers, each responsible for determini ng the orienta tion, w idth, color, and
sltes.slnauer.comtwoWe4e
o ther characteri stics o f the s tripes in one small portion o f the visual field.
In Chapter 4 we will continue this story by seeing how other parts of the for activrues, essays, study
brain combine the outputs from these minicomputers to produce a coherent quesl ons, and othBr study aids.
representation .
2. Perha ps the mos t important feature of image processing is the remarkable
transformation o f info rmation from the circular recep tive fields o f retinal
ganglion cells to the elongated receptive fields of the cortex.
3. Cortical neurons are highly selective along a num ber of dimensions, includ-
ing stimulus orientation,. size, direction of motion, and eye of origin.
4. Neu rons with s imilar preferences are often arranged in columns in prim ary
visual cortex.
5. Seledi ve adaptation provides a powerful, n oninvasive tool for learning
about s timulus speci ficity in human vision .
6. The htunan visual cortex contains pattern an..'1.lyzers that are specific to spa-
tial frequ ency and orienta tion
7. Normal v isual d evelopment requires normal visual experience. Abno nnal
visual experience early in life can cause massive changes in cortical physiol-
ogy tha t result in a devastating and pemlal1ent loss of spa tial \.i sion.
8230336 Aml'la Appa

CHAPTER 4

Perceiving and
Recognizing Objects
WE HAVE BEEN TRAVELING UP THE VI SUAL SYSTEM from the eyes into the
brain. By the e nd of Chapte r 3, we had readied primary visua l cortex (Vl ,
s tri ate cortex), w here we encountered cells that '"'·ere optimally stimulated by
bars a nd g ratings of diffe rent o rlenrations. Of course, when you look at the
world, you d o not see an a rray of bars and g ratingsi you see co herent o bjects
and extended surfa ces. Moreover, you recognize speci fic o bjects even H they
are odd objects, as in Figure 4.1). C hapter 4 continues our journey through
the visua l system and considers how that visu<ll system manages the tas k of
perceiving and recognizing objects.
What and Where Pathways

To begin, let's extend the visual pathways beyond Vl and beyond w ha t was
shown in Fig ure3. 1. Recall that cells in \ '1 are interested in the basic featu res
of the visual image, responding to edges or lines of specific orientation, mo-
ti.on, s ize, and so forth. These neu rons have rela ti vely small and precise re-
ceptive fields . That is, a ce ll wil l respond to its p referred s timulus only if that
sti mulu s is presented in a very specific location relative to the p oint \'•there
the observer (monkey, cat, human) is fix..'l ting its (or his or her) gaze. Beyond
V1 is the extrastriate cortex, a set of visual areas so called they lie
just. outside the prirnary visuaJ (or s triate) cortex. ln the monkey, these areas
are nam ed V2, V3, and so on, th ough they· are n ot a simpl e chain of process-
ing areas and the naming convention breaks do\l\T\ pretty rapidly. Figure 4.2
shows the main visual areas of the m acaque m onkey brain, and Figure 4.3
sh°'"''S one w iring dia gram of these and a few o ther visu.:,J areas. Th e hum an
visual system is no t ide ntical to tha t of a macaque. H owever, the basic plan,
mapped out in Ftgure 4.4 is similar. Basic, local properties are pulled out o f
the image by early s tages of visual cortex, but sophis ticated tasks like object
recogni tion require a great d eal of s ubsequent processing and a large number
of apparently distinct visual processing areas. (See Web Essay 4 .1: Mapping
the Visual Corte><)
We can s ke tch a broad structure of the visual areas beyond VI. In the ex-
trastriate regions jus t beyond Vl (su ch as V2)r receptive fields begin to show
ru1 interest in properties th..i.t \Vill be important for object p ercepti on. Look a t
Figure 4.5. The boundary ins ide the red oval is the same in Figure 45a and
b. But to our eyes, Fig\u-e 45.1 shows the black edge of a black square while
Figure 45b sh ows the light e d ge of a li ght sq uare. A Vl cell tun ed for a dark
ed ge on the left side of its receptive field would respond equally in the two
extrastriate cortex The region of
situations. A V2 cell might not. V2 cells c..ue about 1'bol.md ary ownership." conex lx:fdeting the primary visual
They might respond to the dark edge of an object (Figure 4.Sa) but n ot to a cortex and containlrq mu ltlple areas
dark edge created by a light object on a dark background (Figu re 4.Sb) (Craft Involved In 1Asual processing.
8230336 Arnrna Appa

90 CHAPTER 4
et lt l.t 2fll1}· We have m ore to say about the receptive-He ld properties of

cells in ex trastriatecortex a fter we introduce the broad s tructure of the highe r
stages of the h uman v;sua l system.
From theextrustriate regions of the occipital lobe of the brain, vis ual informa-
tion moves out a long two m ain pathways (see Figure 4.2). One pathway heads
up into the parie tal lobe. Visual areas in this patlnvay seem to be important
for p rocessing information rela ting to the of objects in s pace and the
ac tions required to interact w ith them (movin g the hands, the eyes, and so
on). This pathway is some times knO\'\rn as the u>here pathway. As we \viH see
in C hapter 7, th.is pa thway p lays an important role in the deployment of at-
tention. The o ther pathway head s do"V.'Tl into the temporal lobe and is kno\'\>TI
as the what patlnvay. This p athway appears to be the locus for the explici t
acts of object recogniti o n tha t a re of particular imp ortance in this chapter
(U ngerlei der and Mishkin, 1982). As \o•le m ove d ovvn into the te mporal lobe,
receptive fie lds get much bigger. As the pathway's name implies, idrn/ is in
view seems m ore important than where it is. However, tho ugh it is a useful
organizing principle, one sho uld n ot becom e too addicted to this whnt an d
where distinction. For ins tance, some basic object information is represen ted
in bo th pathways {Konen and Kas b1er, 2008), a nd som e where information is
encoded in the tempora l lobe wha t pathway.
Early evidence for a relati onship bernreen the tempo ral lobe and object
recogniti on came fro m s tudi es in w h.id 1 large sections of the tempor.ctl lobe
FIG URE 4 . 1 We see a world full o f were dest royed (lesioned) in m onkeys. When Khiver and Bucy (1938, 1939)
Identifiable o bjects ewn if we have dJd this, they fo und that their monkeys behaved as thoug h they could see
nevier seen them in this partbular but did not know ¥•.' ha t they were seeing. This deficit, also seen in som e hu-
airangelnQf'lt bQfore.
FIG URE 4.2 The m ain \'i sual areas o f tt1e macaque m onkey cortex. Humans have
comparable visual areas (569 Figura 3. 1). Tha d rawing o f the brain has b99n d is to rted
to show ar'*'6 that lie deep In the folds (sulCI) o f tha cortex. Each abbreviatio n refers
to a. different visual area, not all o f which w ill be discussed here. Visual cortical i::ro-
lesion In referenoe to roorophyslol- cesslng can be divided into two broad stn'.lams. O ne, heading for the parietal lcbe,
ogy, 1. (n) A region of damaged brain. can be thought of as being lntbfested in whGra things are . The other, heading down
2. M To destroy a sectlon of the brain. into the t€1mporal lo be, is concarned with wMtthings are. (After Par\<Qr, 2007 .)

PERCEl\llNG AND RECOGNIZING OBJECTS 91
FIGURE 4.3 A partial dla-

grarn for the main \iis.ual areas of the
braJn . As in 4 .2, each abbrevia-
tion to a diffQt"Mt visual area
not all of which will be discussed here.
The main messag9 o f this figure is that
the visual pathways are w:,ry complex
and that the.re are a multitude o f visual
areas. For a bit of oriGntation. v is.Lia!
inforrnatio n from the eygs enters at the
bottom of the diagram. via the retinal
cells (RGC). Feed-fo rward
informatio n generalty flovvs · upwatd•
in this figure. HOWQVer, visual procr0;ss-
ing is a two-way street. From the LGN
orrNard, there are b oth feed -forward
and feedb..'\Ck connections between
areas (seei Figure 3. 1). (Aft.:f Felleman
and V an Ess9n, 1 QQ 1 .)
36 AmMa Appa
'-"M'-'--'--' RGC
FIGURE 4.4 The main visual areas of

the human cortex (compare \11.'ith Figure
4 .2 and Figurg 3. 1). the convo-
luted, wrinkled surface of the brain has
been flattened. Each abbreviation refers
to a different v isual area 01 the web,
you can find the definitions o f all the
acrcnyms and s9'V9ral oth9r maps that
w ill give you an idea o f the cornplexlties
o f figuring out the map of human visual
processing. H9re you should no tice
the what and pathways, and the
areas specialized fa fac es, b odi9S, and
scenes. (Courtesy of Sabine Kastrigr
and Mike Arcaro.)

92 CHAPTER 4
(a) (b)
FIG URE 4. 5 Notice that the reoeptive fl91d (the roo ovaQ sees the sarnei dalkedge
in both images. Howewr, a C911 In V2 that pref&rn dark edges might respond more
strongly to the one in (8), where the little square CM'ns the dark edge, compared with
the case in where the real edge Is the edgB o f a light square, above and to the
right o f the rec19ptive
man stroke victims, is knmvn as agnosla, though Kl\h"er and Bucy called it
" psych ic blindness.'' Later work pointed to one part of the temporal lobe, the
lnferotemporal (IT) cortex, as particttlarly important in the visual problems of
these monkeys. In the 1970s, Cha rlie Gross and his colleag ue.s began to record
the activity of si ng le cells in this area. \ Vhat they found was quite striking,
We have seen that n eurons in striate cortex a re a di vated by simple sti muli
and respond only if their preferred s timuli are presented in very restricted
portions of the visual field . In contras t, cells in the IT cortex were d iscovered
to have receptive field s that could spread over half or more of the m onkey's
field of vievo.·. Even more s triking were the sorts o f stimuli that activated IT
ce lls. The usmd spots a nd lines didn't work well at aU, but the silh o uette of a
monkey h and worked for some cells. f\.'lonkey faces excited o ther
cells (Figure 4.6 ). There was even a cell with a distinct p referen ce fo1 a toilet
brush shape (Gro.ss, Rocho-Mirand a, and Bend er, 1972).
Findings like this led Horace Barlow (1972) and others to s uggest a hier-
arch ical model o f visual perception in whid1 the sm all receptive fields and
simple feahtres of visual cortex are combined w ith ever greate r complexity
as one moves from striate cor tex to IT cor tex, even tua.lly culminating in a cell
tha t might fire w hen you see your gnmdmother or your Sensation & Percep--
tim1 tex tbook. Indeed , the tem1 grandmother cdl, coined by Jerry Lettvin, has
entered the jargon of the field to stand for any cell that seems to be selectively
respon sive to on e specific object (Ba rlow, 1995).
\Ve d o not know \o\>Th'lt brnin activity achlally corresponds to your recognition
of your gran d1nother. It is probably a bit extreme to imagine that this is the
work of a single neuron in the lT cortex, and more recen t work has cas t d oubt
on the idea tha t IT cells respond to an object independent of its position in
the visual field (DiCa rloa nd Maunsell, 2003). Nevertheless, it does seem clear
tha t cells in tha t part of the brain are critically involved in object recognition.
a'1losia A fallure to recCQntze 1l1e IT cortex maintains d ose connections w ith p arts of the brain in volved
ctJJects In spite of the ability to see
them. Agnosia Is due to brain in mem ory fomlation (not.''1.bly the hipp ocampus). This is important because
damage. those IT cells need to learn th eir receptive-field properties. The recep ti ve-field
properties of primary visual cortex could be \'\-Titten into the genetic code in
lnferoternporal (11) cortex Part of
the cerebral cortex In the IC1'ver por- some manner, but n e urons that respond to g randmothers dearly cannot be
11on of the temporal lclle, Important In hardwired, since everyon e's grand m othe r is different. Nikos Logothetis and
ctJJect reoognltion. his coworkers d em onstrated that cell s in IT cortex ha ve p recise ly this type

PERCEIVING AND RECOGNIZING OBJECTS 93
FIGURE 4.6 Cells in the lnferotemp:>ral cortex of

macaque monk9ys ar9 interested in very specific stim-
uli. In this case, too cell responds vigorously to a rnoo-
ksy faca and to som e othgr stimu fi that seQITI related.
11111111111111111 (After Gross. Rocha-Miranda. and Bender, 1072.)
11111 111111111111111111111
11 1 11 111 11111111111
© .
i
1111111111 111 1111111 1111 11 11 Ill
I I 11111 11111 1111 I
of plasticity (Logothetis, Pauls, and Poggio, 1995). A fter training monkeys to

recognize novel objects, these researchers found IT neurons that responded
with high firing rates to those objects, hut only w hen the objects were seen
from viewpoints similar to those from which they had been leanled.
Human cortex has areas that appear to be the equivalent of monkey IT cortex
(the anatomy of human and macaque monkey brains is not identical, so we
talk about homologous regions). One of the more amazing demonstrations
of this fact comes from a 2())5 study by Quiroga et al. They made recordings
from single cells in the temporal lobe of human observers. Normally we do not
put electrodes into the human hrain, but these observers were patients being homologous regions Brain regions
prepared for brain surgery to treat epilepsy. Implanting electrodes was part of that appear to have the same furci.bn
the treatment plan, and recording visual responses' from these cells involved In different species.

94 CHAPTER 4
Jenni:kr Anlston
JerutiferAn.i.s ton Jennifer A.ni.ston and Brad Pitt l..alU'a Linney
Stimulus
presented
. . lJAL
' ;
I
I
!
I
I
!
lLL
' ; LlL
';
I
I
! !
!
I
I
!
I
I
I
I
Ce ll's
response
Spider Sydney 01--.er,1 Hou se Leaning Tower
8230336 Ann Stimulus
u·,
presented
.,
LlLU
i i i
Lil
',
i i
I I I I
i
I I
Cell' s
response
t
Stimulus Stimulus
ON OFF
FIGURE 4. 7 R9sults of rgcording the activtty of on.e 001 in the kimporal lob9 of a
human patient. This oell responded to pictures of the actress JEf'lnifer Alliston. The
c:eill did not respond to pictures o f anything 91se, including thoS9 o f o tt"lQ'" actr9SSQS.
CAfti;.r Quiroga et aJ ., 2005.)
no extra risk to the patient or interference \Vith treatment. In the experiment,

the observer just looked at a collection of im ages \lvhile the activity of a cell
was monitored . As Figure 4.7 sh ows, like the cells that G ross h ad found in
monkey IT cortex, some of these cells turned out to have very specific tastes.
The cell sh O\vn in the fi gure responded only to th e actress Jennifer Aniston
and to nothing else presented to the obsen 1er. Other cells had preferen ces for
other people, like former Presid ent Bill C linto n. One ce ll responded to the
Sydney Opera Ho use; an othe r, to the Eiffel Tower and the l ea ning Tower of
Pisa but not to other londmar ks (Quiroga et al., 2005).
\Vhile we d o n ot have a lot of systematic d a ta on the responses of single
cells in the human \oisual system , we have a growing volume of fun cti onal
imagin g data that d ocuments areas in the huma n brain th a t appear be
specialized for different sorts of stimuli . Conveniently, as s how n in Figure
4.4, ma ny o f these h ave been given names tha t ma ke th at proposed spe-
cialization dear. Thus, cells in the fu siform face area (FFA ) a re interested
in faces (Kanwis her, McDerm o tt, and C hun , 1997) while an area like the
parahippocampal place a rea (PPA ) has cells that resp o nd to s paces in the
\Vorld like roo ms wi th furniture in them (Eps tein and Ka nw isher, 1998).
T1'1e visual system has m an y differen t problems to solve-like the problem
of face proces.sing-and it appears to have m odules that are specialized for
worki ng on different problem s (Kanwisher and Dilks, 2013). Other e\o;dence

PERCEl\t1 NG AND RECOGNIZING OBJECTS 95
com es from brain lesions ca used by s tro kes o r o the r accid en ts. Like K.li.iver process A process
and Bucy 's m onkeys, humans w ith lesio ns in the tempo ral lo be often s how that carries out a computation (e.g ..
symptom s of agn osi';:i,, the ability with o ut the ability to know w ha t is cb)ect reccgnltlon) one neural step
after another 1 without need fa feed-
be ing seen. Som etimes these a gn osias can be quite s p eci fi c. Prosop agno.sia, back frcm a later stage to an eanler
an inability to recognize faces, is one exa mple that h as receive d a lo t of study stage.
and that w ill be disc us.sed at the end of thi s chapter (Dam asio, Oa masio, and
Va n H oesen, 1982). There are other interes ting subdivi s ions of agnosia as
welt s uch as the ability to recognize anirnate objects (e.g., animals) but not
inanimate objects (e.g., tools) (Newcombe ilnd de Ha:m , 1994 ). The implica-
ti ons of specific agnosi:as in s p ecific pa tients sh ould no t be taken too far.
While 3reas s pecia lized for faces o r place.s seern to be readily d ocum ented
in humans and you mig ht have a reas specialize d for some o ther ca tegories
of objects, like tools (Hutchison e t a l., 2014), you p roba bly d o n ot have a
separa te area for each and e-very ca tegor y o f object tha t yo u can recognize
(Kan wisher and Dilks, 2013).
Some p rocessing that lead s to the catego ri zation of objects and scenes can
be very fas t. Electri cal ac ti vity from the brain can be recorde d froin e lectrodes
placed on the scalp. If we flash a p ic ture to an observer and a s k whether it
contains a n animal, we ca n record a s ig na l in the observer that reliably dif-
fere ntia tes an im al from nonanimal scenes w ithin 150 milli seconds (ms) from
the onset of the s timulus Fize, and tvlar:lot, 1996). That's fas t en ough
to mean that there cannot be a lo t of feed back fr om higher visual or m emory
processes, suggesting that it must be possible to do sorne rou gh object recogni-
tion on the basis of the first wave of ac tivity as it m oves, cell by cel l, synapse
by syna pse, from retina to stria te cortex to extras tri::.lte cortex and beyond .
Tha t feed-forward process mus t be able to gen erate an "an.im a1" s ignal from
a w ide rro, ge of animals in different positions, s izes, and so on (Serre, Oliva,
ond Poggio, 2007).
To summari ze, two path,-vaysernerge from visual cortex. 11,ewlierepathway
"vill be taken up in late r chapters. The -whnt pathway m oves through a s uc- 0336 AMmJ App.l
cession of stages, building a rep resen ta tion of your g randm othe r or the Eiffel
Tower out of the very .specific, very locali zed s pots, lines, and ba rs that interest
the cells in the retina, lateral geniculate nucleus (LG N), and primary v isual
cortex. That' s a but d on 't be lu1led into thinking we fully unders tand
the p rocess of object recognition si mply because we ha ve som e n oti on of the
ne tual pa th ways involved. It is a really difficult p roblem, and in the rest of
the ch apter we will illus trate wh y this problem is ha rd and how the visu al
system. attempts to sol ...-e different parts of it.
The Problems of Perceiving

and Recognizing Objects
Figure 4.8a is d ea rly a picture of a ho use. The imag e may contain som e o ther
things as well, but the chief object of in teres t is the house. It is p retty clear that
Figure 4.8b also sJ, ows a h ouse, perhaps with a s imilar shape a nd chh1u,ey,
though in this case the h ouse is pa rt of a m ore abs tract scen e painted by the
Frend ' artis t Georges Braque in the early twentieth century. Figures
a re two more pictures of h ouses. It is quite d ear tha t Figure 4..Bc is the sam e
ho use as Figure 4.811, w hile Fig ure 4.Sd is not. These seemingl y s imple acts
of object identification require a lot from the wJrnt pathway, and they cons ti-
tute the main topic of the res t of this chapter. Like m any seemin gly s imple
acts, object percepti on is actuall y a collecti on of complex and remarkable
accompli shments.

96 CHAPTER 4
FIGURE 4.8 The problem of object

racognition . ThQ house and the oi l
painting by Goorges Braque in f/:J} look
dlffQ!"&nt, but they are both Consid er Fig ure 4.9a. It s hows yet another h ouse. How do we recognize
c lOOTIY houses. {c) This irnage is also it as a house? First "'ie nee...1 to gather some bclSic visual features. The preced-
vety different from the lmag13 in (a). but ing chapters showed us how single cells in the early visua l system respond
they ae both cl90.t1y th9 $i.llTl9 houSQ. to stimuli suc h as simple lines. In Figure 4.9b each circle is a cartoon of the
(d) This image of a red h ouse cleart-,t
receptive field of a sllnple corticn l cell. Those cells in striate cortex w ill respond
shows a hoUS9 that is diffSfent from
(8) and (C) .
well to the high-contrast lines in the outline of the house. Th(lt limited recep-
ti ve field is like a w indow that allows the cell to "see'' only a small part of the
world . None of these simple cells see a house. Tl,ey jus t collect local feahtres
like horizo nta l, vertical, and oblique lines.
At the very least, the local fea tures''\\dll a house.
1his p rocess could be frnagined as the natural extension of a process \Ve have
already discussed. One way to 11 construct" the lines detected by simp le cells
(a)
ODDD CD
DD
FIGURE 4.9 The prot:Xem continued. (a) Another house. (b) Cells in primary visual
cortQX raspcnd well to the local fQ.ilturoo (circ led} of the housQ. But how do we go
from r.accgnizing a collection of local features t o perceMng a house? (c) In this sligt1tty
m ore oornplicatad sCGr10, how do we know ..,.,hich bits belong t ogeth19r?

is to imagine combining a row of d ots det cteab_y gapglioITcellSi. Dots middle (midevel) vision A
could be grouped together into lines. A pair of lines, each collected by a dif- defined stage of ,;sua1 prooesslng tmt
ferent cell, could be combined by another ceU tha t would sense a This comes after basic features have been
extracted from the Image cr
process could go on a nd o n tmtil we had a h ouse. On closer examination, \'lsloril and before dJject recog-
however, it is dear that this process would not be easy. Consider, for example, nition ard scene un:Jeretandlri;i (high-
Figure 4.9c. Agai.n, sitnple cells would luwe no problem detecting the lines level vlslonl.
and edges in this scene, bu t how do \-Ve know which edges go \o\ith w hich
objects? How d o we avoid considering the s now man part of the h ouse? The
h ou.se a nd s n owman m ake comers, as d o the h ouse and its d oor a nd s teps.
The car and ho u se a re separa te. None of thei r lines toudl . But this is also true
of the outline o f the ho use and its \o\-indo\.vs.
C learly we must have processes that s uccessfull y combin e features into
objects. That is one of the tBsks that defines middle (or mldlevel) vision (as
opposed to low-level vision, whidl was the topic of Chapters 2 and 3). The
next part of this cha pter looks at sorne of the processes of middle vision. 11-ie
followin g part takes the fea ture combin a tions given to us by middJ e-level vi-
sion and asks h ow we come to know w h.:1t th e object is. The act of recogniti on
must involve matching \vha t we perceive now toa memory of somethin g we
perceived in the past. H ow ca n we do that? For exam ple, how d o we knmv
that all of the images in Figures4.8and 4.9show houses? Witho ut having seen
these exact objects before, how d o we go about placing them in the "house"
category? Furthermore, h ow d o we know tha t Figu re 4.& and c portray the
same house, g iven tha t the images de livered to our retin as in the n"-o figures
are radicaJly different? ln the fi na l section of this chapter we..-U disc uss the
high-level visual processes that enable us to recognize familiar objects, nov·el
views of familiar objects, and new instances of farniliar object ca tegories.
Middle Vision
The goa l of middle vision is to orgartlze the elernents of a visual scene into
groups that we can then recognize as objects. Let's begin with the .simples t
case of an object isolated on a simple background. Finding the edge,s of this
object w ill be a good starting place on the road to recogn izin g the object.
Rnding Edges
In Chapter 3 we discussed a t length how n eurons in striate cortex can detect
bits of lines, but how d o '"'·e decide which bits belong to wh ich objects? We
al ready established that '"'e can' t just g roup a ll the edges tha t touch each
other into an object. Because objects a but and overlap other objects, simple
connected ness will n o t work . Worse yet, before we can concern ourselves
with grouping edges, we n eed to worry about the quality of the raw edge
in fo rma tion. In Figure 4 .10, part of the house of Fig ure 4.9 is red u ced to a
simple, arrow-shaped outline. It is easy to see th at arrow. Notice, h owever,
that in som e places the object is lighter than the background, while in other
places it is darker. This means that if we trace the e dge of the object \vi th a
finger, we m11st pass through locations w here there is n o differen ce between
the luminanc-e of the object and the h uninance of the background. In other
words, a t these points the shape has n o edge a t all.
FIGURE 4 .10 In some plaoes this ob)act is darker than the background. In o th.:4"
plaoes it Is lighter. If the c hanges an:i oontinuous, it follows that th9f9 must b9 plac9S
'Wher9 the edge of this shape simply disappears, even if you see 9dgee as
continuous.

98 CHAPTER 4
(a) (b) (c)
FIGURE 4. 11 (a) The Mfind Qdges" function in a popular graphk:::s program finds
gaps in tt1e bordB""s of the image in Agure 4 . 10 - gaps that we do not see. (b) Whffi
a o::>mputB"" algorithm lool<s for the edges in this elephant scene, it finds Wiat Is
shown In (c),
Illusory contour A contour that Lnterestingly, this oo:asional lack of an edge d oesn' t seem to bother our visual
Is percelvOO 6""n though rothlng .system a t all In fact, it m ay be hard to see the gap. Asking sirnpl e computer
c hanges from cne side o f It to the grnphics sofhvare to find the edges in Figure 4.10 wo uld yield som ethin g like
o ther In an Image.
Fagure 4.11a . 11'\e visual system kno\vs tha t the gaps are acd den ts of the lig hting
structuralism A school ofth0tight and fills in the contour. 111e computer is n ot so clever. Fig ures 4.11 b,c s how
belle>Ang that complex objects or
perceptions could be understoo:J by
an other example. In Figure 4..1 lc, no tire that the ed ge-finding software find s all
analysis of the components. sorts of edges. The human visual system , however, is d oing some thing quite
In the early s tages of processing, it is figuring out w hich ed ges m ark
the bound aries of objects a nd w h ich represent s urfa ce fea hues (like cracks on
the rock faces). All these differen t bits of information are then combin ed to
ma ke the syste m's best g uess about the presence of a contour.
ll-1e inferential nature of con tom perception ca n be nppreciated in the m ore
extreme demonstration sho""TI in R g ure 4.12. lhis is an example of a "Kanizsa
figure"-named after Gaetan o Kanizsa (1913-1 993t a n Ita lian psychologist
who spent m any years investigating s uch stimuli. Here it is still e1sy to see
the arrow outline, even th ough the vast majority of the shape's lines are m iss-
ing'! Check-it yourself. There really is n o border between the white fig u re and
the white background . These edges, called Illusory contours, are perceived
because they are the best guess abou t w hat is happening in the world a t that
loca tion. It rea lly does seem likely that a con tom is present, even if there is no
physica.I evidem:.-e that location.
RULES OF EVIDENCE l et's consid er the sorts of regularities in the s timulus

tha t are bk.en as evidence for a contour in the world , be it a real conto ur or
a good guess like those made in Figu re 4J2. TI1is tendency of the visua l sys-
tem to make inferential leaps was problenrnti.c for on e of the ea rliest g ro ups
of perceptual the structuralists. Stn1ctma lists sud1 as Wilhe lm
FIGURE 4.1 2 This .. houS9" outline is constructed from illusory contours . Even
though the contour is doo11y visible , there is no piys1ca1 differsnce betw.ae.n the w hite
background and the vvtli\Q house.

PERCEl\t1NG AN D RECOGNIZING OBJECTS 99
Wundt (1832-1 920) and Edward Bradford Titchener (1867-1927) argued that Gestalt In German.
perceptions a re the s um of atoms of sensatio n-bi ts of colo r, orienta tion 1 and In refe re nce to perception, a schocil
so forth . In the structuralist view, perception is built up of local sensa tio ns of thought stressing that the percep-
tual whole could be greate r than the
the way a crysta l might be built up of an arrny of atoms. A n i1lusory conto ur apparent sum of the parts.
challe nges th is view because an extend ed edge is seen brid g ing a gap wh ere
no local atom of "edgen ess" can be fo1.md. Gestalt grouping rules A set of
rules describing wrrlch elements In an
Over time, it became clear that there are many examples w here the s truc- Image will appear to group togetl1€f .
turalist a rgmnent seems to fail. Inspired by th ese exampl es, a second group of The origh1al list was assembled by
psychologists, led by Max Wertheimer (1880-1943), \ Volfgang Kohler (1887-1 967), members o f the Gestatt school of
and Kurt Koffka (1886-1941), fo rmed the Gestalt school (Wagemans e t a l., thought.
2012). Gestalt theory he ld tha t the perceptual w hole is more than the s urn of
its sensory parts. Perhaps the most enduring contributio n of this school was
to begin the description o f a set of organizin g principles (some times known as
Gestalt grouping rules) tha t d escribe the visual syste m 's interpretation of the
raw retin al image. In the next sections, we will d escribe some of those rules.
More than the specific rules, it is important to remember the overarching goal.
The vis ual syste m is trying to make sense of the vast and often a mbiguous and
noisy inputs from the early st.'lge of visua l processing. These rules are useful
parts of that effort beca u se they reflect regula.rities in the world. l11ey allo\v
the visu.'.ll system to So:'l)i "H the input looks like tha t, 1 ca n infer that this is the
state of the visual '"'·odd."
With tha t in ntlnd, consid er Figure 4 .13a, w he re we h::we a collection of
short line segments. O ne se t of these seen'\S to form a contour, an irregular,
dented loop. The " rule,'' cartooned in Fig ure 4.13b, is th a t we ten d to see
simiJarly oriented lines as part of the same contour (D. J. Field, Hayes, and
Hess, 1992). Pola t and Sagi (19Q3) measured how pieces of a contour s upport
each o ther. The effect w ill be hard to see in print, but the fainter lines in the
(a) (b)
\Vhkh gray line is a likely
oonlinuation of the black line?
Not l: _\/ /y-------'

f'IGU R E 4.1 3 Contour completion.
/ (a) The roughly c irc ular contour saen in

th9 cent9r o 1 this diagram r9sults from
the visual system's applic:ttlon o f the
(c) (d) rule shown in (p). (c) Polat and Sagi
{1993) found that it was QaSiQr to SQQ
Very likely a faJr1t set o f bars if they were flanked
by bars of the sarll9 oriantatbn.
W. S. Getsler and J. S. Perry actuall'/
measlJred the relationships beitween
8230336 qeart?y fine segments In natural SCQOQS.
Tho figurJ shows the probability that a
horizontal tine SQQn')Qnt 'WOUid CO'"()CCur
with othGr line S9Qm9flts in the Image
at different distances and orientations.
ill9 most hK91y pars are ooM n9lr or
nearly so. (Part c aftQf Polat and Sagi.
lQOO; part dafter W. S. Geisler and
Very unlikely P'1fty. 2009.)

100 CHAPTER 4
x
(•) This... (d) Btit this ..
DJ
x
(I>) .. looks more like this .. (e) .. looks less like this ..
DJ
x
(r) .. . not like this . (j) .. .thanlike this .
FIGURE 4.14 The Gestalt princip les

o f goocl continuatiOn and closure:
DJ
Undt< the Gestalt princ iple of good midd le of the trip lets of lines in Ftgu re 4.13c will be easier to see w hen they
continuation, pr·e99nted with an image
are collin ear \vi th the flankin g lin es. ll1ose flankers provide evidence for lines
like the one in '8). th9 brain tends to
int9rpret It as a pair of intersecting
of the sam e orientation in be tween. If a set of lines fo rms a closed s hape like
lines. as in (b), rather than reaching any the roughlycircu1u con tom in Figure4,13n, then the sh ort segments support
o ther conclusion, suc h as that in (c). ead1 o ther even m ore s tron gly (Kovacs and Julesz, 1993). Geisler and Perry
But th9 principle o f c losLira can trump (2009) d ocumented the regtJ arity in the world that s uppo rts the rule shmvn
good continuation. as shovvn in (d-1). in Figure 4.13b. They labeled many, man y contours in na hmd scenes. Then
ig) Laura Williams c leverly exploits good
continuation by lining up the edge of they examined pairs of contour pieces and asked , 11 What is the cha nce th at
tM fiek::l w ith the reflected E!dge In the tliis piece lrere at this orienta ti on is o f the sam e contour as that piece tire re
mirror. lhis allows the good oontinu- at thaf orienk1tion ?'' TI1ese like lihoods are colo r-coded in R g ure 4.13d, and
atlon cut1 to trump the ocduslon cue they s h ow that in the real world, if h¥o contour elements are dose to co linear,
(S99 next section) to produCQ this strik-
they are likely to come from the sam e contour. Sharp turns are much rarer.
ing image.
11,eGestaltists called this the priru:iple of good continuation (Figure 4.14),
and they illustra ted it \Vith examples like the one sh O\-vn in Figure 4.14a. \'Ve
tend to see this figure a pair of intersecting lines (Fig ure 4.14b). There are
many other p ossible orga nizations (Figtue 4.14.c, for example), bu t all else be-
ins eqrml, the tenden cy to see lines continuing in the sam e direction-in other
words, the tenden cy to group edges that have the sam e orientation-supports
the X-like interpretation ill us trated in Figure 4.13b.
PERCEPTUAL "COMMITIEES" TI1e 1' all else being equar' phrase is imp or-
tant. As the Gesta lt psychologists knew, and as we \Vill see, a host o f rules,.
principles, and good guesses contribute to our organized perception of the
world. These seem to operate according to a sort of committee model Every-
one gets together and voices o pinions about how the s timulus ou gh t to be
good continuation A Gestalt unders tood . Fo r example, look a t Figme 4.14d. \'\Then we make the contours
grouping rule stating that two <>ements
,,,;11 tend to group together If they seem
of Figure 4.1411 part of closed forms, good continuation is at odds with the
to Ila on the same contour. principle of closure. Now, the sam e intersecting lin es are interpreted differ-
ently. In this case clos ure is stronger than goo d. continua tion. 11-ie s upremacy
clostre In reference to perception,
closure Is the name of a Gestalt pnn- of closure over good continuation is not absolute. \ Vhen the opini ons of differ-
ciple that holds that a closecl contour ent 11comi:nittee m e1nbers" the results can be somewhat unpredictable.
Is preferrecl to an cpen contour. Somehow, though, a consensus vielv a lmost a lways quickly emerges and we

(a) (b) FIGURE 4. 15 The making o f illusory

contours. The little arrCJWS In fJ:>) and
(f:I} repr9Seot the visual system's best
guess about what is going on in {a) and
(c). The illusory disk in (c) aris4:ls when
the v isuru system combines a whole
collec tion of guasses about the line
rninations . as shown in {d).
(c) (d)
'}>..1.•.4'(
" /
.-
'"
-: '
/
" ;-.(
settle on a sing le of the visua l scene. TI1is committee me ta phor
is \·ery useful and 'vill recur through out this chapter.
OCCLUSION Lf an e d ge suddenly s tops in an image, tol1y does tha t e d ge texture segmentation Carving an
s top ? One reasonable gu ess mi ght be tha t it s tops because som e thing else Image lnto regions of oomrnon texture
properties.
gets in the wa}i hiding it from our vievv. So, returning to the Kanizs.a figure
of Figure 4.12, we can the visual sys tem asking w hy the vertic.J.l line
at the bottom of the figure suddenly s tops. TI1e answer tha t the visual syste m
seems to come up v..rith is tha t there is a nother conto ur occl11di11s the vertical
line, with the ocduding edge oriented perpendicularly to the occluded edge.
llUsgue.ss, combined with a guess that the notches in the circles represent
contours tha t can be extended , lead s to the inference of an illusory contour
(Fig ure 4 .1 Sa,b ). (Note that we also interpret the notches as places where one
object is occluding another.) Rgure 4 .15c sh mvs another example. II eadl black
line genera ted weak illusory contours at right an gles to its e ndpoint, then this
figure would contain a set of ro ughly coli.near line segmen ts (Figure 4.1 5d).
Those segments could be grouped toge the r to form a con vincing ci rcle in the
\vay that the line segmen ts formed a loop in Figure 4,13a. (By the way, illusory
con tom s li ke this rnake exce llent doodles when you would rathe r be d oing
informal p erception exp erim e nts than paying attention.)
Texture Segmentation and Grouping

Connecting s hort line segments w ill get u s only so far in di";ding the raw im-
age into objects. On the left in Figure4.16, you can find a region that differs in
the size o f its e lem ents. An edge detector like the one described. in Figure 4.5
will be obliv ious to the difference beh.veen the large r-grain and s malle r-grain
textures. lnstead, the regio n is found by the visual system' s sophistica ted
mechanisms for texture segmentation (Beck, 1982; Beri;en and Adelson, 1988;
Malik and erona, 19'/0/- Qn the r;iglit-hand side o f Figure 4.16, the texture of
the water looks different from the t ture of the stone surround ing it. Here,
ra thel' th."ln
. deciding w hich feahires go together, the visual system m ay look
FIG URE 4. 16 The problem of texture segmentation. How do we distinguish the

two halves o fth9 11gure? How do we Si9gment out the smaller regions inside each
halt? On the bottom, hO'W might we know that the large area is stone, surrounding a
smaller area of 'Water?

102 CHAPTER 4
FIGURE4.17 ltis easy tos99 the

approximate average orientation of the
tines in the two regions of this figure.
The red lines arie vertical ori average
and the blue lines, horizontal. How9V€<.
you w ill n91iJd to take m orG trouble to
figure out whather there is an indMduaJ
red lina that is perlectty vertical or an
individual blue tine that is pg-fectty hori-
\ \I\ I ----
1"\ I \I\ 11
---
- --
I \ I I \ - ----
-- ...._
- -
zontal . Apparientlyyou can figure out
the awrage o ri4mtation vvithout b€1ing
a...vare of the orientation o f each com-
ponent limi.
...._
at the s tatistics of a.II the features in on e region and find those s tatistics to
be different fro m the statistics in an other region (Alvarez, 2011; C hon g and
Treisman, 2003). Indeed, the visuaJ system ca n d etermine the average of the
fea tures in a region vdth out knowing tnuch about the individual fea tures . In
Fig ure 4.17, for exa mple, it is immediately obvious that the ave rage of the red
lines is near vertica l, while the blue lines are near h orizontal. H owever, you
will have to search to detemline ""·hether a ny individual red lin e is perfectly
vertical or any individual blue line is perfect ly horizonta l.
slmilartty A Gestalt grouping rule Texture segmentation is closely related to the Gesta lt grouping principles that
stating that the tendency of two fea- \.ve 1ve been d iscu ssing . Figure 4.18a illus tr<1 tes two of the stron gest principles:
tures to group toget11er will k1crease as similarity and proximi ty. SlmUarity means that in1age chLmks that are similar
the slmllarlty betwe€(1 them Increases.
to each other w ill be more li kely to gro up togetheL Similar in w ha t ways?
proximity A Gestalt grouping rule Texture grouping can be based on simila rity in a linli ted number of fea hues:
stating that the tendency of two fea-
color, size, orien ta ti on, and, as illus trated here, aspects o f for m. Combinations
tures to group together will Increase as
the distance between them decreases. e'conjunctions" ) of features d o n ot work well (A. Treisman, 1986b). Thus, in
Rg ure 4.18b the texture segmentati on is n ot d ear, even thoug h the left sid e
parallelism A rule for ftgure-ground
assignment stating that parallel con- contains oran ge dia monds an d green squares and the right side contains green
tours are lfkel'/ to to the same dk1Jnonds and orange squares.
figure. 1l1e principle of proximity holds that items near each o ther are more likely
symmetry A rule for figure-grcum to group togeth er tha.n are items m ore widely separated. Proximity grouping
assignment stating that symmetrlcal gives Figure 4.1& its horizonta lly striped c1ppearance. The origina l descriptions
regbns are more likely to be seen as of these Gestalt principles did n ot involve extensive experim entation. Illustra-
fig ure. ti ve demonstrations, like those of Figure 4.18, seemed to rnake the point Since
tha t early Gestalt work, ca reful expe rimen ts have confirn1ed and quantified
what the early demonstrations illustrated . For example, Kubovy and his col-
leagu es (Kub ovy and Cohen, 2001) qui te p recisely measured the strength of
the proxi mi ty e ffect.
Figure 4.19 illustrates two somewhat weaker grouping principles: paral-
lelism and symmetry. Most people see the parallel pair of contou rs (2 and 3) as
a g ro up and th e symmetricnl pair (7 and 8) as roup .p.nd Ag ur.e 4.20
(a) (b)
****** 000000 o <>o 0 0 00 00 0

o o<>o 0 00 00 00
FIGURE 4.1 8 Similarity and r::<oxim- ****** 000000 00 00 00 0 0 0 00 0
****** 000000 OOOOOOOOOOOO
ity. i;a) Grouping by similarity an::! by
proximity is useful. (p) G rouping by a
conjunc tion of color and fotm does not
00 00 00 0 00 00 0
work ****** 000000 o o <>o <>o oOo <>o<>

FIG URE 4.19 Parall91ism and

symme.try. Which pairs of lines g:,
ti;:>geth1;;ir?
sh o\>\"S the grouping effects of enclosin g o r connectin g ite ms (S. E. Palmer,

1992). We can ' t list all the rules (even if \.veknew them all, w hich we do no t) .
More important, for our purposes, is to remember tha t the visual system 's goa1
is to decide which bits o f the input belong \'Yi.th w hich other bits of the input.
, ..........
See Web Activity 4 .1: Gestalt Grouping Principles fo r exa mples of
grouping by common fate and synchrony-two principles not covered here 3 • .-..-..-. .-. •
(S.-H. Lee and Blake, 1999).
FIG URE 4.20 Proximity grouping (li ne 1)
can be overruled by grouping by comm on
CAMOUFLAGE The S.."lme principles that are n ommlly used to help u.s find region (line 2) or grouping by oonnected-
objects in the world can be exp loited to hide them. The art of cam oufl age is, ne6s 3). (From S. E. Palmer, 19Q2.)
to a great extent, the art of getting your features to group '"ri th the features of
the environment so as to persuade an observer tha t yo ur feah.rres d o no t fom1
a group of their O\o\iTI , Figure 4.21 shows som e examples from the
animal kin gd om. The exercise fo r you i s to de temiine which principles of
(c) (d)
FIG URE 4.21 Panel fP) shOlNS the

hiddgn animal in (a). We leaw (c) as an
SX(("Cise ior the reader. What Gestalt
pindp{es are at work here? (d) Som e-
tllTIQS camouflage Is meant to confuse the
\119W'eor. rather than to hide the object.

104 CHAPTER 4
- A!
Decision Demon
Cognitlve Demons
grouping are a t work hiding the camouflaged in each part
of Fig ure 4.2la and c. Carnouflage does not n eed to be entirely
about h.iding things. In World War C navies painted some of their
ships w ith very drarn..1.tic patterns. This 11dazzle camouflage" \v·as
designed to make a d early visible object hard to understand.
Perceptual Committees Revisited

N mv tha t we've seen some of the proble m s the visua l sys tem
A 0 x R H
has to deal 'vi th in pa rsing an image, let's go back a nd flesh o ut
the n otion of perceptual committees in middle vision. Low-level
Feature Demons v6-·ual p rocesses deliver fairly straightforward bits of infonna tion
about th e presence of a line here and a color there. However,
that collection of information need s to be interpreted befo re we
kri o'"r '"·hat we're seeing. Middle vision behaves like a collection
of specialists, each vrith a specific urea o f expertise and indiv idual
FIGURE 4.22 The Pandgmonlum opinions about w h a t the input might mean . The goal is to h ave
for r900gnizing an A. See Web a s ing le ans wer emerge out of this di versity of opini ons.
Activity 4.2 : Pandemonium to view TI1e idea of perceptio n by committee has a long hi story, though the specific
this in action. me taphor takes different forms a t different times. One early version was O liver
Selfridge's (1 959) Pandem oniun1 mode l for le tter recognition (a relatively
simple s ubset of the object recognition problem). Selfridge called his com-
mittee mem bers " dem ons." (I ndeed , the ' ""·ord p(mdemoni111n 1 which ·w·e define
as "noise and ch aos," was originally John J\.1.Hto n's l1 608--1674] word fo r the
abode of all de mon s in Paradise Lost.) Selfridge's theologically n eutral "fea h1re
demons" found vertical lines, acute a ngles, and so fo rth. "Cognitive d emons,"
one for each lette r, had ideas a bout the features of the ir le tters. A cogni tive
dem on looked at the work of the fea ture de mon s and made noise proportional
to the ev id ence for its le tter. A 11decision d em on " listened to the committee
of cogniti ve d em ons and ide ntified the le tte r on the basis o f the loudest yell.
Rgure 4.22 sho\o\'S a screen shot fr orn Web Activity 4.2: Pandemonium . Visit
the book 's website to experience p e rception by conunittee in action .
ambiguous figure A visual stimulus O f course, n o on e !'ea lly thinks that object !'ecognition is accomplishe d
that rise to tw:> cr more Interpre- by de m ons or co mmittees in :a ny lite ra l sense. The physical s ubs tra te o f a
tations of Its Identity or structure. "committee" is probably a massively in terconnected set of ne urons tha t takes
Necker cube An outllrra that Is fn put d.r. bt \rfrtue o f its specifi c connections, produces an output tha t can
perceptually bi-stable. Unlike the s tube read as the committee decision by other processes else\o\·here (1-1ozer, 1991).
atlm with most stimuli , two lnterpreta-
tklns continually battle for perneptual
The details of how this might come to pnss rue beyond the scop e o f this book.
dominance. Still, in the nex t section we w ill identify som e of the workin g ass umptio ns of
these com.mi ttees.
COMMITTEE RULES: HONOR PHYSICS AND AVOID ACCIDENTS TI1ough

we usually come to a sin gle cohe rent inte rpreta tion of the current co ntents of
the sensory world , the d ecisio ns m a de by perceptual committees n eed n ot be
fin a l. An ambiguous ftgure is a figure that generates two or more pl ausible
inte rp re tations. For example, Figure 423a sh ows the famous Necker cube.
(a) (b) (c) (d)
FIGURE 4.23 The w ire-frarne Necker

c ube {a) c an look lik9 eith.w of the
sl1()\,l\ff) in parts f./J} or (c}. It is much less
lik9fi/ to b e seen as a oollec tion of flat
regions o n the p age (d), e\19n though
that's realty what It Isl

Middle-visio n comrni ttees of the visual syste m are willing to e ntertain either
of the two solids sho\o\oT\ in Figure 4.23b and c as interp retatio ns of the w ire
fram e pictured in Figure 4.2311. H m . .·ever, the sys tem is tm\vi.lling to .rei1di1y
consider any of the infin.ite number of other p ossible interpretation s of the wire
irn.age--for exa mple, the flat one in Ftgure 4.23d. Figure 4.24 s hows another
classic arnbiguous figure, now pressed into the service of comme rce.
Necker cubes and duck-rabbits are really the exceptions that prove the rule.
Every image is, in theory, ambiguous, but the percep tual conui1ittees a hnost
al ways agree on a s ing le interpreta tion. Consider, for example, Figure 4.25a. The Duck-Rabbit
This pattern mig ht represent the re tina l image projecte d by the scene depicted
in Figure 4.25b: four s urfaces \'\.; th different shapes, arranged in diffe rent MILK STOUT
orientations, and at different di.stances. For you to see this as a s ingle shape
divid ed into four s quare regions, yo u would have to be viewing the scene
from. exactly one, very precise loca tion- wha t object recognition researdlers
call an ace ldental viewpoint. Any slig ht shift in viewpoin t-say, movin g just
FIGURE 4.24 A d asslc ambiguous
slightly to the left-would d estroy the illus ion of the four equid istant s quare
figure gets a ngw rolQ.
region s. (Web Ac tivity 4 .3: Objec t Ambiguity illustrates this situa tion v.'1 th
an animation that makes the reJiltionship beh-veen and b clearer.)
The perceptual committees knmv about accidenta l vie\.vp o ints. More.specifi-
they know enough n ot to bet on them . l11e d1ances that Figure 4.2.5a is
the hvo-dimensional represen tation of the three-dimensiona l scen e in Figure
425b are so slim that the visual sys tem refuses even to consider this possibility.
A second set of asswnptions made by the visual system involves an implicit accidental viewpoint A '1ewlng
under shmding of some aspects of the physics o f the wo rld . For example, position that produces some regu·
return ing to the Kanizsa fig ure (see Figure 4.12), we infer the arrow-shaped larlty In the visual Image that Is not
prooent in the world (e.g., the sldoo
object, in pa rt, beca use of our implicit understanding tha t solid objects block of two Independent objects lining up
light. This Lmders t.a ndingof o pacity causes us to infer that the lines disappear perfecttf).
because something is blocking o ur view of them _Cal ling this underst.:1 nding
"implicit" means tha t we n eed n o t be able to verbalize th e r ule in orde r to use
it. M onkeys seem to see the subjective con tours \vi thout benefit of instructi on
in physics (von d er Heydt, Pe ted1ans, a nd Ba1m1ga rb1er, 1984), a nd we rn..'ly
a.ss lm1e tha t simifar abilitie.s ex tend far in to the ani mal kingd om. We and our
visual systems " know" this and othe r physica l principles in the same way
tha t a ball "knows" about gravity.
These principles and assumptions may seern so obvious as to be meaningless.
But reme mber that an image has n o mea ning w hatsoever until middle- and
hig h-level-visu l dig into it. One committee uses the knowledge
that opaque objects oca ude other o bjects behind them to genera te p lausible
interpreta tions of image elements like the notched circles and dea d-end edges
e
(•)
(b)
,,.::-·-·-·--."·"/
-"L
----· -
_7'..... -
· ---- +
FIGURE 4.25 Accidental Suppc>SQyou see thgsefour squares. \M!at

is the state o f the \NOr1d that produced this vieJW? ft>) Ma)'be the eye just happens
to b9 in the right position t o see four arbitrary shap9S at arbitrary d9pths lin9 up t o
form the pattern. That would be quite a coincidence. That V¥'0Uld be w hat is called an
..accidental viewpoint."

106 CHAPTER 4
in Figure 4.12. Another committee consid ers all the possibilities a nd d eval ues
any tha t involve accidental vievvpoin ts,.redudng w hat is initiaUy a
unsolvable problem (finding the one rorrect interpretation out of the infini te
number of possible ones) to a potentially solvable one. And so we p roceed
htrther and further into the visual system, until a si ngle, gen erally correct
interpre tation emerges.
Figure and Ground

Armed with this understanding of the g r01.md m.les of oomm_ittee d eliberation,
we continue with the effort to go from simple features to recogni zable
objects. The edge- and region-finding medrnnism s, discussed earlier, would
divide Flgure 4.26in to yellow and red regions wi th out much difficulty. But
FIG URE 4.26 What is fi gure and
how s h ould those regions be und erstood? It is extremely likely that \<Ve would
what is ground and why? inteip ret the illustration as two re<l figures on a yellow background. Tha t is,
we would infer that the yellow wou ld continue behind the red objects if we
co uld Lift them up to check. That need n ot be the case, however. \ Ve could be
looking at yellow and red puzzle pieces, cut to fit ead1 o ther perfectly. Or we
could be looking at hvo yellow objects, a smaller one on the le ft and a larger
one vvith asquiggJy hole o n the right, both si tting on top of a red background.
TI1e ability to disting uish fig ures (objects in the foreground) from ground
{surfuces or objects lying behind the figures) is a critical s tep on the path from
image to object recognition. Like the finding of edges and regions, it is a process
governed by a co llection of principles acting together, ns if in another of these
perceptual committees, to decide ho'"' the visual world s hould be lmderstood.
As before, the governing goa l is to determine the m ost likely reality behind
the im.age on the retina.
figure-ground assignment The Like the grouping princip les, the topic of figure-ground assignment
process of determining that some became important in visual perception because of the work o f the Gesta lt
regbns of an Image belong to a psyd1ologists. Figure 4.27 shows the classic vase/ fare figure introduced m
foreground object (figure) and other
regbns are part of the background the 1920s by the Danish psychologist Ed g.u Rubin (1886-1951). It mus t be
(ground). the best-known illustration from the Gestalt school, and it is analogous to th e
Necker cube in that it illustrates one of those rare cases in wh ich a percep tua l
surroundedness A rule fer figure-
ground assignment stating that If committee has a difficult tirr1e re.i.chin g consensus. All visual sti muli may be
ooe reg bi Is entlray surrcunded by inherently ambiguous, but the processes that determin e figure and ground
another. It Is llkay that the surroo1ded almost always manage to com e to a single conclusion, as they do in Figure
regbn Is the figure. 4.26. \Ve are s urprised w hen the process fails and d elivers two or m ore in-
terpretations. The s urprise we register w hen the Rubin vase 11flips" to a pair
of faces (or vice versa us of the perceptual stability that we usually
take for granted.
\Vhat principles are at work in the ass ignment of regions to figure or
ground ? We can list some of them.;
• Surro1mded11t.>ss. lf one regio n is entirely surrounded by .mother, it is
likely that the surronnded region is the figure. This surroundedness is
a factor for the red region on the right in Pigure4.26.
• Siu. The s maller region is like ly to be figure. 111e cow is s maller than
the fie ld in w hich she stands. T11ese letters that you' re reading are
s malle r than the p age.
• Symmetry. A synunetrk al region is 1nore likely to be as figure. How
likely is it tha t the hvo yellow regions in Figure 4.26 JUSt happen to have
FIG URE 4. 27 The Rubn vasQ/facG figurQ . Do you seG a vase or two facoo in pro-
file? Notice that thQ boundary mark9d by the grn€<1 linQ changgs its allQglanoe whoo
thQ 1igure flips The boundary is •owned.. by the 1igure.

FIG URE 4.28 Why does the beige rEig ioo app01ar to b€I the figure when the beige
and gr9en regions about equal in mo st Parallel oontours arg
taJ«in to belong t o the flgurn.
the sy mme trical conto urs facin g ea.ch other over wh a t would be the red
gap on the left?
• Pnrallefism. Regio ns wi th paral lel contours are m ore likely to be seen as
fig u re (does beige o.r green '\'\rin" as figure in Figure 4.28?). Again, how
like ly is it that tw"o contours would be parallel wi th one an other if they
did n o t belong to the same object?
• Extremal edges. Figure-ground calculations are intended to ans,ver the
question ''ls region A in front of region B?'') n fhe upper leftpan,el 9f
Figure 4.29, the edge of the w hite d isk has sh:Jdin g s uggesting th<1 t the
ed ge is curvin g away from the v iewer and toward the g ray texture. The
viev>'er concludes tha t the disk mus t be closer and must be the figure. ln
the upper right panet the w hite ring has this "extremal ed ge" cue, a nd
it becom es the figure. No te that the cue is s trong enough to overwhelm
cu es like surrow1dedn ess and size (S. E. Pa lmer and Ch ose, 2008),
w hich seem to t1i p the figure-ground assignmen t in the bottom panels
of the fi gure.
• Relative motion. How s urface d etails move relative to an ed ge can also
de tem1ine w hich p ortion of a d isplay is the foregrotmd figure and
w hich is the background (Yonas, Craton , and lhompson, 1987).
(See Web Activity 4.1: Gestalt Grouping Principles.)
Cl•
A COMPLICATED BUSINESS Altl1ough tl1e assignment offigure and grotUld
is governed by r:u1es, it is not a simple process. Cons ide r, fo r example, Ftgure
4.30. 111e s urroundedness, s ize, symmetry, a nd parallelism ndes combine to
fim1ly estabLish the purple squares as fig ures on the g reen background. But
w hat about the circle in the middle of each squa re? Each one is s mall er than
the purple square and completely s urro unded by it Moreove r, each circle is
perfectly symmetrical. Yet, of the three, only the center circle (Figure 4.3LV.i) is FIG URE 4.29 In the top left p anfll,
perceived as being a separa te object (figu re); the circles on the left and right the •extremal eclge· cue makes
are seen as holes in the ptuple squares. the image bok like a sphere o n a
You ca n (a nd should ) specula te about the factors that d etermine whether a background.
circle is seen as a ho le or a sp ot (C. Yin, Kellman, a nd Shiple)'i 1997). ln Figu re
4.30c, the answer has sornething to do vdth the fact that we inte rpre t those
three isolated patches of a dollar as one object, occl uded by the purple square.
Gi ven that theory, the circle must be a h ole. One lesson from this fig ure is tha t
the journey from image features to pen::eptual objects is no t a s imple one-way
(a) (b) (c)
FIG URE 4.SO Which cirdes are figures and \o\lhich ar.e hol.as?

108 CHAPTER 4
(a) Relatable (b) Unrel.1table FIGURE 4.31 Two edges arQ relatable 11 they can be connected with a
smooth convex or smooth concave curve (a}, but not If the connection requires
an S CUTVQ fl>). (After f<Qllman, 1QQB.)
street. Figtu'e-grotmd p rocesses divide up the itn..<tge into regions that can
be recognized as specific objects. At the same time, object recogni ti on
influ ences fig ure-ground assignment (Pe terson a nd Skov1.; 2008). (See
Web Essay 42: The Role of Knowledge in Flgure-GrollldAssignment.)
Traffi c is nmning in both directions. In Figure4.30c, our understanding
that \'v"e are lookin g a t one d ollar bill is based on our unde rstanding that
Elbow Scwve this object continues behind the occluding smfoce of th e purple square.
Dealing vd th occlusion is a n important aspect of middle vision and the
topic of t11e nt:xt section.
Dealing with Occlusion

Objects are rarely kind enough to present the mselves to us in splendid
isolation on blank backgrotmds. In the rea l w·orld, objects a re often par-
tially hidden by othe r objects. Indeed, three-dimensional objects often
hide parts of themsekes. We've a lready discussed how edge-finding
prcx:e.sses can fabricate illusory conto urs on the basis o f the physics of occlu -
sion.. Now, let us thin k about the work re<t uired to connect the visible pieces
of occlud ed objects by in fe rring the presence of h idde n pieces of the object
w hen necessary.
relatabilily The degree to which two 1lle Gestal t p rinci pi e of good continuation comes in to play here (see Figu re
line segments appear to be part o f the 4.14). How d o we unde!'stand the continua tion of a contour w he n it d isappears
same contour. from 'iew, heh.i nd a n ocduder? Philip Ke!Lnan and ll1omas Shipley (1991)
heuristic A mental shortcut. speak about the relatablllty of two conto ur segrnents. Relatability is illustrated
nonaccldental feature A feature of in Ftgure 4.31 . Do the h\.'o black line segm ents in each part look like they belong
an object that Is no t dependent on the to the same ctuve, occl uded by the black square? For Figure 4.3ln, the answer
exact (or accidental) viewing position of is 1'yes." For Figure 4.Jl b, the answer is "no." Kellman and his coll eagues argu e
the observer. th at the cri ti cal difference is tha t the lines in Fig ure 4.3117 can be rebted by
a sini.ple curve like an elbow or a ben d in the road . The lines in Figure 4.3lb
require a m ore complex S curve to make a s mooth connection between them.
TI1e visu al syste1n is wn. .•illing to propose such an elabora te relations hip, so it
concludes that the lines are not related at a ll (that is, they are not parts of the
sa me object). Like the fi gure-ground rules, this heuristic (mental shortcu t) is
not in.fallible (after a ll, some objects really do have S'"5h.aped contours). The
occlusion committee is apparently willing to accept a fe,v m issed comple tions
in order to reduce the vast number of possible comple ti on s \Ve wm.tld have
to consider if we tried to conn ect every pair of occluded ed ges. For more on
dealing with occlusion, see Web Essay 4.3: Dynamic Occlusion an d
Web Activity 4.4: Infant Object Perception.
Additional h e uristics e me rge w hen we m ove from tw"o dimensions
to three, as Figu re 4.32 sh ows. Here we see a pair of box-shaped objects1
one partially occlud in g the oth er. The overlap of the boxes p roduces a
variety of different line junctions1 a ll of w hich can be dassified as Y, T,
or arrow junctions. T junctions a lmost a lways 01.."'C\.tr w hen one surface
occludes another; Y and arrow junctions almos t a lways correspond to
com ers an.d thus d on 't signal occlusions. These rules fail to hold true
only w hen we're v iel'v-in g the scene from an vie\vpoint. Hence1
the variou s jtmction types are knO\-vn as nonaccidental features (Lowe,
FIGURE 4.32 Diff0r€111t sorts of Hni9 junctions
ha\19 dlffflirer1t rneanlngs. T junctions indicate 1985)1 and they provide another p otential tool for use by the perceptua l
ooclusion o f one region b y another. Yan:::t conunittees dl.arged with dividing the scen e into objects and deciding
arrow junctions lndlcati;i corners, if one object is occluding another.

5 5 NNNNN H FIGURE 4.33 Global letters com·
5 5 N H p osed of local letters. Da.iid Navon

argued that perc eptual p-oCBSSQs
55555 N H work from th9 glo b...;J t o thQ local.
5 5 N H
5 5 NNNNN HHHHH
Parts and Wholes
O bjec ts in the wor ld tend to have p arts. Indeed, it is n ot obvious how to de- global superiority effect The nnd-
fine a vis ual object. In so1ne sense, you are an object; so is your n ose, each ing In various exp€flments that the
eye, your shirt, and so forth. [n the lab, stimuli like those in R g ure 4.33 ha ve properties of the whole object take
precedence over the prcperties o f
been used to investiga te the rela tionship of visual objects to the ir comp on ent parts of the object.
parts. Look for Hs in Figure 4.33. Did you find the big (global) H more quickly
than the little {local) Hs? Us ing s timuli like these, Navan (1977) fo und
tha t the global le t ters interfered w ith naming of the local le tters more than
the local le tters interfered with recognition of the global letters. nus global
superiority effect is consistent with an iinplkit ass umption we've been m ak-
ing throughout our discussion of middle vision: that the first goa l is to C.'.U'Ve
the retinal image into luge-scale objects.
Jn the Navon letters, the division into parts a nd w holes, local and global,
is easy. It is n ot so easy to ask h ow to "carve" a h uman form into parts like
legs and arms. On e he uristic is illus trated in Figure 4.34. Don Hoffman and
Whitma n Richards (1984) no ted that w hen one blob is pushed in to another, a
pair of concavities is crea ted in the silh oue tte o f the resulting tv1m-par t object
(Figure4.34a). A process that embodied this bit of physic;: would conclude that
valleys. rathe r than b umps, sh ould be used to mark part boundaries. Thus,
we a re incli ned to parse the object into two parts as shown in Pigure 4.34b,
not into three parts as in Figme 4.34c.
Summarizing Middle Vision

Before moving o n, it is worth briefl y revievdng how we got here. Early-vision
processes gave us the loc.a.I fea hires in the visual world. Middle-vis ion pro-
cesses began the work of lmderstanding w hat those local feahtres might be
telling us about the state of the world. 1lle work of the middle-vision processes
d iscussed so far in th.is chapter m.ight be s umrn arized in five principles:
1. Brins together that whiclr should be bro11ght together. \Ve have the Gestal t
g ro uping principles proximi ty, pamlle lisrn, symm etry,
a nd so forth ), a nd we h ave the processes that complete contours and
objects even when they are partiall y hidden behin d ocduders (e.g., the
rel.."l. tability heuristic).
2. Split m;1111der tlzat which should be split asunder. Complen1ert.ting the
grouping principles are the edge-fin ding processes tha t div ide regions
from each other. Fig ure-grolmd mechanisms separate objects from the
backgrmmd. Texture segmenbti,qr;;i p rocesses divide one region from
th e next on the basis of imag stalistfcs. "
FIG URE 4.34 Find ing parts from
boundari9s. (a) WhQl'"l one blob is
pushed into another. a pair of oonca\'i-
ti9S is c reatsd. We know this {imp licitly,
o f c ourse). 3fld we work backW'afd
from this fact to carve this figure into
the t...vo parts shown in (/;) rathar than
accord ing to a diff€.fo9nf: scheme. such
as the o ne shown in (c).

110 CHAPTER 4
3. Use ·w!ra t IJCm know. Two-dimensio nal ed ge configuratio ns a re taken to

ind icate three-dirnensio na:l corners or occlusion and objects a re
dfvided into p arts on the basis of a n implici t know led ge o f the physics
of image fo rmat ion.
4. A'L'Oid ncddents. Avoid interpretations that require the ass\.Unpti ons
of highly accidental combinations of features or accidental
viewp oints.
5. Seek consensus and m'Oid ambiguity. Every irna ge is ambiguo us. There a re
always multiple, even infinite physical s ituations that could generate a
given image. Usin g the first four principles, the "committees" of middle
vi sion mus t elim inate all but one of the possibilities, thereby resolving
the ambiguity and deliverin g a s ingle solu tion to the perceptual
problem at h and .
From Metaphor to Formal Model

Bayesian approach A W"f of for- 'Ne have been talkin g about "percep tual cornmittees" in middle vision. While
malizing the Idea that our perc eption Is we h o pe tha t is a useful way to thi nk abo ut w hat 's going on, it is jus t a
a com bination of the current stlrnulus me taphor. There are formal, ma the ma tical ways to mod el the way in wh ich
and our knowledge about the condi-
tions of the wortd-w hat ls and Is not
knowled ge about regularities in the world can co nstrilln th e interpretatio n of
lli<Efy to occur. ll1e Bayesian approach mnbiguo us sensory input. One of the m os t frui tfoJ appro.."'\ches is k.nmv n as
Is stated mathlmatlcally as Bayes' the Bayesian approach (Yuille an d Kersten , 2006). TI10mas Bayes (1702-1 761),
theorem : P!AIOJ =RAJx PiO!A)iP(OI an eighteenth-<"ell tury P resbyteria n min ister, \Vas the firs t to d escribe the re l-
whlcll enables us to calculate the evant 1n.:'lthematics.
probability 11'1 that the world Is In a
particular state (A) given a particu lar Look a t Flgure 4.35. O ur visu al system looks at the s timulus and m akes an
cbservatlon (0). observa tion. As noted earlie r, there is an infinite set of hypotheses tha t could
be based on the observation. How does ou r visua l system decide that one of
these is the be.st hyp othesis? The Bayesian approach as ks LIS to think about
tw o factors. Firs t, before you look at anythin g, h ow likely is w hat you are
prop osing? Th is is known as the prior p robability. The prio r probability of a
un icorn is m uch lo\o\>-er th an the pri or probabiJity of a co\\.". 111lis, if you make
an observation tha t seems consis tent with eith er unicorn or cow, you should
be more ind in ed to accept the cow hypothesis. Second , h ow consistent is each
hypothesis \11,rith observation? Suppose you had the hyp othesis that you we re
looking at a red si.1uare in Figure 4.35. TI1e p r ior p robabil.ity of a red sq uare is
n o lowe r th an the prior p rob.."lbili ty of a green square, but the evidence in the
obser vation does not s upp ort the red square hypothesis.
So, in Figure 435, let's consider three hypotheses: Hypothesis A p roposes
E+ tha t w e' re looking a t h.vo green rectangles. The ligh t one has a n ar row letter E
on it, and the dark one ha s a narrow numeral 3. TI1ese just happen to precisely
line up in order to p roduce a B like the one we observed. Tha t is cons istent
Hyp othois:B with the observation, but its prior p robabili ty is low because the Eand 3 would
EB l +c need to line up very p recisely- an accidental viewpoint. Hypothesis B s uggests

that this is a green square with a Con it. That is not particularly unlikely, but
it is not consis tent \vi.th the sensory dnta. Hyp othesis C p rop oses that th is is
a green square w ith a Bon it. Tha t d oes not require anythin g very unlikely in
the world, and it is consistent with the obsen ;ation .
\Vhat is rea lly powerful about the Bayesian t>1pproad1 is tha t these sorts of
ideas can be reduced to fo!'mal, mathematical equ ations. Tha t math is beyond
the scop e o f this text but a brief introductio n can be fo und on the \-vebsite 1
FIGURE 4.35 \"moo the v rual sys- Web Essay 4A: Bayesian Analysis.
tem is faced w ith a stimulus, it tries to
figure out the most likflly situatio n in the FURTHER DISCUSSION of tho Bayesian approach can be found in
vvorld that has produc 9d this particular Oiapte< 6 on pages 11'.&-187.
pattern o f activity.

Material Perc eption:

The Everyday Problem of Knowing What It Is Made Of
We have assembled a rollection of middle-vision tools that w ill b e very
useful li\'hen It comes to recO;Jntzing cbjects, but befcre we move on
to object recognition , let's take a moment to think about what things
are made of. You are very gocd at what we can call "rnatelial percep-
tio n." If you look at Figure 4 .36, you l'.111 be able to say something about
each of these surfaces. For instance. Figure 4 .36b is shiny \'lhile Figure
4.3& is not. From very si mple exampe.s like thls, we can derive some
important principles of material perception. Rrst. material perception is
not about recognizing the objoct and knowing what such objects are
made of. You rould have a rubber ball , a metal ball, or a ball made of
roc:k cr Ice. They \oVOuld all be simpe spherns but you would know from
the usual eense of "object." In Rgure 4.36f, you can Identify sand. Sand,
like snow o r water. lsn1 an object like a rock or a desk. It Is "stuff" (Adel-
son . 2001). but you can stll Identify It from Its surface properties.
Figure 4.36 Illustrates a Secord Important aspect ot mate-
rial perception. You can use the visual properties of a matertal to (a) (b)
perfon11 two related but not ldentjcal task s. You can categorize the
material ; so, Figure 4.3ee is cloth. And you can estimate how the
thing would behave. Would that cloth be rough and itchy against
your skin? W hat wculd it feel like to try to pick up the stuff in Rgure
4. 3ca (Fleming, 2014).
How do you i:ertorn1 these acts of material perception? There
are reliable visual cues that are generated by the ways that light
interacts with phy$cal matertals. Think about a shiny t:el l o r a sur-
face like Figure 4.36/J. There are "higl11ig hts." bright spots, on shiny
surfaces. Thase have specific proper11es. For Instance, they tend to
be less ri chti,r colored than the rest of the shiny suriace. Indeed, if
you take away the a sphere -Mii no longer look very shiny. tel (JJ
Comptrter graphics hwe gotten very good at simulating the physics
of different surfaces. The problem is that It is easier to go from the
physics to the creation of a visual stimulus than it Is to go from the
visual stimulus back to the physics. Think about plastic. You have
a pretty good idea when ycu are looki ng at something made of
plastic, bcrt just think of all the varteti es of su rfaces that fall into that
category, from clear plasti c bags to shopping bags to children's toys
to plastic de.:::::k chairs. The latest animated m0\1e may do a great
job rendering shopping bags and deck chairs. bcrt you won't find
a simple rule that w ill let you categorize all those things as J:Jastlc.
How are you d oing this very tricky task? one possibility is that you
are !earning that specttic features oorrespond to a substance like (e)
plastic in this specific case. Then you are creating a "generative"
model (Fleming, 2014) wh ere you estimate w hat that substance
would look like under a range of different conditions (dimmer light.
light source over to cne sid e, and so on). From this, you build up an
Internal definition of plastic that allQ1;vs you to categorize plastic
FIGURE 4 .36 You are remarkably good at figu ring o..rt w hat things are
made of.

112 CHAPTER 4
the next time you see it, even under quite different conditions. This is
a bit like your definition of object categories-a t op c v-te will return to
later. What. after all, is the definition of "chair"? It might be a thing ''nth
four legs that you can sit on? Unless it has three le;;is or no legs, or it is
a table that you could sit on but shouldn't or it is a horse. However you
do these tasks, you d o them remarkably fast. Given an exp:JSure of 40
rns (1/25th of a second), you can identify many tyl""S of material and
even tell whether they are real or fake (Sharan, Rosenholtz, and Adel-
son, 2009).
Object Recogn ition

Now, a rmed wi.t h tools that s hould be useful for object recognition, le t us
e\t\Pl to Quirogn's discovery of cells that respond to highly specific objects
823033 Q'ulroga et Traveling the what pathway from Vl into the temporal
lobe, we can see a progressive change in the respon ses of cells in different
areas. In Vl , cells respond best to lines and edges in very specific ar-
eas of the visual field . In V2, we ge t early s teps from local features to objects.
As noted before, these cells have a sensitivity to ''borde r O\Vnership." They
are .Jlso sen sitive to illusory contours (Peterhans e t al., 1986; von der Heydt,
Pe terhans, and Balm1garh1er, 19&4). By area V4, cells appear to be interested
in much more complex attributes, like those shm\o'TI in Figure 4.37. No one
has ever d.efinitively de termined the perfect set of stimuli for V4 cells. Figure
4.3711 shows som e geome tric ideas from Galfrmt, Braun, and Van Es.sen (1993);
Figure 4.37b shows a s mall subse t of a m llecti on of bumpy blobs from the Con-
n or lab {Kmutzi and Connor, 2011). The images in Figure4.37 illus tr.ate a fact
about objects: while you can vary o rientatio n or colo r or other bask features
in a systematic way, it is much harder to imagine ho\".' you vary object shape.
What are the dilnensions o f shape? \ Ve really do not know. h1 Figure4.37b the
darkness of the circles indicates how well n specific cell responded to each of
these items. As you can see, this cell seems to have a taste for stimuli with a
(a)
(b) QO QQCCOOOO
OOOOOOOOC)O
0000000000
ocoooc oo
OOQQO 0 0
QOOOOOQOCO
00000 0 0 000
0 0" " 0
FIGURE 4.37 Respons9 o f V4 C€1lls to diffo91'Qllt shapi9s. (a) GaHant, Braun, and Van
Essen tried this set of stimuli. Warm colors indicate m ore respo nse from ong pa.r1icu-
lar ceH. (b) Connor and his ooll.eagues presented these shapes to V4 neurons. Darker
circles indicate rnore respons9. (Part a from GoJlant, Braun , and Van Essert, 1000: b
from Kourtzi and Connor, 20 11 .)

•
feature p ointing to the right. The connection to object perception is illustrated (a) (b)
in Figure 4.30. This V4 cell mig ht respond to a stimulus like the one in Figure
4.38a, but it vi.rill not respond well to a .stimuJus Uke the one in 4.38b, because
that black object d oesn ' t really have a feature pointing right. Its p oint is an
accident of occlusion. TI1e border ownership rules, established in V2, n ow be-
come part of the process of tmderstanding object shape (Bushnell et al., 2011). FIGURE 4.38 A cell in V4 that
Things get more complex as we move farther into the temporal lobe. Figure respondOO well to sharp features point-
ing to the right would r9spond wgll to
4.39 ill ustrares the hopeful h ypothesis tliat a vast family of s hapes like those
'3) but no t to (b) becauss that sharp
in Figure 4.37 might be a basis for object perception just as letters and syllables point Is the ..acctdental" product of
can be the basis for words (Ungerleider and Bell, 2011). We may not know occlusion.
exactly w hat optimally activates individual cells as we progress d own the what
pathway into the temporal lobe. However, as menti one d before, fWlctional
imaging .studies can show u s that different regions of the cortex are activated
better by some categories o f stimuli than by others. One way to show this is
a so-called subtraction method. If you show a human observer a series of
pictures of spaces like rooms and fields and city streets, lots of pieces of the
brain will be activated by these places. Now sh ow the observer other pictures
with similar properties that do not happen to be places-maybe scrambled
versions of the places, maybe objects or abstract designs. Again, large areas of
brain will be active. However, if you subtract the two p atterns of activation,
there w ill be at least one region, now known as the parahlppocampal place
area (PPA), tl1at is specifically and reliably activated more by places than by
other stimuli (Epsrein and Kanwisher, 1998). Other well..,stablished special- subtraction method In functional
ized include the fusttorrn face area (FFA) and the extrastrfate body magneUc Imaging, brain actMty Is
area (EBA) (see 'Pigure 4.4). measured In two oorx:lltlais: one with
Thus, a network of visual areas gives rise to ceUs in the temporal lobe with and one Without the hlvolvement of tile
mental process of Interest Subtract-
tl10se very specific sensitivities that Quiroi;a et al (2005) report But w ha t is ing the two cord ltlons shows regions
going on when a ceU responds to the actress Jennifer Aniston? Could it be of brain specifically activated by tllat
that, in listening to a Jennifer Aniston neuron, the researchers were listening process.
in on the act of object recognition? If the image of Jennifer A n iston fa lls on parahlppocampal place area
your retina, you m ust be able to take that raw image and generate a descrip- (PPA) A regloo of extrastrlate visual
tion that can be matched to a Jennifer Aniston representation in memory. Of cortex In humans that Is speclflcally
course, if you have no idea who Jennifer Aniston is o r what s he looks like, you and reliably activated more by Images
won't recognize h er. Even so, you will still recognize her as :a woman. How? of places tllan by other stimuli.
This is not :a trivial problem, and we can quickly dispense ""'ith some trivial face area (FFA) A region
solutions. You are n ot matching the "pixels" of the Aniston image with the of extrastrlate visual cortex In humans
that la specmca11y ard reliably activated
pixels of an Aniston rep resentation. Figure 4.7 sh owed that the cell responded by human faces.
to various different images of the actress. Moreover, we know tha t a d etailed,
point-by-point representation of objects is n o t transmitted to the parts of the extrasbiate body area (EBA) A
reglcn of extrastrlate >;sual =tex In
brain interested in objects. As we've been discussing. the raw image is analyz.ed humans tllat Is speclflcally and reliably
by middle-vision processes, and it is the o utput of those prcx:esses that will activated by Images of the body other
be available to object recognition processes. than the face.
"'to..,. "'m o"' + "'bile" "'automobile"'
FIGURE 4.39 N9Uron s might be s9'"'1Sitiw to shapgs that can be put togath9r into a
recognizable object evm If no OOl's preferred shape looks like anything that we have

114 CHAPTER 4
Templates versus Strnctural Descriptions

The ide'-1 that we recognize objects by matching every pixel or even
matching every low-level feature of the input to a representation in
mem ory is what might he called naive template theory. lt won' t
work even for si mple objects. The ba:sic idea of a template is rather
like a lock a nd key. As we will see in Chapter 14 and 15, the Jock-
and-key meta phor is quite apt in s mell a nd ta ste, \vhere the ''key''
to be recognized is a molecule like a specific od orant and where
that m olecular key presents itself in mo re o r less the same shap e
every tim.e. For instance, look at the "shape--pattem " theory o f
olfaction in Chapter 14.
Figures 4.40 a nd 4.41 show tha t recognizable visual objects
like the letter A are much less well behaved. Figure 4..40 illustrates
FIG URE 4.40 A basic template. the lock-and-key template idea. A letter A stimulus falls on an array of spot
d etectors. If the A falls on the filled d etectors and not the empty on es, it is
identified as an A. Therefore, the array of detectors serves as an A template.
The difficulty w ith a nalve template m odel is that too many templates a re
A -4A A required.. All of the objects in Figure 4.41 are As. If \Ve need ed a new te mplate
for every lette r in every position and orienta tion, we would nm out of brain
before w e ran o ut of alphabet. A11d just think about the p roblem of building
a aA a a Jennifer Aniston rep resentation i n th is way.

One way out of this problem is to no tice tha t all the As--a t least all the
capitalAs--share abasics tmcture. Cnstead of ma tching each p oint in the image
FIG URE 4.41 Th9 probleim with tern- to 3 point in a te mplate, perhaps we perform 3 more concephMl match. Just
platoo Is that we need a lot of th€m . about any capital A be described by the relationship of its three lin es: the
two flankin g lines me-et, and the third line spans the a ngle cre.:ited by those
two lines. Now the image of the A is being m atd 1ed to a structural description
of an A, a s pecification of an obiect in tem '\S of its parts and the relationships
between the parts.
lvlany versions of s truch.ual'1escription hyp otheses have been proposed. A
key component of each theory is how object parts are re presented in the s truc-
tum l descripti ons. Marr and Nishihara (1971) used "generalized cylind ers'' that
could be scaled longer,. s horter, fatter, o r thinner to rep resent different-shaped
parts. Bied errnan (1987) proposed a set of geons ("geometr ic ions''); Figure
4.42 provides exarnples. Geans are specified as collections o f n onacddenta l
feah.ues (remember accidental features in Figlue 4.25?), so in theory a visual
system s hould be able to recognize a geon equally acctrra tely and quickly,
na'fve tempiate theory The pro- regard less o f how the geon is oriented in space (as long as it's not an accidental
DC'3•1 that the visual system recognizes view}. Geons are fea h.rred in model,
cbjects by matchlr>J the neural repre-
sentation of the •nagewith a stored whid1 we ll now describe in a bit more detai l.
representation of the same In
the brain.
structll'al description A descrip-
tion of an object In terms of the nature
of Its constituent parts and the rela-
tions.hips between those parts.
geon In Blederman's recognition-by-
components model, any of the +lgeo-
rnetrlc Ions· rut of wh lch perceptual
ctijects are built . FIG UR E4.42 111reeofthe36orso
geons in recognition-by-
recognition-by.components c omponents model of object recog ni-
model Blederman's model of obje:;t tio n. Each geion is s hown in three ori-
recognition, which holds that objects entatb n s, Illustrating the fact that thgy
are recognized by the ldentrues and are .:iasily reoognizabkj frorry ·ly. any
relationships of their component parts. viewpoint.

FIGURE 4.43 Combining geons can

c reate a v..ide variety of object repre-
senkillons. (a-c} Three objects •spelled
out• by different combinations of the
cylinder, noodle, and brCk goons. A
d iffer911t vi9w of the coff&e mug (d) and
a different instance of this ob)gct class
(e) 'NOuld lsoct to th9 same strtciural
description re in (a).
Just as the finite set of le tters in the a lphabet ca n be combin ed in vari ous
ways to produce an infinite nmn.be.r of words, the finite set of geons can be
used to construct a very large numbe r of object representa tions . Attaching a
noodle to the side of a cylinder gives us a coffee mug (Figure 4.43a). Moving
the noodle to the top of the cy linder produces d pail (Figure 4.43b). Changi ng
the cylinder to a brick gives us an a ttache case (Figure 4.43c).
Because geonsa nd relationships s uch as'' A is beside B" are designed to be
e<:jually recognizable from ma ny different vantnge p oints, structural d escriptions
composed o f these geons and relations s hould als o be reasona bly viewpoint
invarlant. M.inor .shap e variations also won' t alter the s tructural descrip tions
of geons. .!\s n oted already, this is the great advantage of s tructural descrip-
tions over temp lates: if we can derive the sam e stnictural d escription from
any picture of the object (e.g., the mugs in Figure 4.43a, and Figure 4.430',e ),
then we need to s tore only one representation of the object in memory.
Problems with Structural-Description Theories

We may have made too much progress. Viewpoint invariance is a good thin g,
but UOf!d a:uad 'D !lOU lf you wanted to,
you would be able to read tha t las t sentence wi thout turning yo ur book upside
d m-vn, but even Bied erman would say tha t there is something quite clearly
view p oint-dependent about le tte r recogni tion and, m ore genera lJy, object
recogni tion. There are other problems with strnctural descriptions as '"'""e ll. Fo r
example, it is no t d ear that geons or a ny of the other "alphabets '' p roposed
by s tructural-description models are adequate for the language of object rec-
ognition. Would the geon description of a boo k differ fro1r1 the description of
a box oon taining th at book? This objection mhely p,.oi.Qts out thnt a stnlet\ rq_l
d escription would have to be just a part o f the answer to the problem of ol>Ject
recogni tion .
In response to these questions about structurc1l-description models, re-
searchers s uch as lvlich ael Tarr1 Heinrich BUJ.thoff, and others returned to
template-like representations tha t they cal l ",,; ews." Support for view-based
representa tion s generally comes from experime n ts that use n ovel, ra ther than
familiar, objects. For e..xa mple, Tarr and Pinke r (1990) tra ined observers to
recognize lette rlike objects. During tralning, the objects were always s hown
in the upright orientation . 1llen, in a "surprise" phase of the experi ment, the
observers were asked to recognize rotated versions of the objects. 11le res ults
revealed a linear relati on ship between o rienta tion s hift and the respo nse
time to recognize the object. The mme the object was rotated, the longer the
observel's took to name it. This result s u ggests tha t the subjects may have viewpoint Invariance 1. A property
s tored a te mplate-like representation of the object during the training phase, of an object that does not change
and then recognized. the objects in the s mprise phase by me ntally rotating when otserver \!1ewpotnt changes.
2. A class of theories of object recog-
the m.isoriented objects b£tck to the upright views they h.:1d s tored in m em ory. nition that proposes r€1)resentatlons
Objects that can be identified on the basis of their geons can stlll sh ow of objects that do rot c hanga when
a dependence on vie,,..-point (Hayward and Williams, 2000). Ga uthier e t al. viewpoint changes.

116 CHAPTER 4
••••
FIGURE 4.44 R9cognizing thQSB •gr99bl9s" dOQS not SQQ'Tl to be
lndeper1dEnt, even If they are made up ofgoonlik.e
••• ,
shap8S. (Fran Gauthi9,r 9t al., 1QQ8.)
(1998) mad e novel objects that they named "greebles'' out of

geonlike parts (Figure 4 .44). Recognition of these novel objects
also appears to have a vie'"•point-dependentoomponent (Web
Activity 4 .S: Viewpoint Effects lets you test yourself in a s hort
experiment us ing s uch objects). H owever, view-based m odels
have thei r mvn problem.s. While d e bate be tween structmal-
1111
description and view-based theorists has some times been quite
lively (Peterson and Rhodes, 2003), '""can imagine that a tndy
s uccessful model of object recogni tion w ill incorporate &-peci:s
of both kinds of theory.
Multiple Recognition Committees?

lf d1apter has had one theme, it is tha t no s tep on the road fr om image
to recognition is taken by a s ingle process actin g alone. Prom the groupin g
of similar pi eces of the image to the segrega tion of figme £1. nd grow1d , every
s tep has been based on consens us--a committee decision.. 1t seem s likely that
the £1.d o f recognition is simUar. Indeed, recognition may not be a sing le act.
We can recognize a n object in multiple ways, perhaps simulta neous ly. As an
example, Figure 4.45 shows h vo birds. In the termin ology of Pierre Jolicoeur
(a) (b)
FIGURE 4.45 Two quik1 different

birds (a and b). a yet more diffB'ent ani-
mal (c), and a third Mbird' (cl}.

and hi s colleagues Qolicoeur, Gluck, a nd Kosslyn, 1984), 11bird' 1 i.s the enby# en1ry-level category For an object,
level category for these o bjects--the firs t word that com es to mind w he n the label that comes to mind most
qulcklywhen we It (e.g., 'bird").
we're asked to n.:"lme them . But these objects are also quite clearly different
At the subordinate level, the object
At a s ubordinate level -a m ore specific level ben ea th the e ntry level-Figure might be more spoomca11y named
4.4511 s hows a fox s pa rrow and Figure 4 .45b shows a cardinal. At a s uperordi- (e.g., at the supero rdinate
nate level-a broader level above the entry level-these h¥o objects, as well level, It might be more generally named
as the one in Figure 4.45c, a re all a nima ls. (e.g .. 'animal").
\Ve can imagine that each of these acts of recogni tion ("fox s parrow," "bird ,"
''a nimal") rnight rely on different stored representations and different analyses
of the visual stimulus. And we can imagine a geon accotu1t of "bird/' but a geon
d escripti on of "fox sparrow" would be harder to en vision.
recognition seem s bette r suited to a system in which the p recise de tails of a
particular view of the object are encoded . Recognizing the objects in Figure
4A5a--c as animals seem s like qui te a different act.
lndeed, a number of lud'iC.s }lave .!t.htn-...11 that it takes considerably longer
to recognize objects a t thesubordinllte or superordinate levels than at the en try
level. Even more imp ressively, studies looking at brain recordings provide
strong evidence tha t different parts of the bram are more active w hen people
are engaged in s ubordinate-level recognition than w hen they are recognizing
objects a t the entry level (Pa lmeri and Gauthier, 2004; Rich.ler and Palmieri,
2014). There a !'e several m ore interesting hvists. For exa mple, wh en s hown
an a typical member of a category, s uch as the bird in Figure4.45d, people are
faster to name the object a t a m ore s ubordinn te level ("os trich'1). And when
people becom e experts at recognizing a certain class of objects, s ubordinlte-
level recognition becomes as fas t as or fas ter than en try-level recognition (or,
looking at this a nother wny, we might say that the entry level shifts d ow n
one level when one becomes an exper t). (By the way, bird-watchers and dog
show judges seem to be the most ex tensively studied s uch experts {Tanaka a nd
Tay lor, 19911-) All these results point to the conclusion that the visual sys te m
is e mpl oyin g several different object recogni tion committees, each working
w ith its own .sets of tools, at the same time.
Faces: An Illustrative Special Case

Faces are an interesting special case of object recognition. Take, for exam ple,
the images in Figure 4.46. You sh ould have Little difficulty recognizing these
as faces, but you w ill probably have some difficulty quickly identifying \.vhid1
one has been mo dified . lf you turn the faces right si de up, however, one o f the
pictures will look quite strikingly '"-'rong (Thompson, 1980).
FIGURE 4.46 Whic h o f these two photos has been

altered? As you will see, both faces look fine do"A'n,
but o ne of them looks quite strikingty" when turned
right side up. (After Thompson, 1QBO.)

118 CHAPTER 4
prosopagnosia An lnablllty to rec- lltis is another demonstration of the p oi nt m a de in the previ ous section:
ognize faces. that d ifferent levels of rntegorization often seem to besubserved by d ifferent
double dissociation The phencm - con:mUttees using different types of infom1ation. The p rocesses that recognize
errn In which one of two !Unctions. a face asa face care little about inversion, and they don' t seern to be d isturbed
ard l.Jice versa.

ti!' bpd,jStortio,n of the face. The processes that recognize the face as belonging to
a specific individual (in this case, one o f the a uthors of this book) work poorly
o n inve rted fores a nd are very con ce rned '\.v ith the precise configuration of
congenital prosopagnosla A f0011 eyes, nose, and mouth. (See Web Activity 4.6: The Face Inversion Effect.)
of 'face blindness" apparenlly present
fran birth, as opposed to •acqtirecJ
Neuropsychology pro\>ides further evidence that the processes can be sepa-
prosopagnosla, • whiCh wa.11cl typically rated . Damage to specific areas in the temporal lobe o f the brain can p roduce
be the result of an injury to the navous prosopagnosla, a d isorder in w hich the cannot identify faces. (See
system . Web Essay 4.5: Face Blindness.) Though she m ay be abl e to recognize an
object as a face, s he will not know w ho the person might be. TIU.s is n ot to say
that the two levels of face recognition are s ubserved by completely different
systems (Gauthier, Behrmann, and Tarr, 1999). A neu ropsych ological mark of
truly separa te brain mo dules is double dissociation. Two functions, s uch as
bearing and sight, are d oubly dissociable if one can be damaged wi thout harm
to the othe r a nd vice \·ersa. Thus, you can be blind a nd s till hear or you can
be deaf and s till see. In foce recogni tion, it is possible to lose the s ubordinate
ability to recogaize specific faces w hil e retaining the ability to recognize an
object a fuce_ ft is n ot clear tha t the reverse even makes sense. How could you
recognize a foc-e as yom m other without recogn izing it as a face? Interestingly,
it is possible to be born with a specific impairment in the ability to recognize
faces . The exis tence o f this congenital prosopagnosla is a good in d ication
that there is a specific neural mo du le fo r fa ce recogni ti on (Beh rma nn and
Avidan, 2005). You may think you're suffering from this disorder w hen you
fail to m atch a name to a face. However, tha t is a much more cornm on failure
of memory. You can recognize the focei you just can't remember the na me tha t
goes wi th it. A prosopagnosic would not kn ow that this particular face """ru;
famlHar '"'· h_ile another one was not.
The Pathway Runs in Both Directions:

Feedback and Reentrant Processing
We've been discussing object recognition as a progression of steps down th e
what from Vl into the temporal lobe, an d as mentioned earlier, it d oes
seem to be the case that you can perform some acts of object categorization
in this feed-fonvMd m anner (Serre, Oliva, and Poggio, 2007). 111at said , it is
important to recognize that neura l processin g i.s n othing li ke a one-way street.
Most of th e routine acts of object recogni tion that you perform w ill in volve
information flowin g up and d ow n these pathways. All of the connections in
Figure 4.3 run both ways. Consider that receptive fields get bigger and big-
ger as you ascend the what pathway. TI1e cells care less and less about where
an object is in the visual field or how it is oriented.. They become increasingly
interested in the object's identity as, perh aps, the Eiffel Tower. However, you
know that the Eiffel Tower is right there and it has that red light in thnt s pot.
This precision is p robably achieved by going back do,,..,·n the patlnvay, once
you have some in forma tion about the object, and interrogating earlier visua l
areas about the details of this ins tan c-e of the object (Eps htein, Lifshitz, and
Ullman, 2008). lndeed, there is evi dence tha t interference with this "reentrant''
processing can bl ock conscious awareness of the object (Di Lollo, Enns, and
Rensink,2000). (See Web Activity 4.7: Object Substitution Masking.) Theoct
of object recogniti on should be seen as a con ve rsation am ong many pieces of
the brain r.ithe r than as progression from spo ts and lines to th e activation of
a g rand1nother ce ll in inferotemporal cortex.

Before we leave this topic, it is worth mentioning on e very large part of the
story tha t we've omitted up to n ow. There is good eviden ce that atfention to an
object is critical in recogniti on of that object. For exa mple, a sp ecialized piece
of the brain might be responsible for om· ability to recognize a specific face.
If so, tha t piece of the brain probably requires a single face as its i.nput-hvo
eyes, a nose, and a m o uth. If we're looking at a famil y photo, selective a tten-
tion processes must come into play, lest we deliver a collection of a brother 's
eyes, sister 's n ose, a nd \.m.cle's mo uth to the face processor. cells in
different parts of the visu al system may behave different! y before and after
attention is directe d to a n object (Serences and Yantis, 2006). Fo r example, a
receptive field might change its size to wrap itself around a n a tte nded object
(Moran a nd Desim one, 1985). This makes it pa rt of the nehvork processing
tha t object. The same ne uron may be involved in the processing of a different
object w hen attention shifts a m ornent later. Many of the brain s truch ues criti-
cal for the deployment of attention seem to lie in the o ther m ain path coming
o ut of the primary visual cortex-the on e tha t goes toward the pa rie tal lobe.
This p ath way, a nd th e general topic of a ttentio n , is taken up in C ha pte r 7.
Summary
1. A series of ex trastriate visual areas continue the work of visual process-
ing. Emerging from Vl (primary visuaJ cor tex) are h.vo broad streams of Refer to 1he
processing: one going into the temporal lobe and the o ther into the pari- Sensation and Perception
e tal lobe_ The temporal pathway seems specifically concerned with wlm t a C.Ompanlon Website
stimulus might be. TI1is chapter follows that pathway. (The parietal where
sltes.slnauer.com/woHa4e
pa thway v.• ill be considered in later chapters.)
tcr activifies. essays, stu:ty
2. After early visual processes extract basic features from the visual input, it is
the job of middle vision to organize these features intb the regions, s urfaces, questklns. ard other study aids.
and objects that ca n, in tum, serve as input to object recognition and scene-
unders tanding processes.
3. Pen::eptual "rommlttees" serve as an important metaphor in this chapter.
The idea is that many semi-independent processes are \\'orking on the input
at the same time. Different processes may come to different conclusio ns
about the presence of an edge or the relationship be hveen tv.·o elements in
the input. Under most circumstances, we see the single conclusion that the
committees settle upon. Bayesian models are one way to formalize this pro-
cess of finding the most likely explanation for input.
4. Multiple processes seek to carve the input into regions and to define the
edges of those regions, and many rnles are involved in this parsing of the
image. For example, image e lements are likely to group together if they are
similar in color or s hape, if tile)' are near E'f'"h other!_ at. if they are connec t-
ed. Many of these grouping principles we re fi rst articu lated by members of
the Gestalt school.
5. Other, reL.'l.ted processes seek to de termine whether a region is part o f a
foreground figure (li ke this black 0) o r part of the background (like the
white area around the 0). These rules of grouping and figure-ground
assignment are driven by an implicit understanding of the physics of the
world. Thus, events that are very unlikely to happen by chance (e.g., h.¥0
rontours parallel to each o ther) are taken to have m eaning. lThose parallel
contours are likely to bepartof the same figure.)
6. The proce5..ses that divide visual input into objects and background have to
deal with many complexities. Among these are the fad that parts of otJ;ec ts
may be hidden behind other objects (occlusion) and the fuct that objects
themselves have a structure. Is your nose an object or a part of a larger
whole? What about gL'lSses or hair o r a \•dg?
7. In addition to perceiving the shape of objects and their parts, we are also
very adept at categorizing the material that an o bject seems to be made

120 CHAPTER 4
of-glass, stone, doth, and so on.. We use material perception to estimate

physical properties. \.\'hat would it feel like? Can it be grasped like a bottle
or would it s lip through o ur fingers like sand?
8. Template models of object recognition hold that an object in the world is
recognized when its image fits a particular representation in the brain in
the .,.,•ay that a key fits a lock. It has ah,.,1ays been hard to see how naYve tem-
plate models could work, because of the astronomical number of templates
required: we might need one "lock'' for every object in every orientation in
every position in the visual field _
9. Structural models propose tha t objects are recognized by the relationship of
parts. 'Thus, an H could be defined as h.vo parallel lines with a horizontal line
joining them bet\..'"een their centers. A ca t \..·ould be more difficult, but sim i-
lar in principle. In their pure form, such models are vi5vpoint independent.
8230336 Amma App· ll1e o rientation o f the H doesn' t matter. Object recognition, however, is often
viewpoint-dependent, su ggesting that the cor-rect model lies behveen lhe
extremes of naYve template matching and pure s tructural description.
10. Faces are an interesting special case in which viewpoint is very important.
Upright face. are much easier to recognize than inverted faces. Moreover,
some regions o f the brain seem to be specifically interes ted in faces. They lie
neilr regions in the tem poral lobes th<Jt are important for the recognition of
o ther sorts of objects,
11. Recent physiological work showed very sped.fie responses to very specific
objects (e.g., the actress Jennifer Alliston) in the human temporal lobe.
O ther work shm•.'ed that the first, rough acts of object recognition take place
so fast that they must be accomplished by the first, feed-forward sweep of
activity from the retina to the higher process ing centers of the visual sys-
tem. However, routine perception o f objects requires feedback. from higher
visual areas to those lying earlier in the pa th\.,1ay.

ur ' l 14
8230336 Amma Appa

CHAPTER 5
#
I Sean G"'9ns. G\'dB and Tum. 2010
8230336 Ammd Appa

The Perception of Color
WHEN YOU YOUNG, you probably learned various poems that began
"Roses are violets are blue." Jt turns out that this is n ot really true. A poet
with a background in. color perception rnight rewrite it as L•Roseslook red, v io-
lets look blue; the colors you see are really in you." We offer up this mediocre
verse in order to make the point that, as strange as it may seem , color is n ot
a physical property of things in the world; rather, it is a creation of the rn.ind.
Roses look red because we look at them \<\;th o ur particular visual systems.
The central goal of this chapter is to explain what th.at means.
Basic Principles of Color Perception

Although colo r it.sell is no t a. ph ysical property of things in the world, it is re-
lated to a physica l prnpe rty. As discussed in Chapter 2, humans see a narrow
range of the electromagneti.c spectrum behveen the wavelengths o f about 400
and 700 nanometers (nm). The apparent color of a piece of the visible \vodd
is correlated with the waveleng ths of the light rays reaching the eye from that
bit of the world.
Most of the light U1at we see is reflected light. Typical light sources, like
theslm or a lightbulb, emit a broad spectrum of '"''avelengths that hit s urfaces
in the world aroLmd us. Some wavelengths are absorbed by the s urfaces th ey
hit. The more light that is absorbed, the darker the s W'face will appear. Other
wavelengths are reflected, and some of that reflected light reaches the eyes. The
color of a surface depends on the mix of that reach the eye from
the s urface (an d , as we will see later, fr om the other surfaces and lights that a.re
present in the scer1e). L1 the case of roses, m ore of the longer-wavelength light
(>600 run) is reflected into the eyes of the Even though (almost) all
humans would declare this of wavelengths to appear red, it would be
a big mi.stake to think of specific wavelengths of li ght as being specific colors.
As Steven Shevell (2003) puts it, "There is no red in a 700 run light, just as there
is no pain in the hooves of a kicking h orse.'' Llke pain, color is the result of the
interaction of a physical stimulus \vi th a particular ne rvous system.
Three Steps to Color Perception

Several problem s must be solved in order to go frorn physical wavelengths
to the perception of color, We will organize our discussion around three s teps
(Stockman and Brainard, 2010):
1. Detection . \ Vavelengths must be detected
2. Discrimination . We must be able to telJ the difference between on e
wavelength (or mixture of wavelengths) and another.

124 CHAPTER 5
FIGURE 5.1 The retina contains fou r types o f photo-

receptors. These differ In their sensitivity to the W<Ne- 4al 498 sas 565
lengths of light. Thr99 .COnQS are maxima.Hy sensitivei at • 100
short {S}, medium (M}, and long (l.) wavieleingths. The
.0
si ngle type of rod phota909ptor has its peak sensltMty
between those of the S - and M-conGs.
] 50
S-cone A cone thatls preferentially

sensitive to short wavelengths; col-
loq uially (bUt not entirely accurately)
known as a '"blue oone."
M-cone A oone that Is preferentlaHy 400 500 600 700
seosltlve to middle wavelengths; col - Wavelength (run)
loquially (but not entirely accurately)
known as a "green cone."
L -cone A oone that Is preferentlaly
sensitive to long wavelengths; cdloqul- 3. Appearance. We want to as.sign perceived colors to lights and surfaces
ally (but oot entirely aocuratel)') known in the world. Moreover? we want those perc::eived. colors to go with
as a •roo cone." the object (that rose looks red) a nd not to change dramatically as the
spec1ral sensitivity Refarln;i to the viewing conditions change (that rose sho uld remain red in sun and
seosltlvlty of a oell cr a device to differ- shad ow, for example).
ent wavelengths on the electromag-
netic spectrum.
photoplc Referring to light innensl-
Step 1 : Color Detection
tles that are bright enough to stlrru- Detection was largely covered in Chap ter 2 To briefly review, we have three
late the oone receptcrs and bright types of cone photoreceptors. These cones differ in the photopigment they carry,
ena.igh to "saturate" the rod receptcrs
and as a result, they differ in their sensitivity to light of different wavelengths.
(that Is. drive them to their maximum
resp;yises). Figure 5 .1 s hows those sensi tivities as a function of wavelength. Each cone
type is named fo r the location of the peak of its sensitivity on the spectrum:
scotoplc Referring to lght lnnensltles
that are brght enough to stimulate the The con es tha t have a peak at about 420 run are known as sh ort-wavelength
rod receptors but too dim to stimulate cones {S-cones for sh or t). The medium-wave.length cones (M-cones) peak at
the oone receptors. about 535 run, and long-wavelength cones peak a t about 565 run.
As you can see from Figure 5.t cones are n ot exclusively sensitive to different
parts of the spectrum. That is? even though the L-cone is maximally sensitive
at about 565 nm, the M-cone can detect that wavelength as well. TI1eir spec-
tral sensitivities overlap. As mentioned in C h apter 2? S-cones are relatively
rare, a nd they are less sensitive than M- and L-cones. The combination of
sensitivities of the three types o f cones gives us our overall ability to detect
wavelen gths from about400 nm to about 700 nm (see Figure 5.1). Remember
also that cones work a t daylight (photoplc) light levels. HI, have one type of
rod. photoreceptorj it works in dimmer (scotopk:) light and has a somewhat
different sensitivity p rofile, peaking at about 500 nm.
Step 2: Color Discrimination

500 700 The Principle of Univariance
Wavelength (nm)
We can detect wavelengths between 400 and 700 nm, but how do we distinguish,
FIGURE S.2 A sin1Jle photorecep- for exa mple, lights of 450, 550, and 625 nm? To see how discrimination differs
tor shows different responses to lght:s from detection, let's examine the response of a. single photoreceptor to a single
of dlfferoot wavelengths rut the SarnQ wavelength of light Figu m 5.2 sh ows h ow one kind of human photoreceptor
intens ity. A 626-nm light of this Inten-
sity, indicated by the arrow, produves responds to light of a specific wavelength while the intensity of the light is held
a response midway between the maxi- Because of the properties of the photopigment in the pho toreceptor
mum and minimum responS9S. cell, 400-nm light produces only a small response in each c:ell of this type, 500-nm

8230336 Amma Appa THE PERCEPTION OF COLOR 125
light produces a greater response, and 550-nm light even more. However, 600-
nm light p roduces less than the maximal response, and 650-nm light produces
a minimal response. Light of 625 run produces a response of moderate strength.
So fur, so good. We know rl>at different wavelengths of light give rise to different
experiences of color, and the varying responses of this photoreceptor to different
wavelengths could prov ide a basis for color vision. But there is a problem, as
\
illustrated in Figure 5.3. Supp C16e we change the wavelength from 625 nm to 450
nm. Figure 5.3 shows that an equal amount of 450-nm light will produce the same
response from this pho toreceptor that 625-nm light does. If we were looking at the
output of the photoreceptor, we would h ave no way of distinguishing ber.-.·een
the two lights. But when '.ve look Mth a norrnal human color vision system, the
625-nm light looks orange and the 450-nm light looks bluish. 500
Actually, the p roblem is much worse. Remember tha t Figures 5.2 a nd 5.3 Wavelength (nm)
represent the photoreceptor's response ra te when all wavelengths are presented
FIGURES.3 Ughts o f 460and 625
at the same intensity. Under these conditions, the graphs indicate that light at
nm each €1llclt the sarnQ r9Sponse from
either450 or 625 run produces a response lower than the peak response obtained the photoreceptor w hose responses
at about 535 nm. [f we had a 535-nm light, we could reduce its intensity until are shown here and In Figure 6.2.
it produced. exactly the same level of response from our photoreceptor as the This situation illustrates th€1 prlnciplQ o f
45(}. or 625-nm light did a t the higher intensity. Indeed, we could take a "white univarlanoo.
light" or any mix of wavelengths and, by properly adjusting the intensity, get
exactly the same response out of the photoreceptor.
Thus, when it comes to seeing color, the output of a single photoreceptor
is completely ambiguous. An infinite set of diffe_rent wavelength-intensity
combinations can elicit exactly the same response, so the o utput of a single
photoreceptor cannot by itself tell us anything abou t the wavelengths stimu-
lating it. Titls constraint is known as the prlnclple of unlvarlance (Rushton,
1972). (See Web Activity 5.1: The Principle of Unlvarlance.)
Ob\oiously the human visual system has solved the problem, but not under
all circumstances. Uni variance explains the lack of color in dimly lit scenes.
Remember that there is only one type of rod photorecepto r. All rods contain
the same type of photopigment molecule: rhodopsin. Thus, they all have the prlnclple of uni variance The
same sensitivity to wavelength As a consequence, although it is possible to tell fact that an Infinite set of different
light from dark under scotopic condi tions, the problem of un.ivariance makes w""'"<ngth-lntanslty combinations can
ellclt exactly the same response fran
it impossible to discriminate colors. O ur nighttime color blindness is one hint aslngle type of photoreceptor. One
that color is psychophysical and n ot physical The world seen t.U1der a bright photoreceptcr type cannot make ooor
moon (Figure 5.4) has not been physically drained of col or. The same mix of discriminations based on wavelength.
f=IGURE 5.4 The moonlit wor1d appears drained

of cdor t>oc.ause we have only one type of rod
J::hotoreceptor transducing light under these sco-
topic cond itions. With just one t)'pe of photore-
C€1ptor, wa cannot make discriminations based on
wav&IQrigth, so we cannot SQQ oolor.

126 CHAPTER 5
Wavelength (nm)
FIGURE 5.5 The two wa.wlmgths that produce the samG response from one type.
o f oonG (M) produce dlff9r9rlt pattgrns o f rQSponses across thei three tyPQs of ex>rl9S
(S, M. and L).
wavelengths that produces color perception dtll'ing the day remains present
o n that moonlit night, but we fail to see colors under dim illuminants like
moonlight, because dim light s timulates only the rods, and the output of that
sin gle variety of photoreceptor d oes not permit color vision.
The Trichromatic Solution

We am de tect differences between waveleng ths o r mixttues of wavelengths
precisely because we h ave more than one kind of con e photoreceptor. Figure
5.5 s hows how, with o ur three cone types, we can tell the difference behveen
lights of different wavelengths. Loo.k at the three cones' responses to the two
wavelengths-450 and 625 nm-that produced the same respon se from the
single cone in Figure 5.3. TI1e two wavelengths of light still produce the same
response from tha t type of cone, now revealed to be the M-cone. However,
these two wavelengths produce different outpu ts from the L-cone.s a nd S-
con es. In fact, as you will see if you try Web Acttvlty 5.2: Trtchromacy, a ny
wavelength from about 420 to 660 nn1 produces a unique set of three responses
from the three con e types. Th.is combined signal, a triplet of numbers for each
"pixel" in the visual field, can be u sed as the basis for color vision . (We can
see from abou t 400 to 700 nm, but the very long and very s hort wavelengths
ead1 stimulate only one type of cone.)
In our discussion of the uni variance proble m, we noted that we can make
any wavelength produce the same response as a ny other from a s ingle cone
type by adjusting the intensity of the light. That doesn't happen in the three--
cone_world o f human colo r vision. A sped.fie light produces a sp ecifi c set of
responses from the three cone types. Suppose that the light produces ti.vice as
much M response as S response a nd twice as much S resp on se as L response.
trtchromauc theory of color Vision If we increase the intensity of the light, the response sizes will change but
a trtchrornacy The theory that the relationships will not. There will s till be twice as much M response as S
the colcr of any Ught Is defined In our response and twice as much S response as Lresponse; and those relationships
visual system by the relationships of
will define our response to the light and, eventually. the color that we see. The
three numbers-the ootputs of three
receptor types now known to be the idea that abili ty to d iscriminate one light fro m another is defined. in otu visual
three cones. Also called the Young- system by the relationships among three numbers is the heart of b1chromacy
Hstnhollz or, more elaborately.. the trichromatic theory of color vision.

THE PERCEPTION OF COLOR 127
FIGURE 5.6 Objects in the real vvorld reflect light

across the spectt\Jm in diffenmt arnet.ints, Thls graph 60
plots the reflectances o f raw and cooked hamburger
meat. Vtrnich one boks redder? \Miich one reflects
m ore of tt'"l9 long wavelengths?
40
Metamers
The examples p resented thus far involve the re-
sp onses of the visual system to single wavelengths.
Howeverr y·ou m ay have noticed tha t we keep refer-
20
ring to uw ..n:elengths or rn.ixtures of wavelengths."
That's because \o\'e are not typically exposed to sing le
wilvelengths . Almost every light and every smface
tha t we see is e mittin g or refl ecting a wide range Cooke.:I
of wavelengths . A laser pointer would emit a very
n arrow range o f wavele ng ths, but a m o re nonn al 400 450 500 55() 650 700
si tuation is shovvn in F19ure 5 .6. [t sh mvs the rela- w.welength (run)
tiveam m mtsof light reflected from raw and cooked
hamburger. How can we di.scrim_inate the raw fro m the cooked? When \.ve're
stud ying color vision, this rea l-world con cern gets red uced to a different ques-
tion.: H ov.1 do our cones respond to combinations of wavelengths of light?
To answer this question, consid er w hat h appens if we mix just two wave-
length s. For the sake of this examp le, we will over.simplify by ignoring the
S-<ones and red r<"twi ng the M- and L-cones to make the numbers s impler.
Imagine that we .s hine a wavelength that looks red and a wavelength that
looks green onto a w hite piece of paper so tha t a mixture o f both is reflected
b,,ck to cl>e eyes( Fig...-e5.7a ). Suppose tha t the light cl>at looks green produces
80 tmits of activi ty in the M-cones and 40 in the L-cones {remember, we are
ignoring the S-ames fo r n ow). Jn addition, s uppose that the lig ht that looks
red produces 40 units of activity in the M-cones a nd 80 in the L-cones. If
we assume that we c.an add the cone responses together, then the
"red ,, and "green '' lights produces a resp onse of 120 un'its in ead"t cone. TI1e a
ab.solute value is not importa nt, because it could change if the intens ity o f the
light changes. What is hnportant is these hvo ligh ts, mixed together, produce
a mixture that excites the L- and M.-cones equally.
The key point is that the rest of the nervous system knows 011/y what the
cones tell it. lf the mixture oflightsthat lookredand green prod uces the same
cone output as the single wavelength of light that looks yellow (Fig ure 5.7b ),
(a) Whathapperuif you addthi.slight (b) Th is light that looks yellow pra..iuces
that loo ks red to o ne th.:\t looks green r equal L- <ind M-cone !'esponses.
M-cone -------f'Y\-------11- L-cone

FIGURE 5.7 Jn (a) 1 the loog-wave-
// \ \ l9ngth light that looks ard th9

sho1ter-wav9length Ilg ht that looks
I/ \\
I
I
\
\
\
gr991i mix togeth9r to produ::::9 the
same r9spcnse 1rom th9 cones as
does the m9dlum-wawlmgth Mght that
looks yellow in fP). If two &et s of lights
,,/
y
'-' produce th9 s3me respon:ses, th9Y are
G R rnetamers and must bok identical. so
Wavelength Wave length thg red plus the gro99n w ill look yellow.

128 CHAPTER 5
metamers Dlfferoot mixtures of the n the rnixture and the si ngle wavelength m11st fook identical. M ixtures of
wavelengths that look Identical. More different waveleng ths that look identical are cal led metamers. The si ng le
generally, any pair of stlrnuli that are thatproducesequal M-and L-coneactivity will look yellow, and the
perooved as Identical In spite of physi-
c al differences. correct m ix ture of longe r- and shorter-wavelength lights wiJl also look yellow.
Two quick warnin gs:
1. Mixing wavelengths d oes n ot chan ge the physical wavelengths. If we
m ix 500- and 600-nm lights, the physical s timulus contains 500 and
600 nm. It d oes no t contain the average (550 nm). It d oes not contain
the s um (1100 mn) (w hich we would not be able to see anyway). Color
mi.xture is a m e ntal event, no t a change in the p h ysics of light.
2 For a mixture of a red and green to look p erfectly yellow, we would
have to have just the right red and jus t the right green. Othe r mixes
n"tight look a bit reddish or a bit greenish.
1l"tis example generalizes to any mixtu re of lights. ..\I.I the light reaching the
reti na fro m one patch in the visua l Held will be converted into three numbers
by the three cone types. If those numbers are s ufficiently different fr om the
numbers in ano ther pa tch, you wiU be able to discrimi nate those patches.
Ii no t, those patches will be metamers. They w ill look identica t even if the
""·avelengths are physica lly different.
The History of Trichromatic Theory

From \Vhat you ' ve read so far in this book, you would be forgiven fo r s uppos-
ing th.:i.t clever anatomists a nd physiologists identified the three cone types
and built the tridrromatic theory of color vision from there. lndeed, there have
been beautiful experiments of this sort: For in.s tance, Schnapf, Kraf t, and Bay-
lor (1987) ma na ged to record the activity of single photoreceptors. Nathans,
Thomas, and H ogness (1986) found the genes that code for the different phot-
opigments. David WilLiams a nd his s tudents even developed. n method fo r
photographing and identifying different cone types in the living hum ..·u\ eye
(see Figure 2.1 2 in Chapter 2).
Such research has ceme nted om of the physica l basis of
trichromacy, but the basic theory was established by psychop hysica l experi-
mentation, and theorizing started \vi th Isaac Nelo\'tor{s great discovery tha t a
prism would bre..<i k up sun light into the sp ectrum of hues, and a second prism
would put the spectrum back together into light that looked white. Jn 1666,
Newton understood that "the rays to speak properly a re not coloured.. In them
is no thing else than a certa in Power and Dispasition to s tir up a Sensation of
this or tha t Colom '' (from Opticks, published originally in 1704). Newton knew
that color is a mental event.
TI1e th ree-dimensional nature of the experience of color was worked o ut
in the TI'?"'"' Yow1g (1773,-1829) and subsequently by
He rmann von Helmholtz (1821-19J4).Jn their honor, trichromatic theory is
often call ed tl1e "Young-Helmholtz theory." James Clerk Maxwell (1831-1879)
developed a co lor-matching tedmique that was central to Helmholtz's work
on this topic. (SomehO\v Maxwell missed having his name nttached, or we
would have the Young-tvlaxweU-Helmholtz theory.) Ma xwell's technique
is illus trated in Fig ure 5.8. 1l1e o bserver in a mod ern vers io n of Maxwell' s
experiments would try to u.se different a mounts of "prim ary" colo red lights
(e.g., the lights looki ng red, green, and blue on the right side of the figure) to
exactly match another reference color (e.g., the Ught lookin g cp.m or b luish
on the left si de).
T11e central observa tion from these experiments was tha t only three m ix-
ing ligh ts are needed to match any reference lig ht. Two prim nries are no t

FIG UR E 5.8 In a modi;irn version o f Maxwell' s color·

matching experitn9nt. a color is presented on the left. On
the right, the obse1Ver adjusts a mixture of the three lights
to m at ch thQ oolor on the l€1ft.
enough. Four are more than are n eeded . Lon g before physiology could prove ack:flUve color mlxtl.re A mixture
it, these results led Young and Hel mholtz to deduce tha t three different color of If light A and llgl1t B are both
m echanisms must limit the huma n experien ce of coloJ·. reflected from a sutfaoo to the f1'16, In
the p erception of color the effects of
th:::<le t wo lights add togetr1er.
A Brief Digression into Lights, Rlters, and Finger Paints
The ubiquity of televisions and compute r m onito rs in the h ven ty-fi rs t cen ht ry subtractive color mixture A mix·
ture of If pt)ments A and B
m ay make colo r 1nixing and m e tam e rs reason ably intuitive. lf yo u've n ever mix. some of the light shining on the
d one so, fin d a magnifying glass and take a very close look a t a yel low patch surface v.;11 be subtracted by A, and
on yotu comp uter screen .. You'll find that the patch is actually composed of some by B. Only the rem ainder con-
tho usa nds of inte rmixed red a nd g reen d ots. The 11 red + g ree n =yell ow" tributes to the pa-ceptlrn o f cobr.
for mula is a n ex.ample of additive color mix:b.Jre because '"te are taking one
\"tavelen gth or set of W'1 velen gths and addi,,g it to a nother.
For most of us/ color mixhue begins in kindergarten or earlier, with paints.
In that world/ red p lus green doesn 't make yellow; th.at mixture typ ically loo ks
b rovvn . A finger paint/ or any other pigment, looks a
particular color bee-a use it absorbs some wavelengths,
subtractin g them fro m the broadband ("white") li ght
falling on a su rface covered w lth the pigm ent. \ \.l,en 1. T.lke "white" light
that oontnin.s a broad
a tod dler s mears together red a nd green/ alm ost a ll mixture of wavelength.s .
wavelen g ths are absorbe d by on e pi gm ent or the
o tht>.r1 e per lve., the subtractive color mixture
a dar k color Ii e b rown.
Actua lly, finger pain t mixtures are ra the r com-
plicat&·t with some pigment pa rticles sitting next to
each and effectively add ing their reflected Ught 2. Pass ii through a filt er
that absorbs Shorter
to the result. O ther particles occlude each other, a nd w.w.d.eng ths. Th<t" result
still others are engaged in o ther complex interactions. will look yellowish.
Colored filters, like those you m ight put over stage
lights, are a cleaner example o f subtr.1cti ve color m ix-
ture. Figure 5 .9 shows h ow a s ubtractive m ixture of
yellow a nd blue filters would subtract wavelen gths,
leavi ng only wavelengths in a middle range, which 3. Pass that through a
bluish filter that absorbs
appear green . An add iti ve m ixture of lights that look all but a middle rnnge
ofw,w elength.s.
FIG URE 5.9 In this example of subtractive color mixture.

•vvtiite• -broadband-light Is passed through tvvo fil ters.
The first one absorbs shorter vvavelengths, transmitting a
mix of WCP1elengths tt1at looks yeUow. The second absorbs
4. The w,w eleng:ths that
lo nger and the shortest wavelengths, trans· make it through OOth
rnittlng a mix that looks blue. The wavelength s that can filt ers will be a m ix
pass through both filters without b eing subtracted are a that looks greenish .
midd le rangg of wavelQ("Qths that appear gr9en.

130 CHAPTER 5
FIGURE S.10 If we shine a light that looks blue and a liight that looks yellow on the
same patch of papgr, the wavelengths will add. producing an additive color mixture.
RarTIQfnber that the light that loci-<£ yQ!low is 9quivalent to a mix. of a long wa\IQ!Qngth
and a medium wa\19length, so blue plus yellow results in a mix of shCft, medium. and
long wavelS"lgths. The looks w hite (or gray, if it is no t th9 bright est patch In
,;ow1.
blue and yellow will look \.Vhite {if you have exactly the right blue and yellow)
(Figum 5.10) bec..,use that combination produces a mix of wavelengths that
stimulate tl1e three cones roughly equally. (See Web Activity 5.3: Color Mixing.)
Additive color mixturewitl1 paints is possible. Georges Seurat (1859-1891)
and other Po.stimpressionist artists of the late nineteenth century experimented
with Pointillism, a style of pain ting that involved creating many hues by placing
small spots of just a few colors in different textures (Ftgure 5 .11 ). Viewi ng the
painting up d ose, as illustra ted in the fi gu re, we can see each individual d ot
of color. Like the red, green, and blue phosphor dots on a compute r m onitor,
the d ots in the painting combine additively to produce a wide rnnge o f colors.
Thus, fro n-1 a dis tance the water 's s urface is appropriately s il very grny even
if, up close, it is composed of spots of b lue and yellow that would n ot be seen
on the s urface of a (dean) harbor.
From Retina to Brain: Repackaging the Information

11"1e cones in retina are the ne ur al subs tra te for detection o f lights. \ Vh a t is
the ne ural basis for discriminating between lights \vi th different wavelength
composition? To tell the diffe·rence between different lights, the ne rvous sys-
tem \vill look at differences in the activities of the three cone types. This work
begins in the retina. \'Ve could send separate L, M, and S signals to the brain,
but tha t approach would be less useful than one might think. For example, the
L- a nd M-cones h ave very sirnibr sensitivities (see Figme 5.1 }, so most of
FIGURES.11 Pointillism is usually

il!ustratoct with a picture by Seurat. For
variety, here is Paul Signac(1863-1935)
using additlw oolor mixture in the m anner
of a m odern ccrnputer monitor. See how
small spot s of blue and yellow paint blend
at a distance to give rise to the sitvery gray
of the 'Water's surfaoa.

time they are in d ose agreement: L says, "Lots of light coming from location
X.11 uYes, lots of light coming from loca tion X/' M agrees.
Computing diffe rences behveen cone responses turns out to be a muc h
mo re usehil way to transmit informati on to the brain. The n ervou s systern
computes two differences: (L -M) and ([L+ M] - 5). The difference between L
and M responses contains conside rable info mu'ltion about color. One speculative
idea is that (L- M ) differe nces are particularly w eU suited to appreciating the
differences between different a m ow1 ts of blood in s kin. Blus hing and huning
pale are use ful signals to observe, and our specifi c photopigm ents may have
evolved to he lp us see those signa ls (Ch angizi, Zhang , and Sh.i mojo, 2006).
Th a t is no t the only possibility. The (L-M) difference may also be very u seful
if yo u w ant to tell the d iffere n ce be tween fruit a nd leaves o r different s hades
of green in the folia ge (B. C. Regan et al ., 2001 ).
In addition to (L - M ), we could ereate (L - S) and (M - SJ signals. H owever,
because L and M a re .so similar, a s ing le comparison behveen Sand (L + 1V1)
can cap hue almost the same infommtion tha t w ould be found in (L - S) and
(M- S) signals . Finally, combirUng L a nd f\,1 s igna ls is a pretty good measure
of the intensity of the light (S..cones ma ke a rathe r sm all contribution to our
perception of brighb1ess). Thus, on theoretical grounds, it might be wise to
conve rt the three con e si gnals into th ree new s igna ls-(L- M ), ([L + M] - S),
and (l + M) (Buch$baum and Gottschalk, 1983; Za idi, 1997}-and the visu al
system d oes something reason ably d ose to this.
Cone-Opponent Cells in the Retina and LGN

The ea rliest work on the co mbina ti on of cone s ignals was d one with fish
(Svae tich..i and ,.Macnichol1 1 59). By the 1960s, Russell De Va lois and oth-
ers had m tOsh<n · th t these .E?orts of s ign als actuall y exist in th e lateral
genlculate nucleus (LGN) of m acaque m onkeys (De Valois, and
Jacobs, 1966). As d escribed in C h.a p ter 3, man y gang lio n ce lls in the retina and
the LGN of the tha lam tLS are maximally s timulated by sp ots of light. 1l1ese
cells h ave receptive fie lds w ith a charac teristiccen ter--surround organization.
lateral genlculate nucleus
For exa mple, some cells are e xcited w he n a Light turns on in the central pa rt (l.GN) A structure in the thalamus,
of their receptive field s and inhibited w·he n a light turns on in the s urro und part of the midbraln, tl1at receives Input
(see Figure 2.14 in Chapter 2). fran the retinal gangllm cells and has
A s imilar antagoni stic rela ti ons hip c haracte rizes color. So me of th ese Input ard output coonectlons to the
re tinal and LGN ga nglion cel ls are excited by the L-con e on set in the ir center visual cortex.
and inhibited by M-con e onsets in their s urrotmd . These (L- M) ceUs are one cone-opponent cell A cell type -
type of cone--opponent cell, so narned beea u se different sources of chromatic foun::i Jn the retina lateral genlculate
and visual oortex- that, ln
info rma tion are pitted again st each other. There a re also (M-L), ([M + L]- S), effect subtracts one type of cone
and (S - [M + L)) cells-just the sorts of cells we would like to have to s up- Input fran another.
p ort the repackag ing of cone s igna ls, d escribed in the prev io us section. TI1e
konlocellular Referring to cells In
cells that we re excited by light onset could be thoug ht of as (L + M) cell s. the konlocellular layer of the lateral
Thus , we have the three signal s that we w anted on theoretical grounds . 1l1e geniculate nucleus of the thalamus.
actual physiology is quite complica ted. As rnentioned in C h apters 2 and .31 for Kon/o the Greek for ' dust " refer-
e xample, the S-cone signals go through the konlocellular layers in the LGN ring to the appearance of the cells.
w hile the M- and L-con e oppone nt s igna ls are mostly fotmd in the parvocellular Referrlrg to oolls In
lular layers (Xiao, 2014). the palVOCellular layers of the lateral
geniculate nucleus of the thalamus.
A Different Ganglion Cell Helps to Keep Track of Day and Night P EWO from the Greek for "small" refer·
ring to the size of the cells.
If you have ever fl ovvn a significant d istance east or west, you hsxe probably
experien ced jet lag. The time on your ce ll phone d ock d 1a.nged by a felv h ours clrcadan Referring to cells In the
magnocellular layers of the lateral
one way or the other, but your internal dock didn' t ge t the mes.sage and re- genlculate nucleus of the thalamus .
ma ined on yo ur home time. Over a co uple o f days, you adjusted to the n e\<\o· f.Aagno frcm the Greek fa "large"
time. The primary force adju s ting your intern<ll circadian d ock is s unlight referring to the size of the cells .

132 CHAPTER 5
melanopsin A photcplgment, fcur<l {LeGates, Fernandez, and 2014). Inte restingl y, mice bred to have no
In a class of photoreceptlve retlnaJ hmctioning photoreceptors continued to entrain their circadian d ocks to light.
ganglion cells.
Some apparently blind hum ans also seein to have that ability {Czeisler e t al.,
color space The three·dirnenslcnal 1995i Klerman et al., 2002). How could this be? There has been speculation
space, established because color about photoreceptors in the s kin, bu t in recent years, we have learned that
perception Is based on the outputs of
three cone types, that d esc nbes the there is a previously unknovvn pho tosensitive cell in the re tina It is a gan glion
set o f all colors. cell . It receives input fro m the rods and the cones, but it also contains its mvn
photopigment, melanopsln, so it ca n de tect light even when norma l photo-
receptors are absent. You do n' t see the outputs of this cell. Its outputs go to
centers in the brain that control flmctions like your circadian clock. It is as if
two (or m ore) visual systems were occupying your body, sharing the same eyes.
Indeed , under the rig ht circums tances, a few seconds of blue light can have
an itnpact on cognitive fun ction , even in individuals w ho have no con scious
light perception via th is m elanopsin-based system (Vandewalle et a l., 2013).
Step 3: Color Appearance

Re huning to the visu..1.1 sys te m tha t is giving r ise to our conscious perception
of the visual world, we have seen that the three cones detec t a range of wave--
lengths . Rods m ake a sm .1.U,
. important contribution to color vision, hl1t only
in fairly dim lig ht (Stabell and Stabe U, 2002). The retinaand LGN contain cells
tha t have repackaged tha t in.formation into cone-opponent differen ce signals
tha t constrain our a bility to see differences beh·veen regions. Now is the time
to think about '"1 ha t colors w ill be perceived.
Three Numbers, Many Colors

Pirst, it is useful to have a sys te m fo r talking about all the colors we c m see.
Because we ha ve exac tl y three different types o f con e photoreceptors, th e
light reac hing any part of the retina wi ll be trans lated into three responses,
one for each local population of cones. After that tran s la tion, the res t of
nervous system cann ot g lean a nything more ab out the physical wavelengths
of the light [f the light rays reflecting o ff two surfa ces produce the same set of
cone responses, the hvo stU'fu ces mus t a nd wi ll appear to be exactly the sam e
color. They will be me tame rs, even if their physical characteristics are quite
different. Thus, it is possible to p rodu ce blood.red color on the page wi tho ut
823033£\ Amma Appa
mimi ckin g the physical properties of blood.
\Vo rking with just three nmnbers might not sound very prom ising. but it
has been estimated tha t, with this system, we can d iscrimin;:ite the s urfaces of
more than 2 milli on different colors (Linha res, Pinto, and Nascimento, 2008;
Pointer and Attridge, 1998). A lot of these are lighh1ess variations of w hat ""'e
would coll oquially con sid er the sa me color: b loodred in dim li ght, blood.red
in brighter light, a nd so on. But even if we ignore lightness, we can still dis-
ting uish about 26,000 colors (P-oster, 2011). Unless you ' re in paint sales, you
d on 't know this man y distincti ve colo r nnmes, but you could tell two differen t
co lors apart even if they were both named "pea green'' or "sky blue .11
\Ve n eed some way to talk about all those colors in a n o rde rly way. We
rnn describe each of the colors in th e sp ectrum with a single number-for
exa mple, the light's wavelength. Going beyond the sp ectrum, \Ve have a th.ree-
dimensional color space. The three dimensions s tart with the three numbers
that come from the outputs of the three con e types. The three-din'leJ1Sional
space is a nalogous to the height, le ngthr a nd width of a cube. O f course, it
d oesn ' t make much sense to talk about height, lengt11, a nd width in color
space, bu t the space tha t contains the set of all perceptible colors wi ll be a
three-dimens ional space.

THE PERCEPTION OF OOLOR 133
Picking Colors
Representing that three-dlmenslaial oolor space becanes a practlcal
problem when you am using your computer. Many applcations let you
pick the colors of fonts, objects, and so on, and the app developE<S
needed to devise an Interface that would make that p-actlcal. One sdu-
tlon has teen simply to present a !xed set of options (e.g .. the crayon-
box cdor-picklng tool In many appllcatlcns). However, If the developers
want to p!Wde access to all thl pcsslt>e oclors, then they can pres-
ent cne or more oclor pickers that represent a th ree-dimensional color
space. For example, look at Figure 5.12. It shows two different ways of
uslrg three "sliders" to define oolors (here, borrowed fn:rn Po1NerPoint).
In Figure 5.12a are throo sliders for red. grem, and blue. These ffiB
sliders are useful t.ecause ocmputer mcnltors have red , green. and blue
elemmts In each pixel , and using RGB values is a good W"f to specify
the signal to send to ea.ch of ttose elements. Thus, the green stown In
the figure is what you get from 62 units (out of 255) delivered by the red
elemmts, 180 from the green, and 60from the blue. These RGB sliders
define a three-dimensional space that can be rep-esented by the cube FIGURE 5.12 A oolor picker may offer
shown In Figure 5.12c. Notice that whte Is in the upper ocmer closest to several 'Nays to specify a color in a three-
you on the cube. Black Is nidden on the back comer, farthest frcrn you. dlmaisional ookJr 9PaoQ.l These ..sliders"
In Figure 5.12b, we soothe same shade of groon defined by a frcrn Mcrosoft are based on
an RGB (red, g r9'!n, blue) set of pri maries
dlffernnt set of three numbers. This time , the terms are hue, satumtbn,
and brlgfltness. Part of the resulting HSB odor space Is shown in Figure In '8>and on HSB t>Je, saturation. brlght-
1'19SS) In (b). s... the text f()( dotails.
5. 12d. These drnenslons are very
dlffEfent concepts from red , groon,
and blue, but notice that again the (a) (c)
R.d
system Is three-dimensional. Look 0 0 0 co1o ... 0
at the circu lar. top surface of the
HSB spaces . Hue Is the chromatic
aspect of a light. It changes as you
move around the clrcumfa-ence of
the circle. Each point on the spec- bij j RGB Slid"'' 1:1
trum defines a different hue. As you
can see ti'crn the HSB sliders, the Red
whole family of colors with a hue !!;;Ct 1 ;;;;;;::==:Ji II§]
;;;ci;;;
Graen
of 120 (out of 360 degrees around
the circle) will look like sa-ne version liiiiiiiiiliiiiCiiiiiil
Blue
of green. The saturation dimension
corresponds to the amount of hue ·-!!!!!1]!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! 0
present in a light. In the HSB space
(Figure 5. 12<1), It Is shown as the (b) (d)
distance from the center of the cir-
cle. At the center, where saturation
o.
Is the resUltlng oclor would ap-
pear somewhere en the achromatic
axis from black to w hite. Here, '57
(out of 100) descr1bes a fairly satu- Uj/HSBSlide" 1:1
rated green. The final term. br1ght- Hue
ness , describes that black-white
achromatic axis. The black bottom Saturation
of the HSB cone is a brightness of O
(out of 100). The space tapers to a
point because when br1ghtness Is o.
hue and saturation no longer have

134 CHAPTER 5
meaning. The full three-dimensional space, in this case , would include

a c one, lislng to a white point at a brightness of 100. Slightness Is tne
perceptual consequence of the physical Intensity of a light. For Instance,
the physically intense light of the sun looks brighter than the less intense
moon.
Thus , when you use your color picKer to find just the light cdor,
you are Illustrating the fundamental nature of tlichromacy. There is one
Interesting exceptkm. Aloog v.1th RGB and HSB color many
applications will also give you CMYK. This ls a four-dlmensional system,
v.ith sliders for cyan. magenta. yellow, and black (K because B would
confuse it v.1th B for blUe). You might notice that these are the colors
of thl lnK cartlidges In your color plinter. Color plintlng Is a subtractive
process and CMYK Is designed for plintlng. The black slkJer Is a conve-
nience because using black inK is a better way to get to the black point
In your cdor space than mixing cyan, magenta and yellow inks. We will
leave it as an exercise for the reader to figure out why CMYK uses C, M,
and Y rather than R, G, and B Inks.
The Limits of the Rainbow

The spectrum, that you can see in the "hue" slider is like the w avelength
spectrum that you can see at the bottom of Figure 5.13a, but it is not identi-
cal. There are hues that you can see that do not exist on the wavelength spec-
trum. Figure 5.1 3 illustrates how this comes to be. For example, suppose we
combined a pure 420-nm lig ht with a pure 680-nm light. Thi s combination
would strong ly s timulate the L- and S-cones and produce m..inirnal s timula-
tion in the M-cones. N o single wavelength of light co uld do that, but such a
mixture is visible and must look like something. In fact, such mixtures will
produce purplish magentas that seem to lie naturally behveen red and blue
on a color circle (Figure 5 .13b }. When we vvrap the waveleng th spectnrm into
(b} The color circle
50{)
8230336 AmMa Ap
400 450 500 550 600
Wa,relength (run)
FIGURES.13 lf youcomblne lghts
that look rad a nd bk.Je, the result w(g
No single wavelength
look purp6e, but there ts no on on the spectrum will
the spectrum '8). Purples are • non- have the hue of this
spectraJ c obrs· that join the ends of the mixture of a long and
sp9cirum Into a oolor circle (b). a .s hort wav elength

a color ci rcle, we join the red and blue en ds '""ith a set of colors that are called
''nonspectral hues" -hues that can arise only from m ixhues of wavelengths.
While we are on the topic, it is worth noting that there are cormnonl y perceived
colors that are n ot included in the spectrum's "all the colors of the rainbow."
Brmvn is on e s u ch color. There are no brown lig hts. Brown is seen when a
m.ixh.rre of waveleng ths that would look yell mv, greenish yel low, or orange
is seen in the compan y o f other, brighter patd1es of color. Yoll cannot see an
isolated brow n light in the dark.
Opponent Colors
In the nineteenth renhtry, Ewald Hering (1834-1918) described a curious feature opponent color theory The theay
of color vision: some combinations of colors are illegal. We can have a bluish that perception of cda Is based en
green1 a redd ish yellow (which we would ca1l 1'orange''), or a bluish red (which 1110 output of three mechanisms. each
of them resultlrg frcrn an opponency
we would call "purple" ), but reddish green and bluish yellow have n ever been betvJOOn WK> colors: red-green. blue·
seen, because they d on 1t exist. Red. and green are, in some fas hion, opposed to yenow. and black-white.
each other1 as are blue and yell ow (Hering, 1878). Yolmg and Helmholtz were
describing a trid lromatic theory w ith thJ"ee basic colors (red , green1 and blue);
1-:Iering's opponent color theory h ad four basic co lors in two opp onent pairs:
red versus green , and blue yellow. A black-versus-white component
fom1ed a third opponent pair, but this 11 opp onen cy" isa bitdifferei1t although
you can' t have a reddish green, gray can be descri bed as a blackish w hite .
Figure 5.14 illustra tes this idea. T11e cen ter ring s hows the hu e dimens ion
of HSB space, wrapp ed into the color cirde as in Figure 5.13. llle outer ring
o ffers a ca rtoon of four color m echan isms in hvo pairs. The inner patch o n
the left that looks yellmv in opponent color theory bec.:1 use it stimul ates the
yellow pole of the yellow-blue opponency and does not stimulate e ither red
or green. Move a bit counterclockwise, and the orange patches a dd increasing
red to the decreasing yellow.
YeUow Blue
Red
FIGURE 5.14 Hating' s Idea o f oppo1lent cciors. H€&fing noted that aJI the colors
on the ..color circle• (the center ring) oould repr9Seot9d by two pairs o f opposing
colors: blue versus ye4bw, and red versus greein {cartooned in the outer ring). Thus. a
color could 00 a reiddlsh yellow or a blutsh green, but not a reddish gree<"I or a bluish
yellow. (After Stockm an and Brainard, 2010.)

136 CHAPTER 5
8230336 Anm
(a) Here is .1 Ught th.lt
looks bluish gtl!:en.
(b) lfl add-a bit

of light that
looks red ...
• (J) II the ug ht
looks greener ...
(r) ... more red

will be n eeded
to cancel the
•
•
g fli!en •.•
(c) .. .I can cano?l the • (jJ ....md I will be left

green.and I w ill be with the weaker
left with only the b lue. blue C'01Tlponent.
FIGURE 5. 15 A hue c anc 911atlon expieriment st<:U"ts w ith a oolor (here, bluish green
on the left and a greGner hue on the right) and attem pts to determine how m uch of
thQ opi::on1;mt color of one o f the starting color's com pongnts mus t bQ added to 91irnl-
nat e any hint o f that component from the starting color. In this example, the observer
adds red to canOBI green, 198.ving only' th9 blue com porn;rnt o f the bluish gr9"fl .
Leo 1-huvich and Dorothea Ja meson (1957) revived Hering's ideas and
d eveloped one way to quantify this oppone ncy. Th e meth od, ca ll ed "hue
can cell ation/ is sh own in Figure 5.15. ln this. example, on the left, \.Ve start
\.vith a lig ht that ap pears to be a bluish green. \Ve can can ce l the perceptual
greenness by adding its o pponent color, red. \Ve measure the am o unt of a
light th at looks red that is needed to just rernove all traces of green. The result
wil l appear n either red nor green. It wnI be a shade of blue. Si nce it isa mix o f
lights th at look bluish g reen and red, it will be a ra ther desa turated-looking
blue. If we do this for a greene r color, as on the right of Figu re 5.15, it will
ra ke more dd ded light that looks red to cancel the green 1 and the result will
be an even fainter blue remna nt. If we s tarted wi th p ure g reen, once it was
ca nce.l led, the result woul d be an ach romatic p atch because there would be
n o blue t o be left over. 'h'e coul d can cel the perception of blue in our bluish
green p.i td 'l es by adding li ght that looked yellow. ln this case, the renmant
would appear green.
U \Ve did hue can cellation experiments fo r ligh ts across the spectrum, we
would get resul ts that Jook like those in Ffgure 5.1&-n ot exactly like these,
because these a re Dorothea Ja meson's d ata and everyone has slightly diffe rent
results. If we start a t abo ut 400nm, the lights look reddish blue (or violet) an d
we need to add some green a nd son1e yellow to cancel them. But look w ha t
happens a t about 470 mn. Here is a light that has n o red or green to ca ncel; it
looks perfectly blue. This location on the spectmm is kn o\lvn as \.Uli que blue.
Continuin g to scan a long the spectrum in the fig ure reveals the locations of
tw"o othe r unique hues-hues. that can be described w ith on ly a single color
term. Only fo ur hues can be d escribed in this way. As you might g uess, they
are red, green, yellow_, and blue. Only th ree of them have unique loci on the
spectnun. All the very long wavelengths look red (with_, m aybe, a to uch of
residua l ye llow), The two crossin.gs of the red-green fun ction p rovide the loci
of tmiqu e blue a nd lul.i que yello w (Fig ure 5.16n) . The point where the blue-
urlque hue ""1 of four cdcrs that yellow function crosses from positive to is the locus of tmique green,
can be described w ith cnly a single the green tha t h.:LS n o b lue or yellow in it (and, of course, n o red ) (Figure5. 16b).
color term: red. yellow, green. blue.
Other cdcrs (e. g., purpleor crange) There are exce llent exemp la rs of other colors, s uch .:ts orange or purple1 but
can be described as ocrn pounds (red - they are not unique in the sam e way. O range, n o mdtter how p ure, can still
dish blue, reddish be de.scribed as a red dish yellow.

(a) Unique hues
Wue Yellov.·
Value.s above the

hne mean that you
are adding green
tocancelrednese.
Values below the

line mean thilt you
u e adding red ·to
cancel greenne.ss.
------v------ - - - - - - -
-..-
400 500 600 700
Wnel.ngth (run)
(b) Unique
&"'n
Values above the

line mean that you
are adding blue to
canal yel.lowne&a.
400 500 600 700

Wawlength (run)
FIGURE 5. 16 Here are results from a hue cancellation 9XpQ'irnent The locations
where the hue cancellations cross the neutral midpoint are the locatbns of the unique
blue, green, and yeMow hues- for 9.Xample, the green hue w ith no hint of blue or yel-
low in it. "Unique red" is not defined by just one spectral Jorus.. (After Hurvich and
Jameson, 1Q67.)
Thisrom>onent·process, color-appearance story sounds rather similar to the

L-Mj and ([L + M]-S) story. (L- M) is not the same as red versus
green, and ([L + M] -S) is not the same as yellow versus b lue. If this were the
case, an ([L + M] - S) cell should be a yellow-blue cell, m aximally excited by
unique yellow and maximally inhibited by unique blue; but a more detailed
look reveals that adding and subtracting cone sensitivities to produce those
cells results in cells tha t respond maximally along an axis
extending from a purplish hue to a yellowish greenish hue.
The(L-M) cells aren' t in quite the right place either. TI1e l.rconeend of the
axis is n ear perceptual red, but the M·con e end is a bluish green (EskeWi 2008).
Krauskopf, Williams, and Heeley (1982) call these endpoints the "cardinal
directions" in color space, but they are n ot perceptual reel, green, yellow, and
blue. We n eed another transformation of the color signals in order to make
cone-opponent signals into color-opponent signals. The series of transforn1a-
tions that we've been talking about is depicted in Figure 5.17. Three cones

138 CHAPTER 5
FIG URE 5.. 17 Three stgps to cobr (a) Step 1: Detection (cones)
perception . iia} Detection: light Is dif-
f;eri911tially absorbed by thr99 photopig- 1.0
in the cones. f./J) Discrimina-
..C·.S
• "'
1i.
0.8
tion: dlffefen:::es are taken between
conQ types, creating oone-oppcnent 0.6
mechanisms, in-iportant for wavelength
discriminations. {c) Ap?QaranCQ furttwr
• $ 0.4
riecombnatlon of the sigrials creat9S ]:§ 0.2
cok:ir-o ppon9nt JXOCQSSQS that sup-
port the color-opponent naturQ of color 0.0
appearanoe. (After Stockrnan and
Brainard, 2010,)
(b) Stii>p 2: Discrimination

15
't
·o !l:
(c) S tep 3: Appe.uance (opponent colors)
-·h02·
l1 15
8230336 Amm.:i Ap
05
t1l
-05 -J
"di!<e- -2 504 nm 510 nm
i Ill
500 600 700

Wavelength (mn)
g ive rise to cells that m easu re the differen ces in activity be tween cone types
behveen. Ptuthe r transformations are required to crea te p rocesses that give
rise to prope rtie/;i of color appearance like the unique hues (Stockman and
Brainard, 2010).
Color in the Visual Cortex

Cells in the LG N seem to be cone-opponen t cells, so the transforrnations that
produce the m lor-.opponent processes that .support color appearan1..-e are likely
to be found in the ";sual cortex. On e usefu l transformation of the information
is s how n in Figure 5.18. The LGN has opponent cells, and Figure 5. 18a. shows
a cartoon of one s ud1 ce ll 's receptive fie ld. We will call it L+/ M-1 indicating
that it is excited by redder hues in its center and inhibited by greener hues
in its surround . (For present p·urposes, we w ill no t worry about whether this
ce ll is cone-oppo nent or color-opponent. It is just easie r to talk about red and
green in this case.)

(11)
- -
® -
(b) FIGURE 5. 18 Color-opponent recep·
tiw 'fields. l;a) Sing le ·opponent cells
get excitatory input from one cone
® ® ®®
Best stimtth.lS type and inhibitory input from another.
p) These cei ls respond i,.igorously
0 to regions of the correct color. Here

L+JM- prefers the n9gion that appgars
8
Single-
opponent cel l
rf!ddlsh. (c) The cert er o f each double-
opponent cell gets excit atory input from
one cone type and inhil::Hory input from
Even better anoth er. The pattoarn is rtrversoo in the
®
surround. (d) Th9se cells respond \'igor·
(o) (d) ousty at an OOge between m o colas.
Worst position
Best stimuh.1s
Neutral Neut<al
® ® @®
Double-
opponent cel l
@
Beat poQtion
Figu re 5.18c shows a doubleMopponent cell. Do uble-opponent cells are

first found in the vis ual cortex (E. N. Johnson 1 Hawk.en, and Shapley, 2001).
In the example s hown here, the ce ll is exci ted by redder hues in its center and
by greener hues in the s urround, and it is inhibited by redder hues in the s ur-
round ,md by greener hues in the center. To see why this is interesting, look
r.t Figu res 5 .18b an d d, where we have s uperim posed these recepti ve fields
across a red / green borde r. ln this contex t, the cone-opponent cells, d escribed
earlier, could be called single-opponent cells. A single-opponent cell conveys
info rmation abou t oLa Lyoad ar In this mse, the cell is m ost excited
by red and not green. A d o uble-op ponen t in contra.s t, conveys information
about chromatic edges. Tiuts, the rnos t "exciting" place for a single-oppon ent
ce ll-in Figme S.18b is somep lace en tirely in the red area \v h ere LMcone input is
greatest and M<0ne is reduced. Fo r the d ouble-op ponent ce ll in Figure5.1&1,
the best place is o n t],e red-green edge w here M-conescan have som e input to
the s urround while L-cones d ominate in the center. The double-opponent ce ll
marks w here the color changes. TIU.s information can be useful when you 're
trying to use color to carve the world into surface3 and objects (see 01apter
4). (See Web Essay 5.1: Why Use Opponent Processes?)
Though w e know that color information is transformed in the visual cortex,
it is not dear how the physiology gives rise to perception (Gegenfurtner, 2003).
Many cells in cortex are interested in color but do no t seem to linearly add and
double-opponent cell A cell t;Pe,
fo urd In the visual c ortex, In which one
subtract inputs from different cone types as the cells cartooned in Figure 5.18 reg k::f1 Is excited by me cone twe,
do. There is some evidence tha t they add and subtr<Jct in a nonlinear manne r. cornblnatk::n of cones. or color and
For ins tance, a.cell might be responding to something m ore like L2 - M1+ 5 2, Inhibited by the opponent cones or
thou gh exactly ho w this would produce the colors \Ve see remains a topic fm color (e.g .. R+,G-). Another adjacent
regk::f1 would be Inhibited by the lrst
future work (Horvvitz and Hass, 2012; Wa ndell and Chichilnisky, 2012). For
Input ard excited by the second
now, we may have to agree , .._,i th Conway (2014, p. 201): "There is s till no good In this example, R-!G+).
nelU·al explanation for Hering 's psychologically important col ors ... or ...
single-opponent cell Arother
the unique hues o ften associa ted with them ." wa; to refer to cone-opponent cells,
Perhaps the biggest controversy con cerns the possibility of an area (or In crder to differentiate them from
areas) in the brains of monkeys and hmnans specialized for color vision. In the double-oppcnant cells.

140 CHAPTER 5
achromatopsia An Inability to per- 1980s, evidence favored a separate pathway for colQr..,'Ilwre were "blobs1' in Vl
ceive colors that cause::J by damage where cells did n ot seem interested in orientation bul'-seehled very fnte ·ted
to the central nervo...Js system. in r.:olOJ'. The blobs sent output to " thin stripe " regions in V2 (Livin gsto ne a nd
qualla In phlloocph'f, a private Hube l, 1988) and from there to V4, an area tl1at Semir Zeki argued was special-
consdoos exper1enoe of sensation or ized for color, wi th cells that responded not to wavelength but to perceived
perception. color (Zeki, 1983a, 1983b). More recen tly, functional imaging in monkeys has
been used to uncover color hot spots in visual corte;oc. TI1ese have been named
"globs'' {really! Conwa y, Moeller, and Tsao, 2007), and cells in these regions
also seem m o re intereste d in color tha n do cells ou tside . H owever, altho ug h
these anatom.ical pathways are the:re (Fed erer et al., 2009), it has become less
clear that we can separate color processing from o ther perceptual processes
in cortical anatomy (Shapley and H.:1wken, 201 1). lndeed, you rnay remember
frorn Chapter4 tha t V4is a popular candidate for a "shape " area. This d oesn' t
mean that it is unii:terested in color, but m erely that it is probably n ot only
interested in colo r (Roe e t al., 2012).
FURTHER DISCUSSION of the elegantly named "blobs" can be found in

Oiapter 3
on page 76.
Modern imaging studies sh ow some of the ln.unan visua l cortex tha t

seem partkularly interested in color (Grill -Spector and Malach, 2004), but we
can' t record from sin gle cells in humans under m ost d ri.,_"1.fil1st:::mces. Perhaps
the best evidence for specialized brain ar eas for color in hum an s comes from
certai n cases of achromatopsla, a loss of color vision after brain dam age
1990). In achrom<1 topsia, patients m ay be able to find the boundaries between
regions of different colors, but they cannot report what those colors mig ht be.
l.n these pa tients, vision is largely intact, while the experience of color seem s
specifically impaired.
Individual Differences in Color Perception

Philosophical Problem of "Inverted Qualia"
Thus far, with a little cauti on, we have been talking about color as if we all see
co lors the sa me way, but d o we? Th is is one of those questions that everyone
asks a t one point or an othe r, often as a child . Like many "childish" questions,
th is h as been the topic of rnuch decidedly un childlike philosophical work.
1lle seventeen th-centnry British philosopher John Locke (1 632-1 704) may
have gotten things s ta rted when he asked w hat the conseq uences would be
''if the Idea, that a Violet produced in one Ma.n's Mind by his Eyes, \Vere th e
same tha t a Marigold produces in another f\.fan' s, and vice versa." He suggested
that the answer was n ot going to be readily forthcoming "because one Man's
lvlind cmlid not pass into another Man's Body" (Locke, 1689/ 1975). Today,
we would talk about the qualla (s ingular qua le) of vio let a nd marigold yellow,
rather than the ideas. Qualia is the technica l name for th e subjective aspects o f
experiences like /(violet" or "sweet" or "itd1.. " 1l1e problem that Locke posed
is often called the '' inverted spectrum" problem, th ough "in..:e rted qu.alia"
problem m ight be more philosophically correct (Byrne, 2014). 1l1e problem is
often posed by imagining that the s ubjective colo r qu.alia in a figure like Figure
5.14-have been exchanged lvith the opponent color qualia on the other side of
co lor circle, tho ug h the pairin g of nam.es with specific physical stimuli would
not d1ange. Tims, a region that evoked the qua le of "red " for you \<1.1ould evoke
"green" for an inver t, but that '"'·ould just be his internal experience. He would
have learned- as you lea rned-to call that experience "red," so it would be

very hard or, perhaps, impossible to tell that his s ubjective ex pe rience was
the opposite of ymus. 1lle inverted qualia idea and its variants can be used
to d iscuss a range of significant philosbphica l iss'ues, well beyond our scope
here. This hypothetical situation can also be used to .raise some interes ting
questions about our experien ce o f color.
Loo k back at Figure 5.12 and think abo ut th e sahmition axis of the HSB
colol' spa1...--e. \\'hat is the color halfway from a saturated red tov..:ard the totally
d esaturated v•IUte or gray point? You would probably say "pink." NO\.\irconsider
the o ther side of the color circle in Figure 5.14. Ha lfway betvteen green and
w hite is somet!Ung \.•;e might ca ll "light green" or 'jpalegreen," but '""'e would
not have a commonly accepted, unique color nan,e f ti
di scuss color nam es in a m oment, but for now, think about our friend with
the inverted qualia. Would " pink' ' make sense to him? Where we experience
the lluale "pink/' he would experience the quale "light green," but pink is a
distinct color with some special properties (Lindsey et al., 2010) while lig ht
green is not. CouJd we tell th e difference between individuals \o\oith standard
qualia and those with inverted qualia by care.fully exa mining their responses
to pink? Since inve rts are, m ost likely, a philosoph ic:a l invention, we can ' t
give an fil\S\\'er. However, r aising the question helps us to unders tand that
the relationship, be hveena mix o f reflected from the world and
your s ubjecti ve experience of color, mlght no t be simple (Palnler, 1999).
Language and Color

Suppose you are in a clothing s tore and you find a shirt that you like but you basic color terms Color words
want a different color. You might go to the clerk and ask, "Do you have this in that are monolexemlc (single "oNOrds
blue?" At tha t moment, you a re not worried about the qualia experienced by ' blue" not, ' sky blue;, used >Mth high
frequency, am have meanings tliat
the clerk, but you d o assun1e that you and tha t clel'k agree about the mea ning are agreed upon by speakers of a
of blue. How d o we talk about lan guage to each other? You rn n discri minate language.
on the orde r of milllons of diffe rent colors, but you d on ' t have a separate c ultural relativism In sensatk::n and
word for each of these. There is a vast range of co lor words, but in lookin g perception. the Idea that basic peroop-
for that shirt, you p roba bly would no t ask for 11az ullne " or "cerulean" {both, tU91 exioerlences (e.g., color peroop-
varieties of b lue). You would use a wmd that almos t every speaker of yom tlorQ may be determined In part by the
language would use quickly and cons istently in naming colors. These colors cultural environment.
nam es are the basic color terms of the language. \Vhat makes a color term
basic? Berlin and Kav (1969) as.serted that it must be comn10n (like red and
not like beige), n ot ai; object or substan ce name (excl uding bronze and ofive),
and no t a compmmd word (no blue-3reen or ligh t purple) . TI1ls classification is a
little subjective (is beige that uncommon ?), bu t Berlin and Kay argued that, in
English, these mies }';eld a list of 11 terms : red, green, blue, yi!llaw, black, wlrite1
gray, orange, purple, brown, and pink.
the number of ''basic" color terms differs dramatically across
cultures, d own to as few as 2 or 3. At on e time it was thought that the diffe ring
numbers of basic color terms in di fferent languages meant that color categori-
:z.a tion was arbitrary. This n otion was called c ultural relativism, meaning that
ead< group was free to te its own linguistic ma p of color space. Berlin and
Kay's important d iscovery was that the v arious maps u.sed in different cultures
are actua lly rathe r si milar (Llndsey and Brown ,2006). After surveyin g man y
languages, they found that the 11 basic color terms in English are abo ut as
ma ny as any group possesses. Of oom se, the words themselves dlffer, but red is
rouge (flrend1) is adom (H ebrew). Mmeover, languages do n ot select randomly
among the p ossi ble color nam.es. U a language has only two basic co lor terms,
spe ake rs of the lang uage divide co lors into "light'' and " dark." If a language
has three co lor terms (on e d1rom<"ltic term beyond ligh t anddnrk), what do you
think the third usually is? If you guessed rt>d, you are correct. Typically, the

142 CHAPTER 5
fourth color term would be yellow, then green and This ordering is n ot
absolute but you won' t find a language with, for example, just purple, greeu,
and gray as its basic color terms.
Ill Red How do new basic color tem1s emerge? Berlin and Kay argued a big
Yellow color tenn is l;'artitioned in to tv.,ro sm a lJer terms . Levenson (2000) s uggested
c- tha t ne 'basic...terms tend to emergent the boundary between rn.·o existin g
Blue
Brown
co lor terms, in the area w here neither existing term works vv-e ll. In fact, both
processes may be a t work. Lindsey and Brown (2014) looked a t th e use of
= ••••••••••••• color words in American English by asking 51 Americans to name the color of
Ill Purple each of 330 color pa tches (like paint chips). TI1ese observers were told to use
Gray a single word That ,..,·ord had to be a word that could be used for anything
Teal of that color (you can' t have a "blond" car, can you?). Lindsey and BrO\·Vn
were looking for the sort of word you might use in everyday speech to name
tha t shirt. Those 51 Americans used 122 temls for 330 colors. El'f'ryoue used
the basic 11, but there was evidence that American Englis11 1night be moving
beyond the 11 basics. A colo r like "teal'' may become basic. It emerges in the
no man's land of colors that are neither blue nor green. A term likep11rp!e may
even tually be prutitioned, with a term like lmJender or lilnc taking over some
of the purple real estate in colo r space (Rgure 5 .19).
If o ur set o1 basic color terms increased, would tha t change the way we see
color? If a language has only two or three bnsiccolor terms, do its speakers see
colors differently than we do\•1dth our 11 basic tenn s? Eleanor Rosch (H eider,
1972) studied th is question a mong the Dani of Ne\v Guinea, a tri be w hose
language has only two basic color temlS: mo!n fo r light-warm colors a nd mili.
for dark-cool colors. Now, it is hard enough to ask your neighbor to define the
experience of blue and tl1en to ask if tha t is tl1e sarne as your experience of blue.
It is much more difficult to ask these questions across a grea t cultural divide.
But there are tricks, as the ex:periment illustrated in Figure 5.20 demonstrates.
Suppose you are shown a bluis h color chip and asked to remember it. Then
yo u nre s hovvn hvo test chi ps and asked to pick the color you saw before.
Obviously, the lt"Ss similar the two test colors are, the easier this task is. But
more im porta nt, you v.>i.ll do be tter if the v.rrong choice is on the other side of
a color categoric.al boundary. Color boundaries are sharper than yo u might
think. lf you s how people a collection of colors and ask, "Which are blue and
w hich are green?" people do the task without mud 1 d ifficulty. If you have to
remember a 12olor, as in the task s hoW11 in Figure 5.20, you are likely to give
it a label like "green" or "blue." [f the next color has the same label, you are
FIG URE S.19 Wh4im Unds19y ond more likely to be conf used titan if it has a different label.
Brown C20 14) asked Ame<ic.ans to Rosch found tha t the Dani's perfonnance on s uch tasks reflected the same
name cola patch€1s, QV0fYOne u8'1d color bounda ries, even when their language did not recognize the distinction
the 11 "basic" colors. The other color be tween the hvo colors (a Dani might call all the colors in Figure 5.20 mi.Ii
narneis \Nere used by fewer and fei,.ver
but would still do be tte r vvith the task in Figure 5.20b). This finding leads to
peopla, gcing toward the t:ottom of
the graph. A ftm of these other colors. th e conclusio n tha t color perception is not especially influenced by cu lture
like peach and t eal, may b9com 9 basic an d language; blue a nd green are seen as ca tegorically different, even if o ne's
overtime. language does not e mploy color ternlS to express this difference.
In the late 1990s, Debi Roberson went up the Sepik River in New G uinea
to study the Berinmo, whose lang uage, like the Dani 's, has a limited set o f
basic color terms. Unlike previously s tudi ed g ro ups, the Berinmo have terms
tha t for m novel botmdaries in color space. For example, their nol/wor distinc-
tion li es in the middle of colo rs we ca tegorize as green, and may rough ly
d istin guish Hve from dead or d ying foliage. Moreover, when the Berirnno did
the color me mory task, tl1ey perform ed bet ter across their nol/wor boundary
than across the blue/ green boundary. Eng lis h speakers sh m,.·ed the opposite
result (Davidoff, Davies, a nd Roberson, 1999). More recently, sirnilar results

(a) (b) FIGU R E 5.20 It is 9asief to remern-

OOr w hich of t'NO c d ors you have s.:ien
if the choices are cat egorically differ-
Pnt. For example, supposa you had to
rflirrl9mber the · blue'" patch shown at
the top o f each part o f th'9 figuni;i. Pick·
ing between two ·blues,• as In part (a),
'NOuld 00 rather hard. The t ask would
be QBSier it o n9 o f the chcices \N"Elre
H.uderchoice Eafilerchoice and the o tt1er "green.• as in part
(b). even If the d lstanoe In color space
WQfe the same in the WJo cases.
have been found in comparison of English and Russian subjects across the
Russ ian categorical disti nction between light blue:s (golriboy) and dark b lues
(s i11iy) (\Vi.nawer e t aL, 2007). And even m ore recen tly, evid ence has eme rged
tha t lea rning new color s ubca tegories produces increases in gray m a tter in
parts of the bra in impli cated in color vision (V. Kwok et al., 2011 ). So, can
language or culture influence color perception? For many years, we thought
the answer was "no." Now, thanks to Robe rson's work, we are no t so s ure.
Let's think a little rnore abou t the blue and green patches in Figure 5.20.
Looking across the bottom four disks, you can categorize them as blue, blue,
blue, and green. At the sa me time, you can tell tha t the first and second blues are
not the sarne. ft tu m s o ut that different pa rts of the brain seetn to be involved
in these different ways of aM lyzing color. When Bird e t al. (2014) s howe d
o bservers in a hmctional magne tic resonance ima ging (fMRl ) e xpe riment a
series a f patches of one color, folJowed by a change to a pa tch of a different
color, vis ual cortex responde d to the size of the color change (green to "n ear
blue" < green to 11 fu r blue"). ln contras t1 a location i.n the frontal lobes seem ed
inte res ted in th e ca tegory o f the colo r. G reen to blue produced the sa m e
response, regardless of the s ize of the cha n ge. Blue to blue did not produce a
cha n ge, e\o·en though the change was p erceptib le to the observer.
Genetic Differences in Color Vision

The indh·i dual differences described in the previou s t wo sections are either
small or, in the case of inverted qualia., hypothetical. Under most circmns tances,
if you d eclare two lights to be m etamerically matched, those arotmd yo u w ill
generally agree, even if we can' t make definitive s tatements about their qualia.
There will be some varia tion between individuals. For example, unique green
cru1 vary between observers from at least495 to530 run (Nerger, \ :Olbrecht, and
Ayde, 1995). Some of these differences "111 Pt;due tel fo owlik ag51: which
turns the lens o f the eye yell ow Q. S. \'Verner, Pe terzell, and Scheetz, ii 990). To
a flrs t a pproxi mation , however1 your pe rfor man ce on s tandard measures of
color vision w ill be the sa me as others,,.
l-I mvever, there is a s ignificant excep tion to this unh·ersa1ity of color match-
ing . Some 8% o f the ma le population and 0.5% of the fe male population have
a form of colo r vision deficiency conm1only known as "colo r blindness," in
w hich there is a ma lfuncti on in one or rnore of the gen es coding the tluee cone
photopigments. It's il "guy thing'' becau se the genes that code for the M - and
L-cone photopigme nts are on the X chromosome {Nathans, 1986). h ave

144 CHAPTER 5
only one cop y of the X ch romosome, so if one is d efective, the male in ques-
tion w ill have a problein. Fema les have hvo copie.s and can ha ve n orrna1 color
\,.ision even if one cop y is abnorrna l (actually, su ch \"lornen ca n end up with
four different con e pigrnents) (.Nagy e t al., 1981). 11'1e S.-cone photopigrnent
is coded elsewhere, so everyone h..1s two copies, and therefore S.cone color
d eficien cies are rare (Alpern, K.i tah.ara, and Krantz, 1983). (See Web Essay
5 .2: Experiencing Color Blindness.)
TI1ere are a number of different types of color blindness. One de te rmining
factor is the type of cone affected . A second factor is the ty pe of d efect; either
the photopigment for tha t cone type is anoma lous (different from the nom1) o r
the cone type is missing a ltogether. Although we call people w h o a re m issing
one cone type "colo r-blind,'' it is a mistake to think tha t this m eans they ca n-
n ot see colors at a ll. As you \¥i ll recal l, if you have all three cones '¥i th their
s tand.a.rd pho topigments, you need three priaiary colors to make a meta meric
match wi th an arbitrary p,tttd1 of color. If you have t\vo cone types ra ther than
three, the nom1'-llly th ree-dimensional space becomes a two-din1ensional
space. The world "'ill sti ll be seen in color, but you will have a "fl a tter'' color
experience, d ifferent from that of people v.ri th normal color vision.
Because M- and L-rone defects are the most common, most color-blind indi-
viduals have difficul ty discriminating Lig hts in the middle- to-long-wavelength
range. f:or exn mple, consider the wavelen gths 560 and 610 mn . Neither of
these lights acti vates $-cones very much, and the L-cones fire a t about the
same rnte fo r both . But most of us can distingulsh th e ligh ts on the basis o f
the M-cone outpu ts they elicit, '"·hich w ill be hi gher for the560-nm light than
for the 610-nm li ght (you can confinn these assertions by consulting Figure
5.5). English-speakin g trichromats wou ld labe l the colors of these two ligh ts
as ''green'' and " re ddis h ora nge," respectively.
Now consi d er a deuteranope., someone who has no M-cones. H is pho-
toreceptor output to these hv o ligh ts ""-ill be identical. Following our maxim
tha t the res t of the visu al syste m knm'\is only "''h a t the p h otorecep tors tell it,
560- a nd 610-nm li ghts mus t a nd will be dassified as the sa me color by our
d eu tera nop ic individual.
deuteranope An lrdlvl:Jual wro suf- A protanope--someone w h o has n o L-cones- w iU il.."lVe a d ifferen t set
fers from OOcr blindness tl1at Is due to
of color ma tches based on the outputs of his hvo cone types (Mand S). And
1he aboonoe of M -caies.
a trttanope--\vith no S-cones-""'ill be different again. Gene tic factors can
protanope An irdMdual wrosuffers also make people Color-anomalous individ uals ty pi cally
frcm color blindness 1hat Is due to the
abserce of L-oones. have three cone photopigrnen ts, but two o f them are so si mila r th.a t these
individ uals experience the world in much the sam e way as individuals lvi th
trltanope An Individual wro suffers
frcm color blindness that Is due to the
only h vo con e types.
absen::;;e of S-cones. \Ve actually have some n otio n of exactly what the world looks like to color-
d efi cient indhiduals, because the re are a few, very rare cases o f individuals
color-anomalous A better term
for what Is usually called •color-bllrd. • w ho are color-blind in only one eye. TI1ey can compare what they see w ith the
Most 'color-bllnd" lr-.:JMduals can color-blind eye to w ha t they see with the n ormal eye, enabling us to reconstruct
stlll rnake discrtmlnatlons based on the appearance of the color-blind world (M acleod and Lennie, 1976).
wavelength. Those dlscrlm !nations True co lor blindness does occur but it is very tmusuaL ft is possi ble to
are different frcm the norm- that Is.
be a cone monochromat, with on ly o ne type of cone in the re tina. Cone
anomak>us.
monochromats (who also have rods) li ve in a one-dimensional color s pace,
cone monochromat An Individual seeing the world only in shad es of gray. Even m ore vis ually impaired are rod
with only one oone type. Cone rnono-
c hromats are truly color-blind. monochromats1 w ho are missing cones a ltogether. Because the rods work well
only i:n dim light and are genera lly absen t in the fovea, these individuals no t
rod monoctromat An lrdMdual only fail to discriminate colors; they also have very poor acuity and serious
w ith no cones of any type. In addition
to being truly coior-bllrd, rod rnom- difficulties seein g tmder n om1al d ayl.ight conditions.
c hromats are badly >Asually Impaired In \Ve already mentioned one other very inte resting class of color blindness,
brtght light. 1.."'0 ming not from photoreceptor problems but from damage to the visual oortex.

Lesions of specifi c parts of the visua l cortex beyond primary vis ual cor tex can a{J'losia A failure to reocgnlze
cause achromatopsia. An achroma top sic ind ividual sees the world as drained of ct:>)ects In spite of the ability to see
color, even w hile s howing e viden ce that wavelen gth infom1 ation is p rocessed tram . Agnosla is due to brain
damage.
a t ear lier stages in the visu al pa thway. Brain lesion s ca n a lso produce various
for ms of color agnosia or a ne mia (Ox bury, Oxbury; an d Humphrey, 1969). In anomla An Inability to name objects
In spite of the ablllty to see ard recog-
agnosia, the p a tient can see something but fai ls to know w ha t it is. Anomia nize them (as shown by usage). Ano·
is a n ina bili ty to n a me-in th is case, an ina bility to n ame colors . A patient mla Is l)?lcally due to brain damage.
w ith a nemia might be able to pick the banan a tha t ulooks right" but tu\Elble
color contrast A color perceptlm
to repo rt tha t the banan a is or s hould be yellow. effect In which the ook:lr of one region
Induces the opponent ccior In a neigh-
boring rajon.
From the Color of Lights to a World of Color
color assimilation A color percep-
All the work of this chapter1 up t o this point, has concerned the d etection , d is- tion effect In Vlfilch !'MO colors blee:J
crimin ati on, and appearance of isola ted lights . However, we live in a world Into each other, each taking on some
\>•there regions of one color a but regions of r-mo ther, a nd this p roximity chan ges of the chromatic quality of the other.
the a p pear an ce of colors, as Figure 5.21 shows. 1n color contrast effects, the
color o f one regio n induces the op pone nt color in a neighborin g region. Thus,
in Figure S.21a the yell ow s urround weake ns the yellow of a central sq uare
and s tren gthens the blue. ln color assimllatlon effects, two colors bleed into
each o ther, each taking on some of the chroma tic quali ty o f the other. So, in
Figure 5.21b the blu e in the first column loQL reddish or pw:plisl n the to p
image and greenish on the bottom .
Not only can other colors i.n the scene alter the color of a target region, but
scenes can contain colors that canno t be experienced in isolation. Though it
may be ha rd to believe unless you try it1 you cannot sit in comp le te da_rkn ess
(a) Color oontrMt
•
(b) Coloras.simifation
FIG URE 5.21 Cobr oontrast and oolor assimilation.

(a) In oolor contrast, thG C9ntrol squ an;;i takes on c hro-
matk: attributes that are opposite those o f the surround .
so the gr99r1 central squar.e lcoks gro9Elner on the r9d
background thWI on the green background. (b) In color
assimilation, odors blend together locally. So, in the
S9cond column th.e yellow sq uares look a bit reddish in
the upper squ arG and a bt g reenish in thG bwer squarG.
(A:ft9r Stcx::krnan and Brainard, 2010.)

146 CHAPTER 5
utTelated color A color that can be and see a grny light, all by itself. Tiiot light wi ll look white if seen as an isolated
experienced In Isolation. or unrelated color. To be seen as it mus t be seen i.n relations hip to othe r
related color A color. such as patches of color. Thus it is a related color. \Ve mad e this point earlier, ta lking
brown or gray, that Is seai onty In about brov.rn. In isola ti on, a light mi ght look yellow or orange. Lt w ill only
relation to other colors. Fcr example, to patches. We can dis tinguish a f e\'\i' thousand
a •gray" patch In ccmplete darkness
appears white.
un related colors. AIJm'\>'UlS for contex t effects is what boosts the number of
distinguishable colors to the millions (Shevell, 2003).
negative afterimage An afterimage
w l1ose pdarlty Is the op posite of the
crlglnal stimulus. Light stimuli produce
Adaptation and Afterimages
dark negative afterimages. ColO!S Color contrast effects show how the spatial relations between colors can influ-
are oornplementruy: for example. red ence color a ppearance. Temporal relations ma tter, too. What you saw before
produces green. and yellow prodUJ:es has an influence on the color you see now. You already know this from the
blue.
discussion of light a dapta tion in Chapter 2. Adapting to a bright light ma kes
adapting stimulus A stimulus a moderate light look darker. Adapting to darkn ess would ma ke that same
w11ose removal produces a change In mo derate li ght appear brighter.
visual perception or sensitivity.
neutral point ll1e pdnt at which
FURTHER DISCUSSION of the time course of dark adaptation can be
an opp::nent color mechanism Is found "'1 Chaptel' 2 on pages 4 7-49.
generating no signal. If red-green and
blue-yellow medianlsrns are at their
neutral points. a stimulus will appear Now Jet's extend tha t principle to color. Adaptation can be color-specific,
achrcmatlc. (The black-white prooess as we see in the phe no1nenon of negattve attertmages. If yo u look a t one
' ""no neutral point. ) color for a few seconds, a subsequently viewed ach romatic region w ill appear
to take o n a color opposi te to the original color. \ Ve can calJ the firs t colored
stimulus the adapting stimulus. The illusory color that is seen aften vard is
the negati ve afterimage. (See We b Ac tivity 5.4: Afte rimages.)
The principle is i1lus trated in Fig ure 5.22. Figure 5.22a consis ts of a circle
of g ray sp ots. Now, s tare at th e black d o t at the cen ter of Pigure 5.22b a nd
consider whnt happ ens as you expose one bit of your retina and visual systern
to the red dot at the top of Figure 5.22b. The L-cones w ill be more stimulated
th an the M- or S-cones. L+/ M- oppo nent processes w ill be s timulated. You
w ill see \ Vhen you move your eyes back to fi xate on the black dot a t
the center of Figure 5.22111 the red is w ithdra\'\-11 from tha t area o f the visual
fi e ld. The L-cones wi ll be mo re adap ted tha n M- or S-cones, as w ill the
later processes in the retina a nd brain that were more s timulated by the red
spot. Adapted processes be have as tho ugh they a_re somew hat tired. They
respond less vigorous ly than unadapted processes. The result is a bit like
w hat would happen if you he ld a pe ndulum up and released it. The red-
green oppo nent color mecha nis m swi ngs back toward the neutral point,
overs hoots th is point, and slides over to the green si de. As a consequence,
the gray s pot appears g reenish until the opponen t mechanism settles back
to the n eutral point.
If you look at the green d ot at the bottom of Figme 5.22b a nd then look
back a t the gray image (Fig ure 5.22n), you w ill see the result of pushing the
red-green mechanism in the o ther direction. Other colors '""ill prod uce othe r
results, w hich you sho uld n ow be able to predict. In Figures 5.22c--e, you can
try this '"''i th a real scene. If you s tare at the black d ot in Figm e5.22c and then
quickly m.ove your eyes to the same dot in Figure 5.22d, you will see a
tinted version of the real photograph (Figure 5.22e, tho ugh this effect w ill be
more dramatic if you look a t the version in We b Ac tiv ity S.4: Afterimages).
Notice that we are not attributing negati ve afterimages to just the cones or j11st
one set of cone- or color-opponent processes. Adaptation occurs a t multiple
sites in the nervous system th ough the p rimary generators are in the retin a
{Zaidi et .'1., 2012).

THE PERCEPTION OF COLOR 14 7
• •• •
(•) (b)
•
•••
(c) (d) (e)
FIGURE 5.22 To understard w hat negatiV9 aft91magesare, study the image in

p art (a) and oonvine& yourself that a" the c irclgs are gray. Now star& at the black dot
in part (b). Aft91' 10 seconcls or so, shift your fixation t o the black dot In '8). The cird es
should n ow look cdored . This is a negative aft9rlmage. Wrry does it happen? Of it
didn't hapPQn , try fixating more rigorously. R9El.lly look at the black dot.) Now try with
a real scene. Fixate the black dot In (c), and then fl ick your gyes t o the same spot in
(d). You should 899 a vvashed-out version of th9 colors In {a).
Color Constancy
You m ay recall fro m Ch apte r 2 tha t ad aptation is one way of dealing with the
vast range of light levels in the e nvironme nt. Assuming tha t you are readin g
this on good old-fashioned p aper, w hen you are inside, you see this black text
on a w hite page. If you take this book outside, you still see black on white,
even though the illumination is so mudl brigh ter that the black text, viewed
outside, reflects more light to th e eye tha n does the w hite paper inside. TI1is
is an achromatic example o f the m ore general phe no meno n of colo r con-
stancy, the tendency for the colo rs of objects to appear relatively Wlchanged
in spite of s ubs tantial changes in the lighting conditions. in
this book seem to have more o r less the same colors wherever you read the color constancy The tendency of
book (though there is an entire research area hidden in wha t we might mean a surface to appear the same oolCf
by " m ore or less''; Fos ter, 2011). trder a fab'ly wide range of lllurrtlants.

148 CHAPTER 5
FIG URE 5.23 The S...."1111Q surface (8) (a) Thesurffice

illumir1ated by two lights \b) w ill
gen19rate tiNo diff.Qf&nt patterns o{ activ-
Perce.nl llge ...\ sutilce in the world
ity in the s -. M- , and L-cones (c.---e). re flects different
However, the suriaCQ will appear to bQ of light
re fl ected percent,1ges o f
the same color Lll"ld€< both illurninants. diff;:>re n t wavelengths.
This phenomenon Is known as color
constancy. l;Aft"1r Smithson, 2005.)
(b) Illumin.m tl Illumin.mt 2
Ye llowish sunlight
Enersy and bluish sk)'light
8230336 AmMa App of light are comIX'sed of
differen t mixt ures
of w,1velengths.
(c) Su1faoo x Surfaoe x

illuminant ilhmtin,mt
ReJ.iti,•e reache..s the

amou n t eye is th e surface
of light ret1ect.mce mu ltiplied
b)' the illunUnant.
(d)
Cone Theresulti.s !i<i?en

sensithrity by the three cones.
(e)
This p rcduces two
very different sets
Cone of th1't'e numbers from
responses th.es.:tmesurface.Hov.·
d o Wi' knov.· what
colo r th,1 t surfaceis!
figure 5 .23 illus trates w hy color cons tancy is yet another difficult problem
for the visual sys tem to solve. The heart of th e problem is that the illuminant,
the lig ht th at illuminates a s urface, is not constant. Lighting cha nges as we
Illuminant The light that Illuminates go from inside to outsid e or as the s un moves from the h orizon to high in the
a surface. s ky. Figure 5.23o s hows the spectral reflectance function fo r a surface-the
spectral reflectance function The percentage of ead1 wavelen gth that is reflected from a particular surface. \ Vith
percentage of a particular wavelength its preponderance of long and s hort waveleng ths , this s urface probably looks
that is reflected from a surface. purplish. Let's ca ll it 0 lilac." Figure 5.23b shows the spectral power distribu-
spectral power distribution The tion-the relative a mount of light a t different visible wavelen gth s-of two
physical energy In a light as a function differen t types of "d aylight": s unlig ht and skylight. Sunlight is a yellowish
of wavelength. light, richer in middle and long wave lengths; skylight is more bluish. Figure

5.23c s h m<1i'S that the light reflected to our eyes is the product of the s tuface
and the Wtunination . For example, a surface might reflect OOYo of 650-nm light,
but no650-nm light would reach the eye unless there wassonlein the orig inal
ilhm1ination . Figure 5.23d shows that th ose two different products of s urface
and illumina tio n are conve rted into two diffe rent sets of three numbe rs by
the L-, M- and S-cones (see Web Essay 5.3: Color Constancy in the Lab).
H ere's the prob lent. Even though the th ree numbers from the three cones
are different in the t\1y·o conditions, that lilac s urface will look lilac-colored
under both illumirumts. \'Vhite p aper will look w h ite . A ba nana \vi ii look
yello'""" This color constancy is beneficial because we want to know the color
of the object. Und er normal circums t.:m ces, we donrt care about the spectral
comp osition of the lights.
The Problem with the Illuminant

Le t's think about colmcons tmcy asa m ath problem. In simpletenns, we have
an illumi nant (ca ll it [) and a s urfa ce (S). As shown in FigUI'e 5.23c, what ""-e
can sense is a res ult-the product of 1 >< S-but wha t we \Vant to know is S.
It is as if we \\'ere given a munber (sa.y, 48), told tha t it is the p roduct of h¥o
other numbers, and asked to guess w ha t those h\.•o mnnber.s might be. The
answer could be 12 and 4. Or it could be 16 and 3. Or 6 and 8. Given just the
number 48, we cannot solve the problem. NeYertheless, given the result o f I
x S, the visu..'ll system d oes a pretty good job of figuring out S.
We smnetirnes talk about "discounting " the illuminant as if our w hole
goal '"''ere to throw away the I term an d just see the surface color. However,
this is n ot quite right. You can tell the difference behveen a scene lit by the
tnorn.ing sun and a scene illtuninated by the s un at high noon. Thus, no t only
can you recover the color of the stufcice, bu t you also kn ow somethin g about
the illuminant. H ow d o you d o this?
Physical Constraints Make Constancy Possible

As noted in the previous section, it is impossible to know whid1 hvo integers
are multiplied to produce 48. However, if you a re told that the first number
is behveen 9 and 14, you 're saved . The first mm1be r must be 12, and the sec-
ond, then, mus t be 4. In a nalogous way, color constan cy must be based
on some information o r assumpti ons tha t cons train the possible answers.
There are rnany possible assu mptions that could help. Suppose we assumed
that, in a complex scene, the brightest region was white (Land and McCann,
1971) or th at the ave.rage colo r across the whole scene was gray (Buchsbaw11,
1980). VVe could scale the other colors relative to these white or gray anch ors.
H owever, th is can ' t be entirely right. Think wha t would happen if you were
in a dark romn w ith two spots of light on the wa ll: a red one and a blue one.
Under a simple version of a bri ght-is-white theory, the bright"' SJl .t s h<ru\d pp.:t
look w hite and the o ther spot would ch ange color. That can' t be right (and
theorists knew this, so the actual theories are m ore s ubtle, but they s till do
not work perfectly).
There are other poss ible constraining piece,!i of information. Assumptions
can be made about the illtuninant. For instance, natural light sources (and most
artificial ones, s ud1 as s tand ard lightbulbs) are generally "broadband .., 111.at is,
they contain many waveleng ths, even if some wavelen gth s are no t as intense
cis others. Furthermore, their spectral composition curves (see Figure 5.23) are
usu ally smooth; spikes at particular wavelengths are tmoomm on, thou gh this
generaliz..1.tion is violate d by some artificial light sourC'eS. Highly mmahual
(e.g., m onochrorna.tic) light sources nMke the world look hig hly ulm:Jtural-a
fact exploited in nightclubs the world over. Ind eed, with a. m on ochromatic

150 CHAPTER 5
reflectance nie'peniehtage of !git source, color vision is impossible, though not much broadband light is needed
hitting a surface thal Is re!ected and to get color back.
not absortled nto the surface. Typl- Assump tions can be made about s urfaces. Real s urfaces also tend to be
caJIY rel\ectance Is given as a fLf1ct1on
of wavelength. broadband in their renectances (recall the hamburger distributions of Figure
5.6). It would be very unlikely, for example, to fin d a surface that reflected
100% of 535-nm light, 0% of 538-nm light, a nd 100% again of 540-nm light.
Even surfaces tha t look like s ingle wavelengths of light typicaUy reflect a wide
range of wavelengths. lhat is, the red bars in Figure 5.23 m aybe metameric
with som e thing like a 600-nm light, but that region is sending many other
wavelengths to your eye. TI•ere are othe r limits o n the reflectan ce o f real s ur-
faces. The whitest surface rarely reflects more tha n 95% of any wavelength, and
the blackest rarely reflects less than 5%. The brightest thing in the ' isual field
is like ly to be white. A r 1specular reflection" (like the shiny spot on a billiard
ball) has a wavelength composition very similar to that of the illuminant. Any
o f these facts might help.
Assumptions can be made about the structure of the world . Sh arp borders
in an image are ahnost always the result of boundaries between surfaces, n ot
boundaries bernreen light sou rces. Thus, if you see something tha t looks pink
next to something that looks golden, it is very unlikely to be the result of clev-
erly placed pink and golden light sources (wUe.ss you' re in a thea ter, perhaps).
Shadow borders can be an exception to this rule (Adelson, 1993; Cavanagh
and Leclerc, 1989). A shadow can produce a sharp edge th at is unrelated to
any change in the underlying sUl'face. Hmvever, the change across a shadow
border is typically a change in brightness and not a change in the chromatic
properties of th e regions. [n Figure 5 .24a, you can easily imagine that you' re
looking a t three rectangles with a shadow lying across their middles. Not so
in Figure 5.24b; though the top and bottom in Figure 5.24b are brighter than
the middle, the darker region d oes not look like a s had ow because, for that
d arkenin g to be the result of a s ha d ow, the shadow would have to be coin-
ciden tly aligned with a set of hue changes. Our implicit knowledge of these
sorts of constraints helps us sort out the visual world .
C ues like this, p erh aps in clever combination, can be used to solve the
otherwise unsolvable problem of colo r constancy {Sm ithson, 2005). All of these
assumption s about the state o f the world nUght sound a lot like the "prior
probabilities" that are important in Bayesian theories of the sort introduced
in C hapter 4. Bayesian ideas have been put to work to explain color constancy
(Brainard, 2009; Brainard and Freeman, 1997). The hard part, w hether you' re
working in a n explicitly Bayesian framework or n ot, is to get the prior exactly
right (Foster, 2011 ). What are the precise assumptions that give us our level of
color constancy? We are still not su re.
(a) Luminance change without hue (b) Luminance change ;vith hue
change looks like a shadow. change looks less like a shadow.
FIGURE s. 24 The visual system Mknows· that brightness changes across a shadO'IN
boundary but hue do9s not. As a result. (B) looks like a shadO'IN but (b) does not.

THE PERCEPTION OF OOLOR 151
FURTHER DISCUSSION of Bayesian theories in the context of vision c an FIGURE 5.25 Th9 strawb0tTi9s in
be found in Oiapter 4 on page 110 and in Web Essay 4A: Bayesian a complex scene '!Mii iook red even if
Analysis. there is not muc h long-wavelength light
in the illuminant.
Nonetheless, we can broadly describe how the visual

syste rn mig ht use these assumpti ons to achieve color
constancy. Suppose, for example, that you 're standing
in a kitchen Lllmninated by a light so urce co mpose d
primarily of short a nd rnidd.le wavelen gths (Figure5.25).
None of the surfaces in your visual field wi ll reflect much
long-'""·avelength light, because there isn 1t much of this
light to be reflected . But given the assumption th.a t the
illuminatio n is evenly distributed around the room, you
will s till perceive the s trawberr ies as red, because they
reflect more long-wavelen gth li ght than a ny o the r s urface
does. (See Web Ac tivity 5.5: Illusions of Lighting .)
What Is Color Vision Good For?

H :wing introduced some of the b asics and some of the
complications of color vision, we will '"':-rap up this chapter
by asking about the u sefulness o f color vision. 111e a bility
to use wavelength information h.:1S evolved several times
in several ways during the course of time. Evolutiona ry
theory tells us tha t, for this to be true, color vision must
provi de an '-'dvantage th.."lt m akes it worth the tro ubl e.
Color is n o t a n absolute requiremen t. If we co uld not
make wavelength discriminations, we could s till identify Appa
a lion (Rgure 5.26a), and we could still find o m th.ro ugh the fo rest (Ftg-
ure 5.26b). Although color vision might make the lion stand ou t a bit be tter
from the trees, we would be much m ore impaired if we lacked 11 orie ntation
vision '' or "motion vision." Across the an imal kin gdom, h owever, there seem
to be a t least hovo realms of behavior w here color vision is especia lly useful!
(a) (b)
FIGURE 5.26 A b\ack·and-white lion is still a lion, and you could still find a path in
the 'NOOds w ithout cola"" vision.

152 CHAPTER 5
(a) (b)
FIG U RE 5.27 Finding a raspberry is sasier if you h av9 color\'ision, as is d;;iciding if

that bony r1p<>.
eating a nd sex. More gene rally, color vision see ms to be particularly he lphLI
in visual se..·uch tasks.
H m.; ng color vision d oes seem to m ake it easie r to find c.mdidate foods
and to discrimin.:ite good food from bad food . Comparing the hvo versions
of Figure 5.27, it is dear tha t finding berries is easier \vi th oolor vision. Notice
also tha t it makes it easier to decide which of these berries is ripe. You may
reca ll that we m entioned this p ossibili ty w hen disc lLSsing the L- and M-oon es,
earlier in the chapter. Mo.st diurnal an imals h ave two p hotopig ments: roughly
an S-cone and an L/ M-cone. Some prim.ates have evolved separa te Land M
photopigments an d the ne ural circuitry to exploi t the rather s ma ll differences
between the resp onses of th ose cone types. There has been cons ide rable debate
regarding w hether th is rea lly conveys an advan tage in telling ripe from unripe
fruit or d istingi.U5hing subtle differen le.w••cSome hnveargued
tha t s hape texture information, a long wi th dkhromntic co lor vision, are
good enough for these purposes. How can we find out? Some m onkey species
(e . g., capuchins) have a mix of dkhromatsa nd trid)J'()mats (as d oes the huma n
species). Do the trichromats have an advantage in hunting for food? Rather
tha n test this in cap udtlns, Melin etaL (2013)carefu lly simulate d six varieties
of capuchln colo r \oi.sion in human observers: and had those observers search
for capudtin food in photograph s of those fru its as they would appear in the
Costa Rican forests w here the monkeys live. 1lle result was a d ear advantage
for trichromatic vision..
The food-color vision connection is n ot limited to primates. The colors
of vvi ldflowers did not deve lop to please the aesthetic sense of hum ans. As
a gen eral rule, they are ad vertisemen ts to bees and othe r insects, offering to
trade food for sex (well, at least for pollination o f the flower). ln fact, rnany
flowers have dramatic pattems tha t we can not see, because they are variations
in the reflection of short-\•:avelen gth (ultrav iole t) ligh t, which is outside o ur
range. Bees can see these s hort wavelengths, and it is the bees that flowers a re
designed to attract (Figure 5.28). \,Vhile we're on the s ubject of the relationship
of plants and animals,it is worth a detour to point ou t tha t plants exploit several
sensory systems in an e ffort to attract the attention of the righ t ani mal. The
s mell of a fl owe r is intended as a sjgnal to pollinators, and even the shape of
.specialized le.a.ves can crea te an auditory beacon to a ttract echolocating bats
(R. Simon e t al., 201l ).

(a) Cb)
FIG URE 5.28 Cola"" vision In different species. (B) These b lack·eyed Susans are
shown as they are S9'lf'l with human pho toreoeptors and color v ision . (/:>) A honeybee
can see UVllght. In W. the flowtt's "black.eye· (or maybe the pupl of that eye) is
muc h larg er- a better t arg 9t for th9 bQQ. (Coul19sy o f Tom Elsnar.)
In a dditi on tosea rchi.ng fo r a nd assessin g foo d, a n imals spend signi ficant

time and e ffort sea_rching for and assessing po tential m ates. Here, too, color
plays a central role. ColorfrJ d isplays-from the dramatic patterns on tropi-
cal fish (Figure 5 .29a) to the rail of a peacock (Figure 5.29b) to the lace of the
mandrill (Figure 5.29c)- are all sexual signals. Wha t makes the peacock that
has the most colorful tail the most desirable m<lte for a fernale peacock? \Ve
can' t ask her, of course, but a colo rful tail ntlght somehow indica te tha t this
peacock's genes are better tha n his competitors' . A female peacock that sees
the world in black and \.,.· hite won' t be able to perceive this information and
will therefore be a t a n evolutionary disad vantage. in an o ther example tha t
we menti oned earlie r, the deve lopment of separa te L- and M-cones may h.:we
made prim.a te color \·ision partiaJarly well s uited to d e tecting the amount of
blood in a blushing or blanched cheek (Changizi, Zhan g, and Shlmojo, 2006).
Color vision is accomplis hed in different ways in different species. We are
trich romats, \'\o; th three different types of p hotorecep tors. Dogs appea.r to be
dichromats, with two types of photoreceptors (Neitz, Geist, and Jacobs, 1989). FIGURE 5.29 The colors of animals -
Chkkens,smprisin gly en o ugh, tum out to be tetr.1duomats, with four (Ok..1:.110, from (a) trcpical fish to (b} peacocks to
(c) mandrills - are often
to p oteintial mates.
(ii) (b) (c)

154 CHAPTER 5
{a) Different photopigments can tune photoreceptors

to differmt wavelengths.
400 700 400 700 400 700 400 700
(b) Colored oil droplets can al.so hlne photoreceptors

to different wavelengths.
Light
p p p
400 700 400 700 400 700 400 700

FIGUA E 5.30 Two to make photoreceptors with different spectral SGnslttvi-
tles. (a) Our S-, M- , and are different becausetheyoontaln different phot-
oi;Xgments. (I:>} Some animals haw only one type o f pt'otopgmi1::mt. These animals can
have color vision because colo red oil droplets sitting on top of photoreceptors create
groups of photaQCQPtors with diffsrQOt sensitMties t o waVQ/911gth.
Fukada, and Yoshizawa, 1995). There is not much gain in information if the
number of photoreceptor types is increased beyond 3 or 4 (Maloney, 1986),
which probably explains the lack of octachromats o r doclecachromats- indi-
viduals with 8 or 12 types of cones, respectively.
Our S-, M-, and L-cones are different because they contain different phot-
opigmen ts {Figure 5.30a ). It is also possible to use a single photop igment to
create more than one functional type of cone. The trick is to put a different
fi lter in front of each type of cone so that some wavelengths are s ubtracted
before light read1es the photoreceptor (Figure 5.30b). A cone with a reddish
o il droplet in front of it will respond more vigorously to long-wavelength light
than will a cone covered by a greenish droplet. Chicks and othe r birds have
these droplets, as do a variety o f reptiles (Govardovskii, 1983).
Even fireflies get into this act in a limited way. Fireflies signal each o ther
with bioluminescence: they make their own light. Different species have differ-
ent lights, and each species' visual system appears to be tuned. to its particul..u
wavelength signature. A combination of a photopigment and a colored filter
makes signals of conspecifics {members of the same species) appear brighter
than the flashes of other fireflies in the vicinity (Cronin et al., 2000). With this
sort o f visual system, the firefly will never appreciate the palette of colors in a

sunset. But it will be able to locate an appropriate mate, and that, after all,
the pressure s haping the development of this limited sensitivity to wavelength.
Summary
1. Probably the most important fact to know about color vision is that lights
and surfaces look colored because a particular distribution of wavelengths Refer to the
of light is being analyzed by a particular visual system . Color is a mental Sensation and Perception
phenomenon, no t a physical phenomenon. Many animal species have some Ccmpanbn Website
form o f color vision. It seems to be important for identifying possible mates,
slles.slnauer.com/wolfe4e
possible rivals, and good things to ea t. Color v'ision has evolved several
times in several different ways in the animal kingdom. for aotMtles, essayo, study
2. Rod photoreceptors are sens itive to low (scotopic) light levels. There is o nly questicrn, ard other stu:ty atls.
one type of rod pho toreceptor; it )'i.elds one " number'' for each location in
the visual field. Rods can support o nly a one-dimensional representation of
color, from dark to light. Thus, scotopic vision is achromatic vision.
3. Humans have three types of cone pho to receptors, each having a different
sens itivity to the wavelengths of lig ht. Cones operate at brighter light levels
than rods, producing three numbers at each locatio n; the p attern of activity
over the different cone types d efines the colo r.
4, If two regions of an image produce the same response in the three cone
types, they will look identical; tha t is, they will be metamers. And they
will look identical even if the physic.al wavelengths coming from the h\•o
regions are different.
5. In additive color m ix ture, two or m ore lights are m ixed. Adding a light that
looks blue to a light tha t looks yellow will produce a light that looks ·white
(if we pick the right blue and ye llow) . In subtractive color m ix ture, the fil-
ters, paints, or o ther pigments that absorb some wavelengths and reflect
o thers are mixed. Mixing a ty pical blue paint and a typical yellow paint \<till
subtract most long and short 'vavelengths from the lig ht reflected by the
m ixture, and the res ult ,.,,,ill look green.
6. Color blindness is typically caused by the congeni tal absence o r abno rmal-
ity of one cone type--usually the L- or M-cones, usually in males. Most
color-blind individuals are not blind to differences in wavelength. Rather,
their color perceptio n is based on the o utputs of h\'O cone types ins tead of
the normal three.
7. A s ing le type of cone cannot be used, b y itself, to discriminate between 8230336 Amma Appa
wavelengths of light. To enable discrimina tio n,. information from the three
cones is combined to form three con e-opponent processes. Cones sensitive
to long wavelengths (L-cones) are pitted against medium-wavelength (M:)
cones to create an (L- M) process that is rorigJily sensitive to the ra:lness or
greenness of a region . (L + M) cones are pitted ag;Jinst sho rt-wavelength (S)
cones to crea te a process ror1ghfy sensitive to the blueness o r of a
regio n . TI1e third process is sensitive to the overall brightness of a region.
8. Color appearance is arranged arotmd opponent colors: re..i versus green,
and blue versus yellow. This color opponency involves furth er rep rocess-
ing of the cone signals fro m cone-opponent processes into color-opponent
processes.
9. The visual system tries to disentangle the properties of surfaCES in the world
(e.g. 1 the "red " color of a strawberry) from the proper ties of the illuminants
(e.g., the "golden " light of even ing), e ven though surface and illuminant
in formation are combined in the input to the eyes. Mechan isms of color
co11Stancy use implicit knowledge about the world to correct for the influ-
ence of different illuminants and to keep that strawberry looking red tmder
a wide range of conditions.

CHAPTER 6
•• 82303 Am A
Christcph9r Smril, Rooting, 2014

Space Perception
and Binocular Vision
IMAGINE 11-ltir YOU' RE MOVING QUIETLY TI-IAOUGH A MEADOW, trying to get
d ose enough to a bear cub to get a good picture. Suddenly you discover that 8230336 Al'lmd App.l
Mother Bear is n ot happy about your pho to se86o'ion. Sh e charges, and you mn-
back across the field, through a thicke t of trees. A quick leap across a stream
brings you toa s lope that dm'Vll to the road w here yourparb1e r is waiting
in a Jeep. You dive in to the passenger side and roo.r off d ovm the track to safety.
As the bear lumbers off and your heartbeat returns to normal, your thou ghts
nahrrally turn to the acts of visual space perception that you just performed: You
picked a path through the three-dimensional world tha t brought you to safety.
You behaved as though you knew where the trees were. You acted as though
you tmderstood how far it was across the stream . All in all, you d emonstrated
a sophisticated grasp o f the layout of the physical world arotmd you
Hum an s share this sophistica ti on with a large part of the animal king d om.
Faced vtith the same bear, a rabbit or deer would havesho\vn a s imilar grasp
of the relev..'lnt issues in space perception (v1,.;tl1o ut the Jeep). The a bili ty to
perceive and interact wi th the s truchire of space is o ne of the fundamental
goals of the visual sys tem. It i.s a lso quite a fonn.ida ble accornpli.s lunent, and
in th is chapter we' ll explore how we d o it.
As a stm-ting place, let's ass ume that the external world exists. This is a
phil osophical position kn.own as realism. It is n ot the only possibility. T'he
positivists note that all you really have to go on is the evidence o f ymu senses,
so the '"'orld could be nothing m ore than an elabora te ha lluci nation. For less
philosophicaJ elaborati ons of this vie'\-vpoint, we could consult the writings of
Philip K. Dick and other science fiction authors. ln this book, we'll just ilssume
that there is a rea l world out there to perceive. realism A philosophical position
The geometry of that real world is Euclidean (nmned in honor of the ancient arguing t11at there is a real world to
G reek geom eter Euclid, of the third cenhuy BCE). TIUs means that parallel lines sense.
remain paralle l as they are extend ed in spdce, that objects maintain the same posltMsm A phlloocphlcal position
size and s hape as they move aro und in that s pace, that the internal angles o f arguing that all we really have to go on
Is the evlden:::e of the senses. so the
a triangle alvvays add to 180 d egrees, and so forth. All that s hiff you learned 'MO<ld mlgl1t be nothing more than an
in high school math classes is true in the real world. elaborate hallucination.
Euclidean geometry is n ot the only geometry: Alth ough the real worl d. is Eu-
Euclidean Referring to the geom-
clidean, the geometry of retinal irnage.s of that world is decidedJy non-Euclidean. etry of the world. so named In honor
The geometry becomes n on-Euclidean \Vhen the three-dimensional world is of EL>Olld, the anolent Greek geometer
projected onto the a.uved, two-dirnens io naJ s urface of the retina. Parallel lines of the third century BCE. In Eudldean
in the world do n ot necessarily remain parallel in the retinal image, as Figure gecmetry, parallel lines remain parallel
as they are extended In space . objects
6.1 illus trates. l11e angles o f triangles d on' t always add up to 180 d egrees.
maintain the same sJze ard shape as
The retinal a rea occupied by a n object gets s maller as the object moves farther they move around In space, the lnta--
away from the eyeball. What a ll this means is that if we want to appreciate the nal angles of a trlari;Jle always add to
Euclidean world, we h ave to reconstruct it from non-Euclidea n input. 180 degrees, and so fCfth.

158 CHAPTER 6
FIGURE 6 .1 ThQ Eud id9an gQOmetry

of the thrM-dimtf'lslcnaJ 'WOrld turns Into
something quite diff9rent on the a..nv9d,
t..vo-dimensional In the Euclk:lean
I workl, the angles of a triangle add up to
/ 100 dQgrQQS. In the no n-EuclidQfill W"Orld
of the retinal image, this need n ot be so.
i
/
I
,/
More precisely, genera lly our v isual ex-perience is a recon-

structio n of the world based on two non-Euclidean inputs: the
n...-o distinct retinal images. Close your left eye, stretch yo ur left
a rm out in front of you, and hold up your left in dex finger. TI1e n
hold up your right index finger about 6 inches in front of yotu
face, so th at it appears to be positioned just to the left of the left
index finger, as illustrated in Figure 6.2. Now, quickly open your
left eye and close your right eye. [f you positioned your fhlgers
properly, your right finger ivill jump to the o ther side of your
Retinal image
left finger. Although this demons tration is designed to exagger-
ate the differen t v iews of your two eyes, the point is a genera l
on e: the two retinal images always differ. They differ because the retinas are
in slig htly d!Herentplaces in your head. Just t'IS you and the person s tanding
nex t to yo u see somewhat diffe rent views of the world, so d o yo ur two eyes.
Much o f this chap te r v1rill be devoted to explaining h ow the visual syste m goes
to quite elaborate len g ths both to exploit a nd to reconcile these differences.
FIGURE 6.2 The t¥i'O retinal images

of a world are not
the same. See the tQXt for detoHs.

SPACE PERCEPTION AND BINOCULAR VISION 159
Above (b) Human Front
t
Seen by
.· · ·· fj'-..._
•• both eyes ,1
Back FIGURE 6.3 Com paring rabbit and

hutnan visual (a} A rabbit 's lateral
eyes can see all around its head . ft>) It
c an wen SQQ abov& its head, making
Why have t\.vo eyes a t al1? Perhaps m ost ftmdamenta ll)'i having two eyes its vtsuaJ som&thing like a p lane-
confers the same evolutionary advantage as having two lungs or two kidneys: tarium dome (with ears). (c) The human
you can lose one eye and s till be able to see. A second advantage to doubling visual fie.Id Is mo re like a windshield
the number of eyes is that they enable you to see more of the world 1llis is c ove<ing a large rggion in front of the
ieyes. Light purple indic ates th9 9rlti re
especially true fo r animals like rabbi ts \·vho have lateral eyes on the sides of \'isual field ; dark purplQ, the part of the
their heads. A rabbit can actually see for 360 degrees around its head (Figure Visual field seen by both Qyes.
6 .3a). This explai ns w hy it is so hard to sneak up on a rabbi t. Moreover, the
rabbi t can also see s tmig ht up above its head (Flg ure 6 .3b) (Hu ghes, 1977).
Humans, wi th frontal eyes, still see more of the \Vorld \vi th nvo eyes than
w ith on e. Our visual field is limited to about 190 degrees from left to right,
11 0 degrees of w hich is covered by both eyes (FJgure 6.3c). The field is m ore
restricted vertically: about 140 degrees, 60 degrees up to a limit defined by your
eyebrows and 80 d egrees dmvn to your cheeks, as is s hown in Web Essay 2.3:
C linical C ase: The Man Who Couldn 't React The exact size of your visual
field wHI be limited by the specific anatomy of your cheeks and eyebrows.
Ove!'lapping, frontal, binocular visua l fields give p reda tor anima ls su ch as
humans a better chance to s pot .sma.Lt fast-moving objects in front of them that
mig ht provide d inner. Prey anintttls, like rabbits, are often those with very
wide visual fields allmving the m to m onito r the w hole s cene for predators.
With frontal eyes and overlapping ·d sual fie lds, you get the advantage
of tw'o detectors looking at the sa n1e thing. For example, if t-.vo independent
people each had a 50% chan ce of missing a target, the ch ance that both of
them wou ld m iss it would be SOo/u x Sm'o = 25%. So the chan ce tha t at least
one of the two would find that target is l OOo/o - 25%, or 75%. This is kno\<m
as probability summation. Somethin g similar h appens if the n.vo eyes both
look for the same hard-to--see target. In visio n, this wo uld be called binocular
summation (Blake, Slo.."U1e, and Fox, 1981). Binocular sm11mation may have binocular W th two eyes.
provided the e\·olution ary pres.sure that first moved eyes toward the front of probability summation The
some birds' and mammals' faces. Under most circtmlStances, we do no t ge t increased detection probabll lty based
complete probability summa tion. TI1e increase from a SC»'o chan ce to a 75% m the statistical advantage of ha'1n;i
two (or more) detectors rather than
chance assumes tw'o completely independen t observers, but o ur two eyes are me.
still embedded in on e person. Nevertheless, we wW do better a t many tasks
w ith t.w9 than wi\'1 jus t one R..K Jones and Lee, 1981). binocular surrmatlon The combi-
nation (or wsummatlon; of signals 1rom
Once the eyes rnovM to the front, though, evo lution fo und an a dditional each eye In ways that make parfor-
use for overlapping visual fields . Try this: Talce the top off a pen and hold the man::e en many tasks better with both
top in on e hand , the pen in the other, both about a foo t in front of your face, eyes than with ather eye alone.

160 CHAPTER 6
binocular disparity The differences with your e lb ovvs bent. Now, d ose one eye and try to quickly put the cap on the
between the t wo retlnru lmagoo of the pen . Repeat the sam e task \Vith both eyes open. For m os t (but not all) people,
same scare. Disparity Is the basis fcr th.i's task is e..1 sier with two eyes than with o ne. This is a quick d em ons tration of
sterOCl)sis , a vMd perception of the
three-d lmenslonallty o f the world that the usefulness of binocular disparity-the differences beh.veen the hvo retinal
Is not avai lable with mo nocular 'Asion. images of the sarne world . Disparity is the basis for a vivid perception of the
monocula- With a-10 eye.
three-dimensionality of the world that is not available \vi.th purely monocular
(one-eyed) vision. The technical tem1 for this binocular perception of depth
stereopsls The ability to use bin-
is binocu la r stereopsis. The geometric and physiological bases fo r binocular
ocular disparity as a cue to depth.
s te reopsisare the topi c of a brge p ortion of this chapter (see also I. P. Howard
monocular depth cue A depth cue and Rogers, 2001).
that Is available even v.ihen the world is
>Aewed with one .,.,.,, alone. \.-Vhen you decide you n eed a break from reading this ch apter, take that
break with one eye cl ose d. You s hould be l:lble to notice the loss of binocular
binocular depth cue A depth cue
s tereopsis (and of part of your ,.; s ual field), but a peri od of one-eyed v;sua l
that relies on lnformatlon from both
eyes. Stereopsis Is the primary ex.am- experience w ill also ma ke it clear that binocular s tereopsis is n ot a n ecessary
ple In hurnars. but convergence and co ndition for d epth pe rception or s pace perception. Rabbits d o very well
the ability of 1'X> "1'0• to see more of with very little binocular visio n, and pain ters an d m ovie directors manage
an object than one .,.,.,, seoo are also to con vey realistic impressions o f depth on fbt canvases a nd m ovie screens.
blnorular depth cues.
O n the other h and, binocular s te reopsis d oes add a rid1n ess to perception of
tl,e three-dimens ional worl d, as yMdlXdescribed by O li ver Sacks (2006) in
his artide about "Stereo S ue/' a neurosc1en li.st who re?in&.1 s tereopsis at the
age of 48. We will ta lk about he r later,
[n this ch apter, firs t we describe the set of monocular depth c ues to
three-dimensional s pace. After that, we turn to the mo re complicated topic of
binocular .stereopsis, a binocular depth c ue. Finally, in the last section of the
chapter, we cons ider how the vario lLS c ues a re combined to p roduce a unified
perception of s pace.
Monocular Cues to Three-Dimensional Space

M . C. Escher (189&-1972) titled the drowing in Figure6.4 Rela tivity. Escherwos
a master of the rul es that govern o ur perception o f s pace. Each bit of stairway,
each land ing, every person-all a!'e drawn using
cues that enable us to infer three dimensions from
rn.·o. But when we try to fo llOlv those stc1irs, we
find that Escher's drawing cleverly fails to add up
to a coh erent representation of a place tha t could
exist. Even when no one is try ing to foo l us, it is
geometrica lly imp ossi ble (not to mention compu-
tationally infeasible) for the visual system to crea te
a perfectly faithful recons truction of Euclidea n
space, given the n on-Euclidean input we receive
throu gh our eyes. The best we can do is to use
d epth cues to infer aspects of the tluee-dimensional
world from our h\ 0-d.imensional retinal images.
1
On the basis of the reti na l images and an implicit

understa nding of physics and geometry, we collect
cues tl1at provide hin ts about the like ly struct ure
of the s pace in front of us and the disposition of
objects in tha t space.
FIGURE 6.4 M. C. Esch€11·, Relativity, 1953, Escher

deploys m onorular rues to dEpth in a way that is
essentialty correct at ooch locatbn but that adds up to
an Impossible world.

Unless we' re sh1ck in an extremely impoverished perceptual environment

(say, the Sahara durir1g a v!e of the world provides mul-
tiple depth cues. Usually the cues reinforce ea<:n combining to produce
a convincing and reliable representation of the three-dimensio nal world.
Occasionally, however, the cues are contradictory. Escher could fool us by
deliberately manipulating depth cues and other routine visual inferences. He
arranged sensible local cues into a globally implausible story. lVhat cues does
the visual system use to infer depth relations, and how do we use those cues
to create a representation of the three-dimensional world? (See Web Activity
6.1: Monocular Depth Cues.}
l=IGURE6.5 ()ccluslon makss It easy

Occlusion
to infer r91attve position In deipth.
Some of the cues to the layout of the three-dimensional world were introduced
earlier in this book, because hints to the layout of space can also be hints about
the structure of objects in that space. Occluslon is an example (see the section
titled "Finding Edges" in Chapter 4). In Chapter 4, occlusion was a cue to the
presence of an otherwise invisible edge. As a depth cue, occlusion gives infor-
mation about the relative position of objects. Thus, in Figure 6.5 we are happy
to infer a circle in front of a square in front of a triangle. Occlu.siQn is present
in almost every visual scene (we challenge you to find a situation in nom1al
life where nothing blocks your view of anything else}, and many researchers
argue that it is the most reliable of all the depth cues. lt is "WTong only in the
case of "accidental viewpoints" (remember those, from Chapter 4?). That ls,
the retinal image shown in Figure 6.5 could be produced by a circle and two
oddly shaped puzzle pieces, as shown in Figure 6.6a. That scenario would
require careful placement of the objects and the viewer. [tis mud1 more likely
that Figure 6.5 would arise from a more generic view of a circle occluding a
square occluding a triangle (Figure 6.6b).
We do not know from the occlusion cue alone whether the red square in
Figure 6.5 is in front of a small green triangle, a larger but more distant triangle
(maybe a green tree}, or an even larger but even more distant green moWltain.
Occlusion is a nonmetrical depth cue; it just gives us tJ1e relative orderings
of oocluders and occludees. A metrical depth cue is one that does prm.; de
information about distance in the third dimension.
occlusion A cue to relative depth
Size and Position Cues order In which. for example. one object
The image on the retina formed by an object out in the world gets smaller as cilstructs the view of part of anothe<
the object gets farther away. (See Web AcUvlty 3.1: Visual Angle for a review.} object.
Moreover, your ·d suaJ system knew this fact of pro Jeettve geometry implic- nonmetr1cal depth cue A depth
itly before you ever picked up this book and learned it explicitly. Projective cue that provides Information about
(a)d
the depth ordef depth) but not
depth magnitude (e .g., his nose Is In
frmt of his face).
(b) mebical depth cue A depth cue
that provldoo quantitative Information
0
about distance In the third dimension.
projecUve geome1ry For purposes
of stu<fylng perceptlm of the throo-
dlmenslonal world, the geometry that
descr1bes the transformations that
oocur when the throo-dlmenslonal
world Is Pfqleclro mto a two-dlmen-
slmal surface. For example. parallel
l=IGURE 6 .6 Flgur9 6.6 oould be an "accldgntal• view of the pieces sho"Wn h9r9 In lines do not cmverge In the real world,
lt Is much more lik.Qly. hoWQVQ', that it Is a gM9ric view of cir de, squar9, and trl- but they do In the two-dimensional
angl9, as shown In fP). projection of that workJ .

162 CHAPTER 6
336 Amma Aooa
FIGURE 6.7 This Is a photograph o fa oollsctio n o f Plasticine boJls that are resting
on the sam9 su rface at the same distance frcm the camera Nevertheless, the small
ones appear to be farther a"Nay. Sarne portion of the \'isual system assumes that all
of th9S9 items are the Sm'lG. If one ball projectt;. a smaller image on the retina and if
we assume that the balls really a re the same size, then the smaller cnEt must be far-
thQ'" away. This is the cue of relative siz9.
geometry describes h ow the \Vorld is projected onto a surface . For exarnple,

a s ha dow is a projection of an object onto a s urface. An implicit understand-
ing of the rules of projective geometry ca n be said to undergird many of the
depth cues described here. In this case, the visual system knovvs that, all else
bein g equa l, smaller things are farther away. H en ce, the Plasticine balls in
Figure 6.7 may appear to lie in different depth planes. \Ve ca n call this depth
cue relative size.
The impression of three-d imension ality in Figure 6.8 is more po\.\-·erful than
that in Figure 62 because we've added another rue. The critical difference is in
the organization of the objects in the hvo figures. In Figure 6.8, the rabbits fo rm
an orderl y texture gadlent, with larger objects in one area and smaller objects
in a nother. Because smaller is interpreted as farther away, this arrangement
creates the perception o f a g rotmd plane receding into the d istance.
In Figure 6.9, the rabbits are again arrayed in an. orderl y texture, but here
we get less of a sense of d epth . The differen ce behveen Figures 6.8 and 6.9 is
that the fo m1er includes another depth cue that is n ot p resen t in the latter:
relative height lmagin e that you 're standing in a field of rabbi ts.
Con.sider the rnbbit at your feet (Figure 6.1 0 ). Jt w ill p roject its image in your
lower visual field . TI1esmaller image of a more distant rabbit will be projected
higher in your visual field. Here, then, is another geometri c regularity produced
by projective geometry that the visual system can exploit for objects on the
grou nd plane, objects that are more d istant \-\·ill be h igher in the visu..-i.l field .
relative size A comparison of size

beti.,veen Items without kf'l:Y.Ning the
absolute size of ether one.
te><lure gradient A depth ct"
based on the geometrk::: fact that Items
of the same size form smeJler Images
when they are farther awa1. An array
of Items that change In slZe smoothly
across the image w ill appear to fam a
surface tilted In depth.
relative height "'3 a depth cue,
the observation that objects at dif-
ferent d istances from the 11ewer on
the ground plane will form Images at
different heig hts In the reti08J Image.
Objects farther away will be seen as FIGURE 6.8 This rabbit texture gradient shows that the size c ue is more effective
higher In the Image. wh'1fl size changes systematicalty.

FIGURE6.9 Organized differently, this Hlustratlon o f the same rabbits as those

shown in Figure 6.8 does not produoe the same s9nse o f depth. A slz8 cue Is most
effective when it Is conslsterrt with arranged on the ground, not on a wall.
Texture fields that provide an impression of three-dimensionality are

really combinations of relative size and relative height cues. Remember the
metaphor o f "perceptual committees" in Chapter 4. Different modules in the
visual system perform different tasks. The brain then combines the outputs
to come up with a decision about the state of the world . In the
FIGURE6.10 Relative height asa cue to depth. If we/re looklng down at the
ground planQ, the imag€1 o f a doser rabbit will tie abOV9 the image of a farth9r rabbit
In the retinal image. Because the image is invert9d, the closer rabbit Mes lower in the
visual fleld than the farther rabbit.

164 CHAPTER 6
.1a Appa
FIG URE 6. 11 The rabbit image at the top far left is the same size as th:i one at the
bottom far right. If they don't seem the same sizer then you have been fooled by the
d9pth CUQS.
case o f a tex:hue field, multiple cues interact to produce a final perception.

Figure 6.11 s hows how this interaction can give rise to a size illus ion . The
rabbit at the upper left of the fig u re is actually the s.ame physical size on the
page as the rabbit at the lower right, but the one at the bottom looks smalle r
to most of us than the one at the top . Why? We infer, on the basis of re lative
height, th3t the rabbit at the bottom must be d oser. 1f it is closer and it forms
an image of the same size as the little rabbit at the top, it fo llo'\>VS that th e little
rabbit a t the bottom must be really little.
If we know what size some thing ought to be, tha t knowledge can be a
depth cue in its °""'n right. We infer that the \voman in Figure 6.12a is holdin g
her h."'lnd
. out at theend of an outs tretched arm. \'\'hy do we ma ke this guess?
One alternative is that s he's holding her hand near he r shoulder, as in Figure
6.12b. But if that were the case in Fig ure 6.12n, the hand would need to be a
(n) (b)
FIG URE 6. 12 The cue of familiar size. Tutt hand in (a} looks closer than the one in
{b) because we know how big hands should b9 rslative to heads.

SPACE PERCEPTION AN D BINOCULAR VISION 165
FIG URE 6.1 3 The metrbal cues o f size and height can give the visual sys-
tem rnore lnforrnaticn than a nonmetrical c ue like occlusion can. Not only does the
red sphg-e in this im..'"tQ9 appear to bei c los9St and the grei;;ln sphere fa1thest away,
but also the blue sphere is setn to be d oser to the red sphere than to the green
sphsr0 .
very big hand . He re, our know led ge of the normal relationship of hand s ize familiar size A depth cue based
to head s ize makes all the differen ce. This is the depth c ue of familiar size. en knowledge o f the typical size of
Recall tha t occlusion is a nonme trical cue, providing only depth order. The ct>)ects like humans or pennies.
relative size and relative height cues,especia.lly taken together, prov ide some relative metrical depth cue A
metrical information . This is illustrated in Figure 6.13, where the three balls depth cue that could specify, for
example. that object A Is twice as far
appear to lie a t measurable distances from each other. The blue ball seems ;:may as object B wnhout providing
closer to the red ball than to the green ball in depthr for ex.a mple. Relath·e s ize Information about the absolute dis-
and height d o n ot tell u s the exact distance to an object or between objects. tance to either A cr B.
These a re relative metrical depth cues. Familiar size, h owever, could be an absolute metrlcal depth cue A
absolute metrical OOpth cue. lf your vis ual syste m knew the actua l size o f depth c ue that provides quantlftable
an object and the visual ang le of the object's p rojection on the retina, it couJd Information about distance In the third
{at least in theory) calcula te the exact distan ce frorn object to eye. Jn practice, dimension (e.g., his nose sticks rut 4
however, even if you know that your fri end is 5 feet1 0 in ches tall, the visu al centlrmters In front of his face).
systern d oes no t see m to knO\v that fact with a p recision that would le t you
know he's standing exactly 12 feet away.
Aerial Perspective
ln addition to its implicit know led ge o f geometry and its learned kn owledge
of familiar size, the visua l system " knows'' about properties of the atmosphere.
The triangles in Figure 6.14a give only a faint sense of depth, if any. l11ey are
just an array of identical shapes . Adding some grayscale informati on, how-
ever, as in Figure 6.1 4b, provides th e impression o f something like a mm.m-
tain range receding in the distance. More the fainter m ountains
(triangles) m ay appear to be farthe r away than the darke r ones. d epth
cue at work here relies on an implicit understanding that li ght is scattere-d by mma Appa
the a tmosphere, and that more light is scattered w hen we look through more
i?ltmosphere. Th us, objects farther a\vay are subject to more scatter and a ppear
(b)
FIGU RE 6. 14 Th9 triang!Qs seem to reC9de into depth m ore in (b) than in {a}.

166 CHAPTER 6
FIG URE 6.15 A r9al-world exampl9

o f aerial P9fSpective. Scattering of tight
by the atrn::>sphere mak9s more-dis tant
featur9s appear hazy and bll1e .
haze or aerial perspective A depth fainte r and less d isti nct. Titls cue is kt10\.vn as haze, or aerial perspective. Fig-
c ue based on the Implicit understand- ure 6 .15 sh ows a real-world ex..1mpl e. Sh ort wavelengths (blue) are sca ttered
ing that light Is scattered by the atmo- more than med ium and long wavelengths. This is why the sky looks blue and
s phere. More light Is scatt€fed when
we look throt.gh mcre atmosph ere. why obj ects farther m ...·ay look no t o nly hazy, but bluish. The rea l image gives
Thus, more distant objects are subjsot a s tron ger sense of d epth. By now, it s ho uld be clea r tha t this effect results
to mae scatter and appear fainter, from the contribution of other d epth cues beyond the blue haze- fo r in.s ta nce?
bluer. and less dlsUnct. occlusion a nd the kn own size of boats.
line.er perspective A depth cue
based on the fact that lines that are Linear Perspective
parallel In the tliree-dlrnensklnal wa1d lt wo uld n ot be difficult to imagin e the six lines shO\vn in Figure 6 .16 to be a
will appear to OCflVerge In a two-
dirnmslonal Image.
s ke tch of the view out the Vt.indshield of a car m oving d own a road in a fla t
landscape . The depth cu e in this case-linear perspective-is based on the
rules tha t determine how lines in three-dimension al s pace are projected onto
a n ..·o-dimen.sional irnage. The core piece o f projective geometry in this case
is tha t lines tha t a re pa ralle l in the three-dimen siona l world w ill appear to
con verge in the hvo-dimen sional image, except w hen the para lle l lines lie in a
plane that is para lle l to the plane of the two-dimensional image. Artists of the
ltalian Ren.:1 is.sance are said to h ave'' discovered ,, linear perspecti ve. TI-Ult is not
quite right. Yom d og or cat knows about linea r perspec tive. What Renaissance
artists d iscovered was how to make the rules explicit, write them d own, and
hun linea.r pers pective into a method for generating realis tic d epth in other-
wise flat paintings. Filippo Di Ser Brune llesco (1 377-1446) is typica lly g iven
credit for bringin g linear perspective into Europea n a rt. Leon Ba ttista. Albe rti
(1404- 1474) wrote the first book on the top ic in 1435 (Alberti, 1970). Figure
6.17 shows a n example of w ha t cou ld be d one. This Arcltitectural View is
by Fran cesco di Giorgio a painter from the Italian ci ty of Siena.
FIGURE 6.16 Une ar perspective. If we titled this image .. Driving across Kan-
you would understand, OOcause the converging lines give the impression of
p orallGI lings receding toward th9 h orizon .

FIGURE 6.1 7 Architectl.lrol \li'q..v by Francesco di Giorgio Martini (1477), a very

clear €P<arnple of linear perspective. Paranel tines in thG image plane. such as the
lars at th9 front 0Qflt9r, riemaln parallel in the irnagei. Parallel lines that would r9C9d9 in
dt1)th in tt"'JQ thrge-din"\t!onsional world convqge to a vanishing point in the two-dimen-
sional image. (Staatliche Museen. Berlin.)
Painted a row1d 1477, it is not the greatest work of art_, but it shO\vs
the basic rules of perspective a t work. Para llel lines in the image plane, like
the two front center pillaIS, would be parallel in the world. Look at all those
other lines con vergi ng toward a vanishing point C learl)f these a re in.tended
to s how the parallel lines on the ground or on h,e s'i'des running
in depth_Of course, you can see occlusion, size, and texture cu es at w o rk, too.
Pictorial Depth Cues and Pictures

As befits a gro up of rules th at allowed Martini to paint hi s picture with such
good apparent d epth1 the depth cues discussed so fur a re known as pictorial
depth cues. These are the cues p rod uced by projection of the three-dimensional
\\'orld onto the hvo-dimens ion.al surface of the retina. A realis tic picture or
photograph is the result of projecting the three-dimensional world onto the
two-dimensional surface of film or canvas. 1Nhen that irn.age is from
the correct position, the retinal image (in one eye, at least) for med by the two-
dimensional picture w ill be the same as the retinal image tha t w ould h ave
been formed by the three-dilnensional world , an d hence we see depth in the
picture. In theo ry, this means tha t a picture should look correct fro1n only o ne,
precise viewing position. In fact, pictures look reasonable over quite a ran ge of
dews.. Were this not so, there would be o nly one good seat in the movie theater.
To correctly interpret the shapes of three-dimensiona l objects from two-
dimension al pictures, people take the orienta ti on of the flat s urface of the
image into accow1t. This a llows them to understand that the picture is, in
vanishing point The apparent
fact, a pich.rre and no t the real thlng; at the sa m e time, they can ca lculate an point at parallel lines reoodlr>J In
accmate impression of the thing that is p o rtrayed (Vishwanath, Girshik, and depth <XnVefge.
Banks, 2005). To illustrate this p oint, Marty Ba nks snapped the photo of one
pictorial depth cue A cue to
of thea utho rssta nding alongside a pictme of himself tha t is shown in Figure d istance or depth usoo by artists to
6. 18a; here the"pich.uein the p icture" appears reasonable. In Figure6.18b, the depict thrae-dlmensmal depth In two-
same picture is s hown stripped of its context Now i t appears quite dis torted. d lmooslonal pictures.

168 CHAPTER 6
(•) FIGURE 6 .1 8 Picture in a picture. (a) One of

the authors. Dennis Lwi, is seen standing next
to a photograph of himself. In this panel the
in the picture• appears reasonable. fP)
The framed picture. isolated without the context.
[)OQS the pbture appear dist orteid? (Courtgsy of
Dhanraj Vlshwanath and Martin S. Banks.)
(b)
In Figure 6.1&., om visual system ran compensate for the pel'l:ep tual d is tortion
because there is enou gh context to enable the >v;ewer to attribute the distortion
to the s lant of the picture surface.
llle technique known as anamorphosls, or anamorphic projection, illus-
h·ates that the ability to cope \.vi th distortion is limited. In anamorphic projection,
the mi es of linear perspective are pushed to an extreme. Now the projection of
three dimensions into two dimensions creates a hNo-cHmensiona1 image that is
only from an unusual vantage point (or sometimes with a ctu ved
mirror). TI1e res ults are k.nmvn as anam orphic art. As an exa mple, there is an
odd diagonal s me.:u in the lower center of Hans Holbein 's sixteenth-cen tury
anamorphosis or anamorphic pro- painting Tire Ambassadors (Figure 6.19a). lf you could put your eye in exactly
jection Use of the rules of linear per- the rig ht p ositi on to view the image, the smear would prove to be the s kull
spective to create a two-dimensional s how n in Figure 6 .19b. Despite its s ua:essful recovery of the shapes in Figure
knage so distorted tliat It looks cor- 6.1&, the \·i sual system ca nn ot use knowledge about s urfa ce orientation to
rect only \Mien viewed from a special
rompens.:'1 te for the d istortion in Figure 6.19. In our ovvn d ay, the sidewalk chalk
angle or with a mirror that counters the
distortion.
(a)
FIG URE 6.19 In 1633, Hans Hot·

bein painted the double portrait in (a)
\11/ith an odd objoct (/)) at the feet of
the t'NO men.

FIG URE 6.20 Modern-day anamorphic

art . (a) In this photograph, artBt Leon
Keer creattJs what appears to be a large
three-dirnrensional version of the classic
\1doo game, Pacman. But. as shown in
(j:J), this is just a clever bit of anam orphic
art . It is just a flat image that looks three-
dhnenslonal when viewed from the correct
position.
artist Leon Keer crea tes amazin g anamorphic iinages tha t look spectacula rly
real from the ri ght vantage point a nd spectacula rly d isto rte d from elsewhere
(Figure 6 .20). More on the role of pictorial depth cues in art can be fmmd in
Web Essay 6 .1: Making the lrnpliclt Explicit.
Motion Cues
Beyond the pictorial depth cues, a number of additiona l sources of information
are available to our visual system w hen we \•;ew rea l-\vor ld scene--cues tha t
cannot be reprod uced in a s ta tic two-dimension al pictu re. The first no n picto-
rial d epth cue we w ill discuss is motion parallax. To its power (and
to und erstand why photograp hs of the forest often don' t come out well), the
best thi n g to do is to go outside a nd lie w1der a tree. Gaze up in to the branches
and leaves \vi th one eye covered and your head stationary. You w-ill notice that
leaves and branches form a rela tively fla t text u re. You can see a U the details, motion parallax An Important
but you may have trouble deciding vvhe the r one little branch lies in front of depth c ue that Is based on head
or behind. anothe r. U you open the o the r eye, bin oc ula r s te reopsis (i ntroduced movement. The geometrtc Information
ea rl ier and d iscussed in deta il later in the chapter) \.vill allow the branches and cbtalned fran an f!>f0 In two different
p:E;itlons at two different ti mes Is eJml-
leaves to fill out a tluee-dim e n.sional volllll1 e that was lacki ng before. Close lar to the Information from ™> eyes In
the eye, and the volume collapses ag'1in. Now, move your head from sid e to d iffetent poeltlons in the head at the
si de, and m otion parallax will restore the st>nse of depth. same time.

170 CHAPTER 6
(a)
App a
Time
FIG URE 6.21 Mo tion parallax. As you b ok out the window of a moving train ,
oqects closer to you the flower in this illustration) shift position m ore quickly than
c:b obj9Cls farth.Qf" away (the tr"9) fro m on9 moment (a) t o th1:1 nQXt (b). This r9gularity
can b"' explo itf'ld as a depth a.e.
H ow d oes m o tion p rovid e a cue for d epth? S uppose you 're si tting on a
train, looking o ut the wind ow a t the countrysid e. At o ne instant you see the
scene sketched in Fi g ure 6.21a . A m o m e nt la te r, the scen e has ch anged to
the o ne in R gure 6.2 1b . N o tice that as ymu train m oved fro m le ft to right in
the figure, a ll the objects shifted from ri ght to left. But note that some things
s h.ifted more tha n others. Th e flower (Fin the figu re) m oved a l.most a ll the
way across your retinal inwge, the cow (C) m oved a much shorter
and the tree (T) hardly chan ged positions a:t all. The term para/fax refers to
the geometric relations hip revealed. h ere : when you change y oux vievvp oint
while rolling d ovim the tu1cks, obj ects closer to you s hift p osition m ore than
objects farther away. Of course, you d on' t need to be on a tra in to experien ce
motion pamlla xi jus t movin g your head w ill d o. TI1e geometric info rnmtion
obtained from a n eye in two d iffe rent posi tions at hvo d ifferent times (m o tion
para llax) is simila r to the information &om hvo eyes in different p ositio ns in
the head at the sa me time (binocular stereopsis) (Durgin et al., 1995i Rogers
and Colle tt, 1989),
Motion pa rallax p rovides rela tive m etrical in formati o n about how far
away objects are; and as the experiment with the tree bran ches proves, it can
prov ide a compelling sense o f d epth in some situations in which other cues
are n ot very effective. 1l1e d ow ns ide of moti on parallax is that it works only
if the head moves (j us t rnoving the eyes back and fo rth won ' t d o, as you can
eas ily prove to yourself). Now you know why a cat might bob its head back
nnd forth as it plans a s pecta.cular leap from the sofa to the table, Other mo tion
s ignal s produce in fom1ation ab out depth. For example, objects get bigger and
optic flow The pattern of apparent
motion of objects In a visual scene s maller as they get cl oser a nd farthe r awa)'i so an object that is simply gettin g
produced by the relatwe motrn bigger on a scree n ca n appear to be looming towa rd you. If everything is
between the oboorver and the scene. looming at once in a large field .. this optic flow may rna ke you feel like y ou

(11) Con\'ergence (b) Oi\•ergem:e

•
a Gets bigger.
Convergence
Oi\•ergence
a Gets smaller.
FIGURE 6.22 (a) As WQ shift foa..1s from a far to a point, our 9yes
oonvgge, (b) As we go from near to fur, th.a e')"9S diverge. Th€1 sizie o1 the angle
(labek3d Qa") is a cue to depth .
<Jre m o\oi ng toward the screen rather than like the objects on the scree n are
tnoving toward you. These top ics are discussed in more detail in 01t1pter 8.
Accommodation and Convergence Appa

Like a ca mera, the eyes need to be focused to see objects at different disUmces
clearly. As we learned in Chapter 2, the human eye focuses via a process called
accommodation, in whid1 the lens gets fatter as we d irect our g.nze toward
nearer objects. We aJso need to point o ur eyes differently to focus on OOjects
.at different d is ta n ces. As the schema tic eyeba lls in Figu re 6.22 move from the
red dot to the blue dot, they rotate inward-a process ca lled convergence
(Figure 6.22n ); refocusing on the red dot would require rotation outward,
whid1 is known as divergence (Figu re 6.22b). accommodation The prooess by
If we could monitor our s tate o f accomm odation and / or th e extent to which the aye chan;ias Its focus On
w hich our eyes were converged, we could use this information as a cue to w hich tha kins gets fatter as gaze Is
the depth of the object we were tryin g to bring into focus: the rnore we have directed toward nearer dljoots).
to con verge and the more the lens has to bulge in o rder to focus on the object, convergence The ablllty of tha
the closer it is. ln fact, we d o use this infom1ation . When we focus on objects two to turn inward. often used
more than about 2-3 meters away, the lens is as thin as it can get and the in crder to place the two images of a
feature In the wor1d on correspond -
eyes are diverged about as much as possible, so nei ther cue provid es much
ing looatlons In the mo retinal Images
useful information. But carefu l s tudies have s h o""n that the vis ual sys tem (typically on the fovea of each eye).
takes advantage o f both cues for objects closer than this limit. Convergen ce Conv€<gence reduces the disparity of
is used more than accommodation (Fisher a nd Ci uffreda, 1988; Owens, 1987). that feature to zero (cr nearly zero).
Moreover, in principle these cues can tell us the exact distance to a n object. divergence lhe ability of the t wo
Humans are not particularly precise about measuring the exact angles s how n eyes to turn outward, often usecl In
in Figure 6.22, Chameleons, on the oth er hand, do use the absolute metrical crder to place the two lrna(,les of a
d epth infonnatio n frorn con vergen ce to ca td l prey insects \'Vi.th their sticky feature In the wor1d on correspon::l-
lng lccatlons In retlnal Images
ton gues. Harkn ess (1977} sh owed this by fitting a cha me leon with glasses that on the fovea of each eye).
d istorted the angle of convergence. 1l1e result was that the poor chameleon o;,ergerce reduces the disparity o f
flicked out its tongue to the \vrong distance and missed its intended dinner. that feature to zero (cr nearly ZHO).

172 CHAPTER 6
FIGU RE 6.23 (a) (b)
•
soen9 Illustrates how geometric
n9gularitii9S ar'9 by tl').e visu<ll
systr::m to achle\19 st ereopsis from b in -
ocular disparity. (a) The \llq1A19r, Bob, is
assumed to be fixing his gaze on the
ried crayon . (/;) This t op \1ew traces the
rays of lig ht boun cing o ff the red crayon
onto Bob's retinas.
•
•
a·
Binocular Vision and Stereopsis
As defined earlier, the tem1 bh1oculnr di<Spnrity refers to differences betv"·een
the images fa lling on mu h•m retinas, stereopsis refers to rl1e impression of
th.ree-.di mensiona li ty---of objects " p opping out in dep th"-tha t most h tunan s
ge t w hen they view real-world objects with both eyes. Like the acco lmt.s o f
othel' dep th cues, the s tory of the route from b inoc ular disparity to s tereop sis
is a s tory of the visu al syste m exp loiting the regula rities of projective geom-
etry to recover the three-dimens ional world from its projections-this time,
onto a pair of two-d im ensional surfaces. \ Ve w ill illustrate the transJation from
d ispority to stereopsis usin g the s itua tion shown i n Figure 6.23a, in w hid1
the viewer (call him Bob) is facing a scene that includes fou r colored crayons
a t differen t dep ths. Suppose tha t Bob is focusin g his gaze on the red cr3yon,
as show n in Figure 6.23b. The two lines in this fig ure trace the pa ths of th e
light r<1ys tha t reflect off the red crayon an d onto Bob's two retinas. (Sim ilar
experiences \vith crayon scenes are also demonstrated in Web Acttvlty 6.2:
Binocular Disparity.)
Beca use the visu al system is d esigned so tha t the object of our gaze ab..wys
full s on the fovea, the rays fr om the red crayon fall on the fovea in each of
Bob's eyes. Figure 6.24 s h ows the retin a l image for the crayons in each eye.
The red crayon is in the center of both images. \ \le'veadded a das hed vertica l
FIG URE 6.24 The ovi9flapping por-

tions of the images falling on Bob 's left
and right re tinas. Because the retinal
image is re\19rsed, the blue and p...1rpe
c rayons on the right side of the scene
in Rgure 6.23 projQCt to the left sid9 of
each re tina. whereas the bro"Wn cra-fon
on the left side of the scen9 projec ts t o
the right sid9 of each retina The siz9
d ifferences between the retinal im ages
o f thg crayons in the two retinas ar9
exaggerati:id in this figure com pared
w ith the differences we would observe
if we saw this scene in the real world . Left retin al im age Right retinal im age

• FIG URE 6.25 Bob is still gazing at

the red craycn. Th is \'iew from above
traces the light rays reflecting from
the red and b lue crayons onto Bob's
retinas. The btua crayon projects to
Horopter (Yteth.-Miiller ci:n:le)
C01T9SPOnding retinal pdnts-posi-
tlons that ;:)(0 equidistant from and on
the same side of the fovea The same
wouk:I b9 tru9 o f any object falling on
the gray c urve shown in the figure. (The
horopter and Vieth-MUiier c ircle are not
exactly the sarnc;i, but they would 00
w:.<y slmUar in this case.)
8230336 AmMa Appa
line in fro nt of this crayon in each image, to emphasize the fact tha t this is the
location of the fovea.
Now consider the retinal images of the blue crayon .As you saw in Chapter
2, the optics of the eye reverse left-right and up-dmvn (see Figure 2.2). Thus,
the blue crayon on the right side of the scene in Figure 6.23 fa lls o n the left
si de in each of the hvo retinal images s ho\o\.n in Figure 6.24. In our imaginary
scene, the b lue cr1Jyon is p laced so tha t the m onocular retina l im:.ages of that
crayon are fonned at the s.,1 me distan ce from the fovea in both eyes. \Ve say
tha t this crayon's images fall on corresponding retlnal points. The same can
be said of the images of the red crayon, ""hk h full on the two foveas.
In fact, a ny object l};ng on the Vieth-MOiier circle-the im agi..na ry circle
tha t nms through the two eyeballs and the object on w hich Bob is fixated-
sh ould p roject to corresp ond ing retinal poin ts. This imaginary circle is dra.wn
in g ray in Figure 6.25. Objects that faU on corresp onding re tinal poin ts are
said to have zero binocular disparity. H the tvrn eyes are lookin g a t one s p ot corresponding reUnal points
(such as the red crayon), then there will be a s urface of zero d isparity nmning Two monocular Images of an cbject
through tha t spot. Tha t s urface is knmvn as the horopter. Any object placed In the world are seJd to feJI on ccrre-
on that irnaginary s urface in the world w ill fo rm images on correspond ing spondlng pdnts If th<>le points are the
same distance from the fovea In both
retini.ll points. As it happens, the horopter a nd the Vieth-Millie r d rde are n ot eyes. The 1'No foveas are aJso co1re-
quite the sarne. lf you a re extremely fond of rather complicated you spondlng pdnts.
may want to pursue this topic in one of the follow ing sources: L P. Howard
Vieth-MOiier circle ll1e locatlcn
and Rogers, 1995 or 200 li or Tyler, 1991. O then.vise, the imp ortant p oint is of objects whose images fall on geo-
tha t there is a surface of zero disparity whose p osition in the world depends metrk:eJo/ corrooponding pdnts In the
on. the current s ta te of convergen ce of the eyes, two retinas. If life were simple. this
O,jects that lie on the horopter are seen as sing le objects when viewed '"rith c irde would be the horopter, but life is
bo th eyes. Objects significantly closer to or farthe r away from the surface of not simple.
zero disparity fo rm images on d ecid ed ly noncorresponding points in the hvo horopter ll1e loca!lcn of objects
eyes, and we see two of e..1 ch of those objects. Thi s d ouble vision is known as whose Images lie on corresponding
pdnts. The surface of zero disparity.
diplopla. Objects that are dose to the horopter but n ot quite on it can s till be
seen as sin gle objects. This region of space in fron t o f and behind the h.oropte r, dlplopla Double vision. \'lslble
In both eyes, stImu Ii falll ng outside
witl1m w hich binocul ar single vision is possible, is known as Panum's fuslonal
of Panum's fuslonal area will appear
area (Pan tun, 1940). You can check this quite simpl y, by ho lding a red crayon dlploplc.
(or pen ) directly in front of you with your left hand, at a distance of about 20
Parum's fusional area The region
crn,a nd keepin g both eye.son it. Now hold a blue crayon {m p en) about S cm of space, in tront of and behind the
to either side o f the red one with yo m right hand, and slowly move it nearer horoptE!f , within which binocu Br' single
to your eyes and then farth e r away, while maintnin careful fixation on the red vision Is possible.

174 CHAPTER 6
FIG URE 6.26 Light rays proj9cting (a) (b)
•
from th.a brown (a) and purple(/)) cray-
on s onto Bob's retinas as hQ epntinu9s
t o gaze at the red crayon.
one.. You sh o uld initially see the blue crayon / pen as single, w hen i t is about the
sa me dis tance fro m you as the red o ne, because it is \\•i thin Panum's fusi o na l
area. Hmvever, w hen it falls outside Panum's it will appear d ouble.
PanumJs area provides a little roorn. for s mall errors in eye alignrnent, w hile
sti ll maintaining s ingle vision.
Am1ed 1ivith th is tenninology, let's return to Bob and his crayons. Consider
the retinal images of the brov•m crayon, lying just off the horopter. As Figul'e
6.24 a nd the view from above in 6.26a s how, rays of light bouncing off
th.is crayon d o not fall o n corresponding retinal points: the crayon's image is
farthe r away from the fovea on th5 lef.tµotina than on Refati ve
to the horop ter, this crayon foh:ris -re\tn'31 -lm;:tg with a h 'o nzero binocular
d isparity. The purple crayon is even fur ther off the horopter (Figure 6.26b); it
forms retinal images that a re even more disparate (Figure 6 .27).
1lle geomehicregularity that the visual system uses toe>.trnct metriral depth
information from binocular disparity should now be growing 111e larger
the disparity, the greater the distance in depth of the object fro m the horopter.
Land R R LandR R L
FIG URE 6.27 Supa-pooition of Bob's left (Wand right (R) retinal
images of the crayons in Figure 6.24, showing the r9lattv9 dispar·
lty tor each crayon. Size differences are ignored here. The red and
blue c rayons sit on the horopter and have zero d isparity. They form f------71
retinal irnagoo in ccnespcnding locations. Th9 brown crayon forms H
Images with a small binocular disparity. The p.;rpe crayo1l, farther Zero Big Zero Snl.:,u
fro m the ho ro pter, llas larger binocular disparity. disparity disparity disparity disparity

(a) Cros&ed di.spa1ity (b) Unc.roS1Sed disparity FIG URE 6.28 C rossed and
uncrossed disparity, '8) Here Bob is
foveatlng the red crayon. In the Bob's-
f,flfe \'iQWS, the clOSGr, blue object is
:' \ S9efl to the right In t1·19 left eye and to
the l13'ft in the right QYQ. This situatic:n
//\'\
is crossed disparity. Pl Here Bob h:;tS
shifted his gaze and his horopter to thtio
blue crayon. In the Bob's-eye views,
the farthtf, reel object is. s..een to the
dJ 0
left In left eye and to the right In the
right eye. This situation is LJncrossed
dlspanl'f.
Disparate blue bar: sparate red b.ln pa

Right in left eye, left in right JUsht in right eye, left in ieft
The di rection in depth is given by the sign (that is, "crossed " or " Wlcrossed ") crossed disparity The sign of dls-
o f the disparity, as illustra ted in Figure 6.28. Suppose that Bob i s l ooking at a P"'ltY created by objects In front of the
red crayon with his eyes converged so that the red crayon fulls on the fovea plane o f fixation (the Tl1e
term crossed Is used because Images
in e,1ch eye. A doser, blue crayon w ill form images on noncorresponding, of objects located In front of the raop-
d isparate p o ints. On the le ft retin a, blue \vilJ lie to the left of red . Because the ter appear to be displaced to the left
im age is reversed, thi s means tlmt, viev..,ed from the lef t eye, blue is to the right In the right aye, and to the right In the
of red Viewed from the right eye, blue is to the left. Right in left, and le ft in left aye.
right. This is kno",' as crossed disparity (Figure 6.28a), and crossed d isparity uncrossed disparity lh9 sign of
al ways m eans "in front of the horopter." ln Figure 6.28b, Bob is lookin g a t the disparity c reated by objects behind
blue cr.:iyon, and the red is seen to the left \Vith the leit eye and to the right th& plane of fixation 110ropt€<).
The term uncrossed Is used because
w ith the right. T11a t's uncrossed disparity, and uncrossed disparity ahvays
Images of objects located behind the
means '1Je1Und the h oropter." horopt« will appear to be displaced to
1110 right In ti... right eye. and to the left
Stereoscopes and Stereograms In the left fffe.
Inte resting ly, although scientis ts had s h1died the geometry of binocular vi- stereoscope A device for simulta·
sion for millennia (the geometer Eudjd was at it in the third cenh1ry BCE), neousty presenting one Image to cne
not until the nine teenth cenhuy \1'1 as binocula r d isparity properly recognized aye and anoth€< Image to the other
as a depth cue. [n the 1830s, Sir Charles Wheats tone invented a device called f!J'fe. Stereoscopes can be used to
preoont dlc hoptlc stimuli fa steroopsls
the stereoscope (Figure 6.29) tha t presented one image to on e eye and a dif- and binocular rivalry.
FIGUR E 6.29 Wheatstone's sterooscope. Tug v iewer

would bring h9r n ose up the verticul rod at the CGflt'91'"
of the apparatus so that each gye was looking at the
image reflected in one of the two mirrors. (From I. P .
Howard and Rogers, 1905.)

176 CHAPTER 6 8230331_; Amma Appa
(b)
FIG URE 6.30 Steroopsls for thi:i masses. (a) This Holmoo stereosccpe -among
others-brought st91'"90 photos into many mid-nin9t99nth-0Qfltury hom'1s. sh11-
rieo photo of South African Light Horse. a scouting regiment o f thiB British Army, on
Adderly Slr99t In Cape Town, South Afrba. in 1900. If you can freE1-fuse (9.Xplain9d
latiEW· In this secUon), you will be able to si:i.e this scene jump out in depth,
ferent image to the other eye. The stereoscope proved tl1'.1t the visual system
treats binocu lar d isp•nity as a depth cue, regardless of whether the d isparity
is produced by adual or simulated images of a scene.
For the average citizen the s tereoscop e was not science; it was hom e en-
tertainment. The \Vheatstone s tereoscope he ld two diffe ren t hnages in two
different places. In the 1850s, however, David Brews te r and Oliver \ Vendell
Holmes invented viewers (Figure6.3oa) that held a card with a d o uble image
like that sh own in Ftgure 6.30b. The double images '"'ere cap tured by cameras
""-i th two lenses separated by about 2 ind1es, the di stance between the nver-
age htunnn 's eyes. This arra ngement allows s tereo cam eras to take a pair of
pictures th.n tmi.mic the images produced by the projective geometry of human
binocular vision Photogr..1phers tra\.·eled the ""Tor Id \>vi th these stereo rn meras,
c.a phuing far-off scenes in a way tha t enabled a London schoolchild to see, for
example, a vivi d three-dimensional image of the British Am1y in Cape Town,
South Africa. (For a guide to more of these historical im:.ages, as well as many
o ther s tereogr-an1s, see Web Essay 6 .2: Stereo Images on the Web.)
A stereoscope is very he lpfot but you don 1t actually n eed one to experience
stereopsis. You c.m teach yourself a technique knm.vn as tree fusion, whid1
John Frisby (1980) called " the poor man's stereoscope." Figure 6.31 contains
two a lmost identical pictures. U you cross yom eyes hard1 you sh ould see four
sets of squares. Titls is the phenomenon of d ouble vision (diplopia), described
in the p revious sectio n. Two of those sets are seen with the left eye and hvo
with the right . The trick is to relax jus t a bit until you see just three sets of
squares. The far- left set is seen o nly in the leh eye; th e far-right set, only in
the right; but the middle set is the fusion of h ·VO sets, on e seen by the left eye
and another seen by the right eye. 1ltis fusion of the separate images seen by
the two eye:s: makes s te reopsis p ossible.
Ad1leving the perception of three, instead of fom or two, sets of squares
in Figme 6.31 is the first step toward free fosi on.. The second s tep is to bring
the middle set into foc us. As n oted ear lie 1:. con vergence and accommodation
free fusion The technique of con-
verging (crossing) or diverging the "'fOO normally work in lockstep, so crossing ymu eyes automatically lead s your
In order to v1011i" a stereogram wtthout a ciliary muscles to rnake your lenses more s phericaJ (im.less you are presby-
sta-eoscope. op ic--see C hapter 2). Similar problems (in the opp osite direction) will occur

FIGURE 6.31 Try to conv9rge {cross)

or div9rge (uncross) your €1'/00 so that
r. ' you seei exactty three big blue squa roo
h9re, rath91" than the two on the page. If
you succ9iEld, you \Nill probabt'f be atje
to see that the three little l/\ihite squares
lie at different depths in the second o f
those three big squares.
if you dive rge your eyes. To see the middle set of squares demly, you h ave
to decoupl e accommodation and convergence. If yo u ca n m anage it1 then the
image \'\<ill com e into focu s and the three w hite squares wtll appear to lie a t
differen t depths in the middle set of squares. \ Vhen you \.iew them normaU y,
no tice that the w hite s qua res in the le ft and right panels look misali gn ed in
opposite directions. Those are the m on ocular v iews. When yo u free-fuse,
the opp osi te misalignrn_e nts become the binocitlar disparity, a nd your visual
systern converts that disparity into a perception o f depth.
The depth that yo u see depends on whether you converged or diverged
your eyes. We de.scribed crossing, or con verging, the eyes. It is also possiWe to
free-fuse the images in Figure 6.31 by dil:rerging yo ur eyes. Divergence requires
focusing on a point beyond the p lane of the page so that the im<1ge of the left-
hand set of squares falls on the left fovea and the image of the right-hand set
falls on the right fovea. Because the im ages fullin g on the n.vo retinas in the
di vergen ce me thod are reve rsed compared with the con vergen ce s ituation ,
the d isparities are reversed a nd the perceived d epth will be 1,.£...yqu
converge, the top square v.,.ilJ be the farthest back. lf you diverge, it \Vill appear
1maAppa
closest to you . Either con verging or diverging will produce n elem stereoscopic
effect, so give it a try.
Before we go on , we sho uld note that approximately 3--5% of the population
lacks s tereoscopic d ep th p e rcepti on-a. condition knmvn as stereobllndness.
Stereoblind individuals rn.ight be able to achieve the pe rception of three sets of
squares in Figure 6.31, but the little w hite squa!'es \vill not p op out in depth .
Stereoblindness is u s ually a secondary effect of childhood visual disorders
s uch as stra bis mus, in w h ich the two eyes are m isaligned . See the sectio n
"Recovering Stereo Vision " below. lf you had s uch a visual d isorde r during
d1ildhood and / or you 've been diagnosed with stereoblindnes.s, we apologize
but you jus t won't perceive depth in the stereogra ms presented here and on the
website. ll1at said, many people who try and fail to see d epth in s tereograrns
h ave " normal" vision (wearing glasses d oesn' t cotmt as ''abnormal" in th is
case). Those people just need practice, so d on't give up . Web Activity 6 .3:
stereoscopes and Stereograms pro,·id es more stereograms for practice,
stereobllndness An Inability to
and Web Essay 6 .2: Stereo Images on the Web leads to another web site make use of binocular disparity as a
w ith more tips for free-fos ing. depth cue. This term Is typically usoo
to describe Individuals with vision in
FURTH ER DISCUSSION of strabismus can be found in Oiapter 3 on both eyes. Somecne who has lost one
page 86. (Cf both) eyes Is not typically descrlbOO
as ''Stereobllnd."

178 CHAPTER 6
Recovering Stereo Vision

As noted earlier, about 3-5% of the population are stereobllnd, usually
due to earty childhood visual disorders. Can steroo \1sion be recovered
later in life? Meet Stereo Sue and Stereo Bruce.
Susan Bany, a professor of neurobiology, had strabismus as an in-
fant and had been stereoblird essentially all her life. Her book FiXJi1g My
Gaze pro1;des a fascinating, Informative. and beautifUlly written account
of her ao::::iuisltlon of steroopsis following vision therapy. It describes her
transfonnattve journey from the many visual, social. and psycholcgical
dlallenges of a turned eye (a sq uint or strabismus) early In life. to the
sudden enrichment of her perceptions of the world follovAng success-
ful unoonventional visual therapy begun at 48 years of age. (An earlier
article about her visual recovery i.vas published In The New 'i'brker under
the of "Stereo Sue" by 01;,,er Sacks.)
B'lrryvMdly reoou nts how acquiring stereoscopic vision led to
a dramatic improvement of her perception of depth. cr the apprecia-
tion of "the si:ace between" objects. A partlcular1y valuable insight is
her argument for the inability of pecp/e vAth normal vision to appreci-
ate the visual experience of being stereobllnd. NaNely one might think
t11at th<; experience would be duplicated slmr:;ly by c losing one eye so
all info!Ttlation abrut depth was oorrveyed by mmcx::::ular cues. Not so,
however, Bany argues: the monocular experience of a typically reared
person who c loses one eye has been Informed by a lifetime of exper1-
aice wlth stereoscopic vision and so ls far different from that of a person
who is As a result, Barry's new stereosooplc vision brought
much more to her life than Just depth perceptio n: Ol::jects became
dearer, motion perception more veridical, her ability to move around the
WC<1d more confident. Even more pdgnant Is her vMd description of the
enhanced sense of touch she had developed over the years and Its key
role in informing her newly acquired sense of stereo vision. Bany did not
simply "recover· stereopsls bLrt rather had to releam to see with stereo
vision. As blind or deaf indi>Aduals·often descrt>e, !J\dMdual of
a sense are not just .,missing" a sense. Rather, the\/have develOped an
entirely different '""I of oonslng the wood . Upon sensory restitution, a
fascinating but rather disturbing experience unfolds as the brain has to
adapt to a new way to function.
Even more dramatic is the experience of Stereo Bn..ce (Bruoo
Bridgeman). a very perceptive vision scientist who had been stereo-
deficient all his life. Remarkably, he recovered stereopsls attar watching
the 30 movie Hugo (Bridgeman , 2014). Whether this sort of immersive
experience. with very large dispsrities along with many other depth cues,
v.; 11 be a generally effectr,,e treatment for abnormal stereopsis remains to
be tested. However, these caoo stLdies, along v.;t h lab studies of per-
ceptual l6"mlng that result in the reoovery of stereopsls (Di ng ard Levi,
2011 ), call into question the notion that has been the received wisdom.
that rew1ery of stereopsis can only occur during early childhood. The
Idea, dating back to the early twentieth century, has been that there is a
"crttical period" of dev8oprnent wtien the visual system is still plastic and
capatle of change. After ttat . it was thooght. our basic \'isual capabilities
are fixed. This led a number of practitioners to tell Susan Bany and her
mother that "nothin;J could be done" about her >Asian (one suggested
that she might need a psychiatrist). binocular neurons are pres-
ent in the visual oortex of primates wit hin the first v.ieek of life (see the
"Development of Bi nocular Vision and Stereopsls" section below), Bany
surmises that some of the innate 1,viring of her binocular connections
remaina::l Intact , and that vision therapy taught her to move her eyes into
position for stereo visi on, 4 flnally giving these reurons the infonnation
they were >Mred to receive" (Bany. 2009) .

FIG URE 6.3 2 A ster80 photograph

o f a woman's face.
Random Dot Stereograms

Fol' 100 years or so after the invention of the stereoscop e, it was generally random dot stereogram (RDS) A
supposed that s te reopsis ocn1rred relative ly late in the p rocessing of visual stE!feogram made of a large number
(otten In the thousands) of randomly
stimuli. The id ea was that the first s tep in free-fusi ng images such as those in
placed dots. Random dot stereograrns
Figure 6.32 wo uld be to analyze the input as a face. We would then use the contain no monocular cues to depth.
slight d isparities be tween the left-eye and right-eye ima ges of the n ose, eyes, Stimuli vtslble stereoscopically In ran-
chin, and other objects and p a rts to enrich the sen se tha t the n ose s ticks out dcm dot stereograms are Cyclopean
in front of the fa ce, that the eyes are slightly s unke n, and so on. stimuli.
Bela Jule.sz, a Hungarian radar engineer w h o sp ent m ost of his career a t
Bell Labs in Ne w Jersey, thought the conventi onal v.risd om might be backward .
He theorized that s tereopsis might be used to discorer objects a nd s ur faces in
the world . Why wo uld this be usef ul ? Julesz thought tha t stereopsis might
help revea l objects. A m ouse might be the sam e color as its
background, but out in the o pen it would be in front of the bdckgro und . A
ca t tha t could use s tereopsis to break the m ouse's ca mouflage would be a
m ore s uccessful hunter. (Cats d o have s tereopsis, by the way [Blake, 1988;
R. Pox and Blake, 1971] .) To p rove his point, )ltlesz (1964, 1971 ) made use of
random dot stereograms (RDSs). A n exa mple is s hmvn in Figure 6 .33. 1f
you ca n free-fuse these images, yo u will see a p a ir of squa res, o ne s tickin g
out like a bump, the oth er lookin g like a h ole in the tex ture twhich one is
the bump and w hich is th e h ole dep en ds, again, on whether you converge
or di verge your eyes).
FIG URE 6.33 If you can free-fuse. th is

random d ot ster90gram, you VYiH seg
two rgciangular re gion s.: on9 in front of
the plan e of the page, the o ther behind
th'11 page. Which is which d eplWlds on
w heithe- you conV€<ge or diverg e In
order to fuse th.e two squar01S.

180 CHAPTER 6
Cyclopean Referring to stimuli llle importan pofut abo ufR i s tha \Ve cannot see the squares in either
that are delned by binocular dlepar- of the componen t images. We cannot see the sq uares u sing any monocular
lty alone. Named after the one-eyed depth. . a.1es. 111ese are shapes thn t a.re defined by binocular disparity alone.
Cycops of Homers O:tyssey.
Julesz called sudt stim uli Cyclopean,, after the Cyclops of Homer's
Odyssey. Wheatstone s howed wi th his s tereoscope thn. t binocular disparity is
a necessary conditi on for stereopsis. Julesz demonstrated t\-•ith the RDS that
disparity iss11Jficient for stereopsis. To \.mderstand how RDSs are m ade, visit Web
ActMty6.3: stereoscopes and Stereograms, whid1 also explains 11magiceye''
and 11wallpaper" s te.reogra:m s, know n m ore as ''a utostereogram s."
Stereo Movies, 7Y, and Video Games

Whereas the Brevvster stereoscope provided h om e entertainment in Victorian
times, in our d ay 3D movies, TV, a nd d deo games are the n ew s tereoscope.
Achrnll}; 30 m ovies h ave been around for more than 60 years. In general, 30
mov ies make use of s tereoscopic photography (which records the images as
seen from two slightly diffel'ent perspectives) a nd 30 g lasses (whid1 sepa-
rate the images to the hvo eyes, so tha t dispa ri ty provi des additi on al cu es
to d epth ). Some of the ea rlies t 3D mov ies, were viewed \vi th 1'an aglyphic"
glasses, with a red lens for one eye a nd a blu e or green filter for the o ther eye
(Figure 6 .34). When the images on the screen we re filtered through the diffe r-
ently colored filters, one eye saw one set of images, and the o ther eye S..'lW the
other set. Yo u've may ha\·e seen the an aglyph.ic me thod used in comic books.
La te r, fi lmmakers used p o briw ti on to separate the irnages to the two eyes,
and p ola rizing glasses were used to "·iew the films. A recent advance on the
anag lyph ic m ethod is the use ofinterference filters to present s pecific wave-
lengths of red, green, and blue to the right eye, <'l nd different wavele ng ths of
red , g reen, and blue to the left eye. Very expensive goggles that filte r out the
specific wavelengths enable the filmgoel' to three-dimensiona l image and
enable the movie thea te r to cha rge a few extra d ollars per ticket.
Man y modem 30 m ovies, T\ j and video gam es use special glasses that
contain liquid crystal d isplay technology (known as LCD shu tter glasses) and
FIGURE 6.34 An audierlcQ watching a st9r49o rTl0\.149 in the 1Q50s.

FIGURE 6.35 It Is possible gffQCtiVe stQl'"eoscopic imaQQs of terrain by tak-

ing tvilo aerial from t\NO, quite w idely separated viewpoints.
e nable the p resentation of images to each eye tha t a re synduoni zed with the
images on TV, or computer screen. In this metho d, alte rna te fra mes
of the rnovie are presen ted to different eyes. All of the systems described here
have the d rawback that they require special glasses or goggles to see in three
d im e nsion s. H mvever, ce rtain clever m e thods d o no t requi re g lasses . For
example, in a m e thod knovrn as le nticuJaJ printing, the images a re digitally
split and interleaved (left, right, left, right, ... )at a fixed spaci ng. A special
array of man y tiny lenses placed on the screen ens u res tha t the left eye sees
only the 11.eft'' image and the right eye sees only the "right '' imL'lge. This setup
permits three-dimension ril viewin g w itho ut glasses or o ther p arrip hernalia.
The d eve-lopme nt for the cons ume r ma rke t of vi rtual reality (V R) technology
is makin g 30 gam es and m ov ies more widely availab le. YR technQJbgy ma Appa
ables the user to be imme rsed in a v irtua1 30 world . I ts comme rcial potential
was evi dent w·he n, in 2014, Facebook bough t O culus, the VR company tha t
d eveloped the p ioneering O.:uh.1s Rift system _
Using Binocular Stereopsis

The appli ca ti on s of s tereopsis go well beyond ente rta inmer1t. The rnilitary
h as kt1 0'"'11 for a lon g time that you can get m ore UUormation out of ae rial
surveillan ce if yo ur view of the g rotmd is s tereosco pic. However, if you ' ve
ever looked out the Yvind o,,,. from tho usands of feet in the air, you may have
n oticed the g round looks rathe r flat. This is yet m ore geome try. Binocula r
s tereop s is can provide useful info rma tion a bout me tric d ep th only for dis-
tances up to 40 m eters (Pahnisano et al., 2010). \.Yith eyes a few cen timeters
apa rt, you d on ' t get adequa te dispari ty from more dis ta nt targets. \ Vhat you
need are eyes sepa rated by hundred s of feet. Titls can be done if you have a
plane and a special cam e ra . Figure 6.35 reprin ts a fi gu re from a 1951 issue of
Popufar Mecliauios in w hich Colon el George W. Godd ard showed the public
how images taken from ti,.1,.-0 vantage po ints produ ced s te reo images durin g
the Korean Wa r. \.Ye ca n also use stereop sis to have a be tter look ins ide the
bo dy. Figure 6.36 sh ows a s te reo pair of mammograins, X-rays o f the breast
used to d etect breast can cer. Actually, in order to crea te an othe r free.. fus ion
d emonstration, we' ve prin ted the image of the left breast twice, to the left and
the to right of the irnage o f the rig ht breast in th e middl e. lf you free-fuse the
images so tha t you see four breast one of the t\vo renter images w ill

182 CHAPTER 6
FIG URE 6.36 Thisster19omam-

rnogram was c reiatQd by taking X-rays
of a woman's breast frOn1 rm the correct disparities (depending on whether you diverge or con verge
points. If you can free-fuse, you will see
your eyes to free-fuse). The correct one \•,.. ill show the w hi te wire-a marker
the structures in the lma.ge seoparnte
in depth, m3king it easlsr to deddQ for the s urgeon--on top o f the breas t. You can see tha t the breas t tissue is
wheitl"lef th.a filament is part o f the chilrac terized by a ne h vork of intersecting s tructures. This is like our earlie r
breast tissue or is a diffgr90t structur9, exam ple of lookin g a t the little brand1es of a tree w hile lying on your bade
(Courteey of David Getty.) wi th just one eye open . It ca n be very ha rd to tell w hich line-like s tructures
actually intersect and whid1 o nes lie at different depths . This turns o ut to be
important w hen reading a mammogram. A s tarburs t structure might be a s ign
of can cer, but not if it is a n accidental vielvpoint (see Chapter 4) of s tructures
at diffe rent depths in the breast th:Jt jus t happen to form a s us picious pattem
in a hovo-dimen sional projecti on . Stereopsis can d is.a.mbigua te this s ituation. If
you can free-fuse these images, you will see the texture in three dimensions,
and you 'vill be able to detennine how different s truc tures rela te to each othe 1·
in depth. S tereoscopic displays are beginning to be used in radiology (Held
and Hu i, 2011 ), and they can reduce the e rror rate in these import.1nt tasks
(Ge tty, D'Orsi, and Pickett, 2008).
correspondence problem In bin- Is, binoc ular s tereops is useful in everyday life? In p eople wi th normal
ocular vision. the problem of figuring binocular vision, "isually guided hand m ovements are s ignificantl y impaired
out which bit of the Image In the lett when dewing is restricted to one eye (Field er and Moselyi 1996), likely mving
f!Ye sh:.<1ld be matched with which bit
In the right eye. The problem Is partioo- to the fact binocular deptl1 thres hold s ru·e about factor of 10 better than
larly vexing when the Images consist of monocular thresh olds (McKee and Tavlor, 2010). These results are mirrored
thousands of similar features, like dots in patients v.'ith amblyopia ("lazy eye;;.) for w hom many observed visuomo-
In random dot stereograrns. tor deficits are due to impaired stereopsis, and in particular irnpaire<l visua l
feed back contro l of moveme nts, rather than visual acuity loss (Grant and
Moseley, 2011). Loss of stereop sis rn.ay also result in tmstable gait, especially
reduced accuracy w hen a change of terrain (e.g., s teps) occu rs, ruid difficulties
for children in playing some sports.
FURTHER DISCUSSION of stet""eo sensation can be frurd in Chapter 10

(sound localization: pages 2{J1-2Q5) and Oiapter 14 (binaral rivalry in olfac-
tion; page 443).
Stereoscopic Correspondence
•••
If you s uccessfully free--fused the rand orn d ot patterns in Figure 6.33, yo u
solved a truly da\.mti.ng p roblem. Even if yo u didn't, if you have norrnal bin-
ocular vision, you are solving the correspondence problem all the time. The
cor resp onde nce proble m is the problem of figuring out whid1 bit of the image
in the le ft eye s ho uld be matched with which bit in the right eye. Figures 6 .37
FIG u RE 6.37 Is this a simple pic - and 6 .38 use an extremely sirnple sih1ation to il lus trate w hy corresp onde n ce
ture or a complicat9d computational is so tricky. The re are, of course, just three dots in Figure 6.37. Figure 6.38a
problem? traces the paths of the rays of light from the printed circles on the- p age to the

(a) The actual situation (b) 11/h at the visual system knowi> (c) Another plausible i.nte rpret.1tion
0 0
FIG URE 6.38 Interp reting the v isual lnforrnah:m from the three c ircles In Figure
6.37. It would rgquire careful placgm'9nt , but four dots in the world could producg
three dots in each gye as in (c).
irn..1ges on the viewer's reti n'-'S. TI1e re tinal images of the circles are labeled to
ma ke it dear w hid' image on the left retina corresponds to which im age on the
right retina, but your visua l sys te m has no such labels. All it knows about is
the retinal images, as sh ow n in Fig ure 6.3Sb. Fig ure 6.3&: shows an other pos-
si ble geome tric interpreta tion of the situation: if the left retinal im age o f circle
2 is ma tched to the r ight reti nal image of circle 1, and the left retinal imrige of
cirde3 is m a tch ed to the right re tinal image of circ le 21 you will perceive/our
circles, with the inner pair o f cirdes pen...--eived as fl oating in front of the oute r
pllir. (In fact, you may be abl e to exper ience this fo r yourself if yo u rnn cross
your eyes correctly.)
Wi th only three e le men ts in the visual scene 1 it isn' t hard to im agin e how
the vis ual syste m might adli eve the proper co rresp o ndence: firs t m atch the
two ci rcles w hose images fall on the foveas; then ma tch the two images to the
left of the fove..'l.s vdth ead1 o the r; the n m atch the two images to th e right of
the foveas. Befo re random dot stereogram s, a s imilar logic seemed reasona ble
for more complex scenes too. Go back to the face in Fi gure 6.32. O ur visual
systems could solve the correspondence p roblem by fi rst find in g the p a rts of
the hvo faces and then ma tching n ose to n ose, mouth to rnouth, and so fo rth.
The RDS in Figure h wevi;µ:. thousa nds of identical black and
d ots falling on eat'h retina. How h n l-\-"e be sure that the dot in the center
of the fovea o f on e eye corresponds to the dot in the cen ter of the other eye?
Even if we knew tha t, could we really match each d ot in the right eye \.\'1 th
ju st one d ot in the le ft eye? If there were a little dirt on the page, would the
w hole process colla pse? How in the world does ou r visual system succeed in
ma king the p roper m atches?
Ma tching thousa nds of left-eye dots to thousands of right-eye dots in Figure
6.33 would require a lot of work for a ny computa tiona l system . . H m·vever, the
problem is simpler if we look a t a b lurred version of the s te reogram . Blurring
lea ves onl y the low-s pa ti al-freque ncy info rma tion . Figure 6.39 sh ows the
low spa tia l frequen cies of theste reogra m from Figu re 6.33. Now, ra th er than
thou s.ands of d ots, we have just a fevv la rge blobs. Now you could ima gine a
process tha t, for example, ma tched the black blob in the upper left corner of
the left im age with th e very similar bloh in the right im age. C rude m atches

184 CHAPTER 6
FIGURE 6.39 A low-sp citiaJ-fr9qlJen-

cy-1itterOO version o f the stiereograrn in
Figure 6.33.
8230336 Amma
of this sort could act as anch ors, allowing the visu::tl system to fill in the fine r
(high-spatial-frequency) ma tches from the re.
In additi on to s tarting w ith low-spl\tia l-frequency inform a tio n , Dav id
Marr a nd Poggio (1 979) s ugges ted h vo mo re he uris ti cs for solving
the correspondence p roblem . They called. these the uniqueness and conti.Jiuity
constraints. The uniqueness constraint acknowled ges that il feature Ul the
world is re presented exactly on ce in eac h retinal image. \Vorking in the op-
p osite d irection, the vis ual system knows tha t each m onocula r image feature
(e.g., a n ose or a d ot) s ho uld be paired \.vith e xactly one fea ture in the othe r
monocular image. Notice that Figure 6.38c would n ot violate unique ness. Eadl
d ot in the world would be rep resented exactly once in each retinal image. The
odd thing is that two d o ts in the rea l world could be rep resented by the sam e
d ot in the retinal im age. The continuity constraint holds that,. except a t the
ed ges of objects, neighb or ing p ollltS. in the world lie a t s imilar dista nces from
the viewe r. Acco rding ly, dispari ty s hould d rn nge s moothly at most places in
the image. (These cons traints are difficult to illustrate o n a s ta tic pnge, but Web
Activity 6 .4: Stereoscopic Correspondence p rovides dy na mic explanations.)
With those con strain ts, the correspond enc-e proble m is not entirely solved but
it is m.ad e mu ch m ore tractable. The re are not so many p ossible solutions.
The Physiological Basis of Stereopsis and Depth Perception

Now th at we know something about the theoretical basis of s tereop si.s, we
can ask h ow it is implemented by the human brain. The most fundam ental
requirement is that input from the two eyes must con verge onto the same cell.
As no ted in C ha pter 3,. this conve rgence d oes no t happen lmtil the primary
visual cortex, w here most ne urons ca n be influenced by input from bo th the left
and rig ht eyes--i.e., they are binocular {Hubel a nd Wiese l, 1962). A bin ocula r
uriqueness constraint In stere- n euron has h vo receptive fie lds, on e U1 each eye. In binocular primary vis ual
cpsls, the cbservatlon that a feature
In the wcrld Is represented exactly
cortex ne urons, the receptive fie lds in the h vo eyes a re genera lly very similar,
cnce in each retinal Image. This con- s haring nearly ide ntica l orie ntati on and s pa tial- freque ncy tun in g, as '"''e ll as
stralr"lt simplifies the correspondence the sam e p referred speed and direction of motion (Hubel a nd Wiesel, 1973).
problem Tirns,. these cells are we ll s uited to the task of m a tching images in the hvoeyes.
conlirulty constraint In sternopsls. Man y binocular n eu ron s resp o nd bes t w h en the retinal images ar e on
the obsecvatlon that, except at the corresp onding points in the rn.·o retinas, the reby providing a neural basis for
edges of objects, relghborlrig points In the h oropter. H O\vever, many o the r binocula r neurons resp ond bes t vvh en
the world Ila at similar distances from s imilar images occupy slightlydiffr!1"ent positions on the retinas of the h vo eyes
the viewer. This is one o f several ccn-
stralnts that have been proposed as {Ba.rim'\', Blake more, and Pettigrew, 1967; Pe ttigrevv, Nikarn, and Bishop, 1968).
helpful In soMng the correspcnd€f1ce Jn other words, these ne urons are tuned to s pa rticular binocula r d is parity
problem. (diagramed in Figure 6 .40).

(a) FIGURE 6.40 In th9S9 simplified dia-

grW'Yls o f receptiV9 fields fcr two binoc-
ular-disparity-tuned neurons in primary
visual cortQX. the red miuron ..
stimuli falling on the red receptive
and the blu9 neuron respon ds to stimuli
falling on the receptive (these
fields overlap en th9 right
retina). ip) The ov"1fall picturQ, sho\'\ring
the fixation point in relation to the tvYQ
retinas. (/:J) The red neuron responds
best to a stimulus c los-9r to and slightly
to the right of the fixation poi nt. (.c) The
blu9 nsuron responds best if its pre-
ferred stimulus is behind and slightly to
th0 l9ft of fixation.
Recall the d istinction, fr om earlier in this ch ap ter, between me trical and

nonmetrical d epth cue5. Stereops.is can be used both metrically and n o1m1e tri-
ca1ly. Nonmetrical stereop sis mig ht just telJ you tha t a feahue Ues in front of
or behind the plane of fixation. Gian Poggio and his colleagues (Poggio and
Talbot, 1981) fo und d isparity-tune d neurons of this sort in V2 (which stands
for"visual m ea two" ) and some higher cortical areas. Some neurons responde d
posi ti vely to disparities near zero-tha t is, to images falling on correspo nding
retinal points. O ther ne m o ns were broadly tuned to a range of crossed (n ear)
or uncrossed (fu r) disparities. On the other hand, stereopsis a m a lso be used
in a very precise, rnetrical m ann er. Indeed, stereopsis is a "hyperacuity'' like
Vernier acui ty (see Web Essay 3 .1: Hyperaculty) \vi th thresholds smaller
than the size of a cone. Both of these fom1s of s te reop sis have their uses, and
functi onal magnetic resonance irnaging (fMRI ) d ata s uggest that the dorsal 0336 Amma Appa
u·here p athway (see Ch apter 4) is most interested in metrica l s tereopsis, while
the ventra l what pathway makes d o with more categoric.:i l, near-versu s- far
information (Pres ton e t a L, 2008).
The neural bases of o the r depth cu es have a lso been investigated. For
exa mple, w hen we d iscussed motion parallax ear lier, we su gges ted m ovi ng
your head back and fo rth w hile looking into the branches of a tree in orde r to
create a m ore vivid impression of the depth relation.ships a m on g the branches
and tw igs. To exp loit th a t c ue properly, you need to know how your head is
moving (see Chapter 12) and how items in the visual field are moving (see
Chapter 8). Nadler, Angelaki, and DeAngelis (2000) looked for the neural
s ubstrate of parallax in the middle te mporal a rea (area MT) of the brain of
1nacaque monkeys. As 1'>/e will see in C hapter 8, this area is very important in
the perception of motion. Nadler e t a l. set up an apparatus ""· here the monkey

186 CHAPTER 6
was rnoved from s ide to si de whi le items on the screen also moved . lf the
monkey was integrating s ignals about its head rnovement with the motion
signalsr then the objt:ds on the screen should have been seen in depth. Other-
wise, they \·vould have been seen as just rnoving in the plane of the screen. It
turns out that cells in area MT can signal the sign of depth (near 0 1· far) based
on this m otion para llax s ign.al alone.
Other visual also contribute to the complex business of inferring the
three-dimens iona l world from h vo-dimensiona l retina l images. Anzai and
DeAngelis (2010) suggest that e.:i.rly visual areas1 par ticularly V2, are involved
in computing d epth order (wh o's in front?), based on the contour completion
and bo rder m\.,1ers hip p rocess we di scussed in Chapter 4 (see Figmes 4.5
and 4.13). lnterrnediate visual areas s uch as V4 ( meaning "visual area fo ur")
encode depth intervals, based on relative disparities, and higher cortical areas
s uch as 1nferotemporal cortex are involved in the representation of complex
three-dimensional sh..1pes.
FURTHER DISCUSSION of phase as it reates to vision can be found in

Oiapter 3 on pages Sensitivity to i:hase is also important fa tempo-
ral ooding of soond frequency (Chopter Q, page 278).
Co mbini ng Depth Cues

lf the chapters of th is book v.:.-ere novels, this d 1apter cou ld be said to have
8230336 Amr t:l\e ;Jtame p lo t as the discussion of object i·ecognition in C hap ter 4, but w ith
different characters. ln Chapter 4 we ta lked about a se t o f cues that en able us
to group locaJ features toge the r into p ossib le objects and then to recognize
those objects. We d escribed the process as a sort o f committee effort in wh ich
different sources of informa tion a Ucontribute their opinions and w here we see
the ccmunittee decision without necessari ly knowing how that decision was
read1ed . ln this chapter we've covered multiple sources of depth infomtation
and they. too, need to be combined . Any or all of these cues might be avail-
able to the visual system w hen we' re viewing a ny visual scene. None of the
cues are foolproof, and n one work in every possible si tuation . For example,
relative height produces inconsistent or misleading information if we
see the point a t w hich an object to uch es the ground . All we really have is a
collection of guesses about p ossible d epth rela ti ons behveen different objects
in our visu..1.I field.
By carefuUy combining and \\•eighting these guesses, the visual system
generally arrives at a coheren t, a nd more or less accurnte, representation of
three-dim ensional space. Helmho ltz, \vriting in the nineteenth century {an d
trans lated into English in the t\11,rentleth), called this automatic cu e combination
process uunconsdous infe ren re" (Helm.holtz, 1924). ln recent years, a number
of vision researche rs have been a ttempting to put this sort of argument on
Bayesian approach A W'i# of for- the more rigorous m a thematical footing of the Bayesian approach that we
malizing the Idea that our perception Is
mentioned in C hapter 4 and that is the s ubject of a m ore de tailed essay on the
a cx:mbinatlon of the current stlrnulus
and our knowledge about the condi- website, Web Essay 4A: Bayesian Analysis.
tions or
the wond-what Is and Is not
likay to occur. The Bayesian approach The Bayesian Approach Revisited
Is stated mathematically as Bayes' Reca ll that the basic insigh t of Reverend Thomas Bayes '"'·as th at prior know l-
=PIA) x P(OIAV ed ge could influence estimates of the p robability of a current observation. Let's
P(O)-whlch enables us to calculate
the probability IP! that tl1e v">rld Is In this idea to a concrete, depth perception example. Suppose our visua l
a particular state IA) given a particular systern is confronted '-vith the retinal image shown in Figure 6.41 . There are
observation (0). infinite possible ways to prod uce this retina l image. Actually, thi s is a bit o f ..i

problem for the use of Bayes' theorem. We do n' t n.."Ylfly know the prior prob-
abilities (M. Jones and Love, 2011). Still, we can acknowledge that limitation,
and we can s till make good u se of the' basic ins ight that som e hy potheses are
m ore likely than others and that the;e prior probabilities can shape o ur in-
terpretation of the world. Tiuee hypotheses nbout o ur pennies are sh own in
Figure 6 .42. f\faybe the h.\•o pennies are the same size, but the on e on the left
is slightly farther away than the one on the right (Figure 6.42" ). Maybe the
penny on the right is mud' s malle r, but also m\.1ch closer, than the penny on
the left (Figure 6.42b). the two pennies are equidistant, but the penny
on the left is smaller and has had a bite taken out of it (Figure 6.42c). If you
FIGURE 6. 41 Retinal Image of a
d on' t see h ow the set o f p ossibilities could be infinite, reme1nber that s ize slmpe v isual scene.
and distan ce can vary continuously over a large range. ln Figure 6A2b, the
big penny cou ld be on the rnoon (but it would h.ave to be a really big pe1my).
How d oes the visu.:1.l syste1n decide what we' re actua lly seeing? Which
interpre tatio n seems 1nos t likely? That is the core of the Bayesia n approach
(except thatit'sall automatic; our conscious selves do not get to make th e deci-
sion). [n mu: exp e rience, all pennies are the sam e size. This cue of fami liar size
is o ne source of prior know le dge in this case. This makes the prior probability of
the hyp otheses shown in Pig ure 6.42.n higher than the prio r probabilities of the
other hvo hypo theses. Furthermore, for the scene in Figure 6.42r to produce
the re tinal image in Figure 6A1 , we wou ld have to be seein g the scene from
o ne of those unusual and unlikely 11aocid entc1l viewpoints." It is mud1 m ore
likely that the points of contact between the images of the h.-vo p e nnies reflect
occlusion. If we were to plug all these probabilities into the math of Bayes'
equa ti on, \ Ve would find th.:1t,given the image in Figure 6.41, the m ost like ly
answer is the scene d ep icted in Figure 6A2n .
(•) (b) (r)
1
Identic.tl pennies PeruUes of A penny with a bite
,1t .slightly different di:ffarent siz.es at out of it, right next
d'Ptru very different depths to ai nonnal penny
uuu
FIG URE 6.42 Three of the infinite number of scenes that could generate the retinal
imag9 in Flgur9 6.4 1. Which o f thooe ts th£ii most probro!Q?

188 CHAPTER 6
!\!\!\!\!\
FIG URE 6.43 In which image are the two horizontal lines the same length?
ln thinking about combining d epth cu es, our choice of the me taphor of a

committee is n ot arbitrary. We could have talke d about an election, but that
would have itnplied something like "one cue one vote." On a co mmittee,
you might have one member w ho is s tronger than the othe rs and vdn.s all
the arguments. You might give more '"'eight to the committee member \"tho
comes prepared with the best in.formation. The committee nUght defer to one
me mber on one top ic and another mernber on a different topic. Somethlng
like this option is d escribed by He ld, Cooper, and Banks (2012). Binocular-
infom1atio n can be very precise, but that is only true ne ar the plane
of fixation (re.m e mber Panum 's fusional area?) . Blur can be quite a good cue
too, but it is actually better aw.J.y from the plane of fi xa tion . When Held e t
al made s timuli th.a t had dis pa rity cues, blur cues, o r both, they found th.J.t
disparity drove respon ses where di sparity was more reliable and that blur
drove responses in parts of the three-dim ensionaJ world where it was more
reliable. Th.is is different frorn just letting every cu e have its say, and it makes
us realize that the visual system mus t be estimating h ow reliable each depth
cue might be .
Illusions and the Construction of Space

If oul" visual perception o f the '"'orld is our best guess about the ca uses of
visual input, then interesting things should happ en when a g uess is w rong.
In some sense, as vdth the pennies jus t discussed, a guess is wro ng w hen-
ever we look at a two-dimensiona l picture and see it as three-dim ensional
As noted, however, we are not really fooled into thinking that the picture is
three-dimens ional. Lt would be m ore accurate to say that '""'e make a p lausible
guess about the three-dimensional world that is being represented in the two-
dimensional picture.
\.Vhat about a situation like tha t shown in Figure 6 .43? One of the five pairs
of horizontal lines (and on ly one) shows two lines of the sam e length. Can you
p ick the correct pa ir (vd thout a ruler)? ln fact, it is the second from the left _
Odds are you picked the third or fourth pair1 even though the bottom line in
both of those images is physically longe r than the top line. This is known as
the Ponzo illusion, naine d after Mario Ponzo, who described the e ffect in 1913.
What causes this illusion? For many years, a p opular famil y of theories has held
that the illus ion is a guess gone wrong-a situation in which '"·e overinterp ret
the depth cues in a hvo-dim ensional image. The basic idea is illustrated in
Figure 6.44. Maybe the two tilte d lines that induce the illusio n in each image
of Figure 6A3 are being interpreted b y the visual sys tem as linear-p erspective
cues like the train tracks in Figure 6.44. lf so, then objects that were the sam e
s ize in the two-dimensional image \o\-'ould represent objects of diffe rent sizes
in the three-clime n.sional world .
8230336 /. SUch accounts are very cornpelling and exist for a wide range of visual
illusions (Gillam, 1980). (See Web Essay 6.3: The Moon Illusion.) They are
consistent with the view that the job of the visual system is to use available
cues to make an intelligent guess about tl'e world (Gregory, 1966, 1970). jus t

FIG URE 6.44 Th9 t...vo p 90ple t>Jjng across th9S9 train
tracks ar9 the same size in the irnage. You c an vt1orlfy
this by rnaasurk1g th.em. the more distant
pgrson INOUld n94;1d to b9 muc h larg'-C" in the rM threa-
dlmensional v-rorld to produce thB Image in the two-
figure.
becau se a n ansvo.·er is plaus ible, however, doesn' t m ean it's right. ln Figure
6A5a, line B looks longe r than line A That m akes sense if we' re interp reting
these lines as lines lying '1t d ifferent distances on the w all of the colo1made. As
in the Ponzo illusion, if line Bis farther away than A, then the same i.rnagesize
implies a larger size in the real war.Id. But what '1 bout lines C a nd D? Surely
D would be inte rpreted as farther away than C. Does it look convincing ly
larger? Ftgure 6.45b reveals \.\.'hat you probably guessed-that alJ fi ve of the
lines in Figure 6A5a are the sa me leng th.
Prin zmetal, Shimamura, and Mikolins ki (2001) use a demon::;tration like
this as pa rt o f the ir argument that the Ponzo illusion is not reall y a by-product
of de pth cues. They argued that it reflects a m ore general asp ect of the visual
system 's response to tilted Hnes and is related to iJlusions like the Zollner and
Hering Wus ions illustrated in Figure 6.46 {which we w i!J '101 try to exp lain;
(a) (b) FIG URE 6.45 All o f the red lines in

this illustration (e) are the same length,
as you can Sie9 In f/;JJ. Does e look big-
than A? Does D look bigger than
C? In 98.ch case, which ling is farthi91"
aw<:tf if (a) sho\'llS a colcnnade?
A I I
B E
I
0
. mm•App•
1
c

190 CHAPTER 6
FIGURE 6.46 Despite their appear- (a) The: Zollner illusion (b) The He-ring illusion
ance, the vertical llnoo are paraJlel In (a),
as arei thie horizontal linQS in fP}.
see Prin zmeta l and Beck, 2001). The p oint is d ebatable. After a ll, line E looks
very big dmvn thereat the apparen t end of the colonnade in Figure 6.45a. \Vho
is right? Jt could be tha t both positions ho ld a piece of the truth. Perhaps the
visual system's response to tilted lines is related to the role of those lines m
creating an impression of depth. Going back to Figure 6.44, that would mean
that the Ponzo illusion '\.Vas n o t based o n some version of the railroa d track
s tory but that the processes that give rise to a three-dimensional interpretation
of Figure 6.44 also give rise to illusions like those in Figures 6.43, 6.45, a nd 6.46.
Binocular Rivalry and Suppression

The p recedin g sections demonstrated that objects in the wor ld o ften project
Un.ages on o ur two retinas that d o notoverk:ip (that is, the images fall on n o n-
corresponding retina l points) and tha t the visu a l system is physiologica lly
prepared to deal with these discrepancies via disparity-tuned neurons in s tri-
ate cortex and beyond. But wh at happens w hen completely differen t stimuli
are presented to the two eyes? You can answer this question for yo urself by
fixating on a s mall object across the room, su ch as a clock, and moving your
hand up so tha t your fingers occl ude the object in the right eye (makin g sure
the left eye still has an unobstructed view}. It would seem a bad id ea to fose
the fingers and the clock into a single perception of something tha t d oes not
exist in the world. Accordingly, tJ1e visual system drnoses instead to suppre.55
one image and peri,:eive th e other. Jn the present situation, you probably see
the clock as though you ,..,1ere lookin g throu gh a hole in your ha nd. (Fo r an
even m ore com pelling perception of a h ole in your lu1.n d, dick on "An other
Dernonstration'' in Web Activity 6.2: Binocular Disparity.)
How d oes the visual system "decide" wh C!. t to see? The more interesting
of the t\.vo s timuli is likely to be d ominant. Interesting in thi s case has several
mean ings. 1l1e mos t important factor is \'\1 hich stimulus more salient to th e
early stages of cortica l visual processing. High contrast is m ore salient than
low contrast1 bright is better than dim, moving objects are more interesting
tha n s tationary ones, and so forth (Fahie, 1982). TI1e meaning of the.stim ulus
also h as an effect (Yu and Blake, 1992lr as does wha t you ' re a ttending to (Ooi
and He, 1999).
The compe tition between the two eyes for control of visua l perception,
!mown as binocular rivalry, is never completely won by ei ther eye (Alais and
Bla ke, 2005; Wheatstone, 1852) o r eitl1er s timulus (Blake and l<>gothetis, 2002).
lf you stare at the combination of the dock and hand long en ough, your fingers
\viii eventually conquer the vis ua l territory, only to surrend er it back to the
binocular rlvalry The ccmpetltlon clock a mo ment bter. The battle is easier to see if the two combatants are more
between the two ayes for contrc> of
visual rerceptlon, which Is e\ldent closely matched . If you free-fuse the two panels of Figure 6.47, yo m visua l
when completely different stlmull are systern wi ll not actually combine the perpendku lar stripes in the two center
presented to the two eyes. squares. Jn.stead, you will see a battle behveen the vertica lly and horizontally

I
-·I "I I· "" .1
""'""'
I
I
FIGURE 6.47 Binocular rivalry. If yo.J fre&fuse these two you will be able
to watch the blu9 w rticals and orange horizontals engage in the pet"CQptual battle
known as blnocular rtvalry.
striped patches, with regions of dominance g rowing and shrinking time,

as illustrated in Figure 6.48.
Binocular riva lry might seem an odd situation tha t would arise only in a
vision lab or a perception course, but a m om ent's reflection should con vince
you th.a t the stimuli for rivalry are actually very comm on. Recall that Panum 's
fusional area is the region of s pace surrounding the horopter w ithin which Left eye
=J
images on the two retinas can be fused . As n oted earlier, objects located too
far off the horopter--outside Pan um' s area-are s ubject to d.iplopia. ln fact,
the proportion of the three-dimensional visual world that fall s inside Pan urn' s
area is actually fairly small. Think about tw"o pairs of corresponding retinal
points: AL and AR, and BL and (the Land R s tand for "left" and "right,"
respectively). Now imagine tha t one object falls on two noncorresponding
retinal points: AL It follows that somethhigelse must be forming fill image
on AR. There will be noncorresponding images on the corresponding p oints
AL and AR (and, similarly, on Be and BR). Those are the conditions for rivalry.
The fact that the IB•o eyes are seeing different images brings us to an an-
cient problem: the problem of binocular sing le ";sion. 'Why; when we see one
elephant in one eye and one e lephant in the other eye, don' t we perceive h.vo
=
elephants? Given what we've learned in this chapter, ""-e' re now in a position
to a nswer this question. If the elephant is \vi.thin Panum's area, we fuse its
two images into a single s tereoscopic perception. If it is outside Panum's area1
we normally s uppress one o f the copies. Why do n't we see the rivalry? In
part, because we aren't looking. Our a ttention and eyes are typically directed
toward the foveated object or toward objects falling on roughly corresponding
points in each eye. Moreover, acuity is so bad in the periphery of the visual
fields tha.t, even w hen objects are vying for binocular precedence, the rivalry
is quite indistinct.
The classic demonstrations of rivalry pit a s timulus in one eye against a
stimulus in the other eye. More it has become dear that r ivalry is
part of a larger effort by the visual system to come up with the most likely
version of the wodd, given the ourrenl'!"etihalimages (solUlds Bayesian again,
FIGURE 6.48 If blu9 w rtlcal bars shOIM1 to on9 &y& whla orange horizontal
bars are shoWl to th& other. the t'NO stlrrull will battle for dominance.

192 CHAPTER 6
d oesn' t it?) (Clifford, 2009). lf you c,m free-h ise F1gure6.49a,

you will see the monkey and the text battle each o the r. That
is jus t s ta ndard binocular rivalry. Interes ting ly, you will see
a similar m onkey / text riva lry if you free-fuse Figure 6.49b.
ln that case th e monkey is bei ng put together frotn bits in
th e h vo eyes. Your brai n is not trying to p ick a wi nner in a
ba ttle between the eyes. It is trying to fig ure out the world,
and if that interes ting monkey is seen w ith one eye in one
spot and the other eye in another spot, the brain can put those
different bits together into a co herent percep tion (Blake and
\Vilson, 2011 ).
Ri valry is a very useful tool for probing one o f the more
vexing problems in the neuroscien ce of perception: what parts
of the vis ual syste rn give rise to the conscious e.xp erien ce of
seein g somethin g? For these purposes, the g rea t fea ture o f
rivalry is th<'lt it di ssociates the s timulus on the retina fro m
the s timulus tha t you see. Now, imagine tha t you record
from sing le cells som e'",.here in the m onkey vis ual system
(Logothe tis ond Sch all , 1989) o r you use fMRl to look at
the working human b nlin. If you train the rn onkeys or ask
lnuna n.s to m onito r their p erception (''Do you see vertical
or hor izonta l?''), you ci.ln as k whether the neiual signal in
a sp ed.fie p a rt o f th e vi s ual pathw r.y foUows the physical
FIG URE 6.49 Binocular ri valry is no t s timulus or the perceived stimulus. ft is a complex and evolv-
just a light between the two gyes. lf you ing story, but one tha t clearly s hows that conscious vis ual is not
c an fre e fu se these pairs o f images,
4
some thing that hap pens in one d iscrete s tep in a chai n of visua l processing
you w ill S9Q a fyplcal example of rtvalry.
(a) Dlff€<e<rt lmagg.s In each eye s truggle (Tong, Men g, a nd Blake, 2006).
for d ominarlCQ. Ths. chimp and the text
struggle for 00min3flc e In panel (b) too.
but this Is pattern rivalry: the two eyes Development of Binocu lar Vision and Stereopsis
are actl1alty coop erating to put to gether
coheHmt \'isvvs of the c himp or the t ext.
What is binocular "ision like in infants? Babies are bo m w ith two eyes, but a re
(Fro m Kovacs et al ., 1006.) they born with s tereo psis? If n ot, how d oes binocul ar function deve lop? In a
wonderfu l con versation a bo ut visual development, Davida Telle r a nd Tony
Movshon (l 9S6) recalled a lecture by a disillusioned deve lopmental psych olo-
gist, John McKee, who argued tha t the field could be s ummed up by three la\'\--s:
1. As children get o ld er, they get better a t tl1ings.
2. \ Vhatever it is, gi rls d o it before boys.
3. a lon with ever hing else.
To these "laws,"Tetler added a s umm statement: "Things s tart o ut
then they get bette r; then, after a lon g time, they get worse again" (Teller and
Movshon, 1986).
As it tllrns o ut, research over the last 25 years or so has s hown th at visual
development provides su pport for the fi rst and second "ktws," bu t n ot fo r a
s trict for m of the third. The development of binocula r vision and s tereop sis
provides o ne of the s trongest violations of that third law.
Most vis ua l functions indeed s ta rt o ff badl y (but not as badly as w e used
to think) and then imp rove steadily w1til they read\ adu.ltlevelsi however, the
d evelopment o f stereopsis is s urprising.. in that infants are essentially blind to
d isparity until about 4 months of age. At that point, s tereopsis a ppears qui te
s uddenly-almost out of the blue. Of rneasuringstereopsis (or any thin g
else) in infants is no easy taskr but de velopmental psychologis ts have been
very inventive in designing methods for assessing deve loprnent.

FIG URE 6..50 The ons9t of stEM"90psls. This tigure shO\<\IS the P9fC'1rtage of D Study l: Llnestereogr.uns
Infants dem onstrating stereopsis 1or the first time as a function of their age. In thr"*1 0 Study 2: Dynam ic RDS
separate studies (the three different colors), almost al l infants showed stereopsis D Study 3: Dynamic RDS
for th9 fir st time bet\rv9en 3 and 5 months. (Mer Birc h. 1QQ3.)
Despite di_fferences in technique.sa nd procedu res, m ost investiga tors

agree abo ut the onset o f s tereopsis. Fjgure 6.50 s ummarizes the results of
several s tudies, showing the age a t wh ich stereop sis can first be de tected .
These s tudies (a nd others like them) looked for ed dence indicating that
infants could reliab ly d etect a large binocular disparity (typicalJy on the
order of 30--60 arc minutes). The agreem ent among the s ti.1dies is remark-
able. l.nfants are essentially s tereoblind before 3 months, with m ost infants
shovli.ng a su dden onset of s tereopsis between 3 and 5 m onths.
Stereopsis is not an all-or-none phenomenon. Jus t as an individual's acuity
is a measure o f his ability to resolve sp L'l tia l detail, stereoaculty is a measure
of the smallest binocular disparity th at can gen era te a sensation of d epth. Age when stereopsis was first
Once an infan t d evelops s tereopsis, stereoocuity increases rnpidly to near detected (mo nths)
adult le,·els (Figure 6.51 ). Birch a nd Petrig (1996) fo und that s tereo.,cuity
rose from essentially nothing before 4 months to near adult levels by 6 m onthst
Th.is time course is very differen t from th e time course in the development o f
simple acuity. TI1ough coarsely p resent a t birth, basic acuity takes years to rerich
adult levels. The same differen ce between basic acuity and stereoacuity is seen
inmonkeys(O'Dell and Bootl1e, 1997), butthe qver Urat of eve! R'Jentis
faster. Interestingly, not onl y stereoo.cu ity but several o ther vtsual functfOns
d evelop at a rate approximately fo ur times fas ter than in humans 1 as if one
m onkey week \Vere the equivalent of one human month. In keeping with th is
rule of one human month being equal to one monkey week, s tereops is can stereoaculty A measure of 1110
be detected in ITton keys "";th in the (i.rst 3-5 weeks of life, compared wi th the smallest binocular dlspartty that can
3- to 5-month wi ndow of onse t observed in humans. generate a sensation of depth.
Adults
0:1
F IGURE 6.51 Th9 d9wlopm 9nt o f ste.reoacuity. Ste-

rno.acuity develops to adu lt 19vels w ithin the first 6 - 7
months of llfEi, Data points bebw the dashed line lndicata
Age.(months) unmeasurable sterooaculty. (After Birc h and P(lttig, 1096.)

194 CHAPTER 6
dichoptlc Referring to the presen- H ow, then, d o "',.e explain the sudden e mergence of s tereop sis in humans
tation of two different stimuli, one to at about 4 months? Although a newborn infan t makes convergence eye move-
each eye. Different from blncx;ular ments to track a target as it approaches he r n ose, accurate and consiste nt
presentation, which could Involve toth
eyes looklrg at a elngle stimulus. convergence probably d oes n ot occur unti l 3-4 rnonths of age. But \Ve can ' t
concl ude tha t inaccurate convergence prevents stereopsis from developin g
earlier than 4 mo nths, because convergence need not be very accura te in order
to detect large disparities. Moreover, severa l s tudies used repeating gratings,
which would be hised at some disparity even if con vergence were inaccurate.
Before 4 months, babies d on' t respond to these s tereoscopic sti muli either.
A.n alternative view is thnt the failure of s tereopsis to develop prior to 4
months might m ean tha t some part of the visual system is immah1re. Disparity-
sensitive n eurons in the primary visua l cortex (Vl ) are one plausible candidate
for that inunature part, but recent a niltonUcal and physiological data suggest
th at we need to look beyond Vl for an explanation of w hy infants d o not
exhibit s tereopsis.
Yuzo C hino and his colleagu es made the most d eta iled quantitative study
of the binocular responses of Vl neurons o f infant monkeys (Chino e t al.,
1997). TI1ey presented a pair o f drifting s ine \\·ave gratin gs dichoptlcally {one
to each eye). The sine waves were identical in spatial freq uency, orientation,
con.trnst, and velocity, and th ese values were ch osen to maximize the cell's
response. A phase difference between the two monocular gratings wil l also
create binocular disparity. Some cells in the visua l cortex are sensitive to this
phase s hift a nd Freeman, 1986b). Whe n Chino et al.
the reliltive spatial phase of the two drifting sine waves (Figure 6.52a), the
response of a binocular neuron waxed a nd w(1ned (Rgure 6.52b). The das hed
lines in FigLUe 6.52b s how the levels of response for the two eyes, stim ul ated
alone ot:Gxt that fo r somejQ terocular phase differences, the binocular re-
sponse was considerably higher than the response th.rough ei ther eye alone.
A t other interocular phase differences, the binocular response \\laS lower tha n
the response through either eye alone. lhis sinusoidal bincx::ular phase tuning
is the hallmark of this type o f binocular neuron in the visual cortex. Using
tlUs sensitive method, Chino e t r1l. (1997) found thri t within the first \\>·eek of
life-well before the onset of stereopsis-infan t monkeys h ad practica lly the
same prop o rtio n of phase disparity-sensitive neurons th a t a dul ts have in
primary visual cortex.
In addition, the ocular dominance properties of these infant monkeys were
essentially iden tical to those of adults. Other investigators have also found th.a t
oc ular dominance colunms in the input layers of Vl a re essentia lly adultlike
a t birth (Horton and H ocking, 1996).
FURTHER DISCUSSION of ocular dominance can be found in Chapter 3

en page 72.
\Vha t d o these s tudies tell us about the d evelopmen t of s tereopsis? TI1e

resu lts su ggest th at the neural app aratus in Vl of nev1i•borns is capable of
combining signals from the h\'O eyes and that it is sensitive to interocular
disparities. So wh y are newborns blind to disparity? One possibility is that the
ex traction of rela tive dispa rit); w hich is nee ded for s tereoocuity, takes p lace
beyond Vl , possibly in V2. At this time we d o n ot know much about h ow V2
neurons in newborns resp ond to disparity, but emerging evi dence s uggests
that other receptive-field properties mature later in V2 tha n in Vl {Zhang e t
al., 2005; Zheng et al ., 2007).
Another possibility is that the problem is in Vl. Al th ough Vl ceUs of
newborn monkeys are adultlike iJ' their response to interocular phase dispar-

(•) FIGURE 6.52 Interocular phase dif·

ferenoe gratings are used to study dis·
parity tuning. l hie sinusoidal grating
drifts difffll'"lfltly in one eye than in th'11
othi9t' . producing a si nusoidal c hange
in spatial phas9 disparity. lh9
reisponse o f a binoet.ilar neuron, Notice
that the binocular respcnse varies with
phasa, and the m onorular responses
do not. Olino et al .. 1997 .)
(b)
Binocular 1-eis:ponse - -
Relative Sf'lliaJ phase
ity, these neu rons rem ain immature in several import.1.nt ways. They do n o t
have adult sensitivity to mon ocular spatial frequency or direction of m otion .
Moreover, they are mud-1 less resp onsive overall than are adult neurons (thr1t
is, their peak firing rates are consid erably lower}. In addition, these neurons
d isplay m ore interocula l' s uppression th an adult neurons do. Thusr it is also
possible that because of the inunahtrity in Vl ne urons, the signal s they send
to the n ex t stage of processing are too weak or confused to suppor t stereopsis.
Abnormal Visual Experience Can Disrupt Binocular Vision

The p resence of a ll this binocular ha rd ware, even if it is immature, strong ly
suggests that extensive binocular visua l experience is n o t necessary for
lar connections to fonn in Vl. l11ese connections are present at birth or very
shortly thereafter, so we don 1 t need to 11 lea rn '' or develop binocuJar vision.
However, the normal d evelopment of adult binocular vision and stereopsis crltlcal period A period of time dur-
d oes require visual experience. ln Chapter 3 we lear ned about Hube l and ing development when the organism Is
particularly susceptible to developmen-
Wiesel's work on the critical period, the period during early visual d evelop- tal change. There are c ntl°'" periods
ment when nomlal bin ocular visual stimulation is required for norma l cortical in the development o f binocular v1sloo,
d evelopm ent. During thisperiod the visual cortex is highly susceptible to a ny human language, and so en

196 CHAPTER 6
Fixation FIG URE 6..53 Left esotropia, Thie patl911t 1Nants to fixate en the brick, but tM
left 910 Is turned too far toward the nose: as a result, the left fovea Is pointing at a
ferer:it location f1ere th9 purpl9 pg-itagon). while th9 image of the ygHow brick falls to
the right of th9 left fovea
disorder that a lters norma l binocular visual experie nce. In cats and monkeys
this critical period is approximately th e first 3-4 m onths of life.
'We canno t s tudy a child's visual cortex the way we mig ht study a monkey's
or a cat's. How, the n, can '"'e es timate the critical period in humans? Some
humans are born wi th n...·o eyes tha t do not p o int at the same s pot in the
world . This no t unco nunon k.J,o,..,,..,, as strablsmus, as we learned
in C hapter 3. The mcidence JS about 3 o. In esotropia, one eye is pointed
too far tov.'ard the nose ("cross-eyed "). In exotropla, the d e\.;ating eye is
pointed too far to the side. 1l1ere '-lre various ways to lTeat strabi smus. For
example, it is p o.ssible to s urgic.Jll y correct the posi tion of the eyes. For the
present discussion, however, the important point ts that it is possible to test
ad ults who had misaligned eyes a t different tim esu nd for different durations
during childhood .
Reca ll from Ch.."lpter 3 tha t exposure to lines tilted to one side of vertical
'\vill make vertical lin es appear tilted to the othe r side. This is known as the tilt
aftereffect One ch amcteristic of the tilt aftereffect is that it shows interocular
tr.Jnsfer (transfer of the e ffect from one eye to the other). If we show the adapt-
ing lines to one eye, \'\'e can measure an aftereffect through the other eye. This
result is genera lly to show that the cells resp onsible for the effect are
binocular: they receive input from both eyes (for some de tails, see \Vo lfe and
He ld, 1981 ). Individuals w ho e xhibited s tTabismus during the firs t 18 months
of life d o n ot show normal interocular trans.fer (B..1.nks, •.\slin, and Letson , 1975i
H ohmann a nd Cre utzfeldt, 1975). This result provides an ind.irectestim..'l.te of
the period during w hich binoctJar connections in humans are susceptible to
input.
Let's explore in a bit more de tail w hystrabismus disrupts bin0ctJar vision.
\'Ve will u se left e.sotrop ia as an example. In left esotropia (Figure 6.53), the
le ft eye is turned in. As a conseque nce, although the object o f fixation (th e
yellow brick in this case) lands on the fovea of the right eye, in the left eye
it lands on a region in the " nasal" retina. (The nasal retina is the half of the
retina closer to the nose.) This means tha t the irnages of the yellow brick are in
n on corresponding po.in ts in the two eyes. \ Vhat will the patient see? If an adult
becomes esotropk (perhaps because of an injmy), sh e will experience diplopia
strct>ismus A misalignment of the (do uble vision ), seeing nvo bricks instead of one. However, people wh o exhibit
two fi'/es such that a sing le object In s trabismus ea rly in life often experience no such problem. Why ? No tice that
space Is Imaged on the fovea of one
&ye and on a nonfoveaJ area ofth9 something is present at the fovea of the left eye. In Figure 6.53, this some th ing
other fi'/0. is a purple pentagon. Thus, in s trabismu.s, nom1ally corresp onding points in
the hvo eyes receive conflicting informatio n (th is si h1atio n is knmvn-no t
esotropla Strablsmus In which one
eye deviates Inward. unreason.:1bly-as: "confus ion"). To eliminate diplopia and confusion, the brain
suppresses one of the hvoirnages. lt is simply n ot consciously perceived. ln
exotropla Strablsmus In which one
"'f0 deviates outward. esotropia, the most com.mon pattern is s uppression of the input from the eye
th at is h1rned in. So, in the examp le in Figme 6.53, the patient wo uld m ost
tilt anereffe<:t The perceptual lllu-
stn o f tilt, produced by adaptation to likely s uppress visual input from the le.ft eye.
a pattern o f a given orientation. Binocular rivalry is a form of s uppression, so some suppression is an
inlportant part of normal visua l experience. Unforhmately, ea rly-onset stra-
suppression In -Aslon. the Inhibi-
tion of an unwanted Image. Suppres- bi.smus can have oth er, more serious effects on the developing visual ne rvous
sbl occurs frequently In people with system , and on visual performance. For exampl e, strnbis mus grea tly reduces
strablsmus. the ntunber of binocular neurons in the visual cortex (\\'iesel, 1982). Cel ls that

SPACE PERCEPTI ON AND BINOCULAR VISION 197
FIG URE 6 .54 Dw9lopTient of st9r90p sis in normal infants (red line) and in esotro -
pes (blue). Beyond 4 m onths, very few esotrop k:: Infant s dernons trated st€11'eopsis.
(After stager and Birch , 1Q86.)
- Normal
- F.sotropic
would no nnally be driven by both eyes are d onU.na ted by only one. You would
be cor rect if you su spected that th.is s itua tion would disrupt s te reop sis. Birch
and he r colleagu es (e.g., Stager a nd Birch, 1986) followed the d evelopme nt of
stereopsis in norma l and esotropic in fan ts. Figure 6.54 illustra tes the percentage
of infan ts w ho s howed a meas urable ability to perceive stereoscopic depth.
The red line s ho""-s tha t by about 6 months of age, alm ost all norma l infants
d em onstra te s tereopsis. ln contrast, theesoh"opes (all of w ho m were diagn osed
w ith esotropia by 6 m.o nth.s of age} initially de mo nstl·a ted a n orma l pattern of
stereop sis developmen t. Afte r 4 months, however, very f ffi>v esotrop ic infants 10 12 14
ted stereop sis. Age (months)
This res ult has an in teresting pa rallel in corti cal physio logy. Chino a nd his
colleagu es (Kumagam i e t al , 2000) m.ade otl1erv..·ise nonna1 m.onkeys stra bism ic.
They fo und tha t a brief period o f experi me ntal strabi.sm us shortly after the age
of onset o f stereop sis produced a grea ter loss o f d ispa rity sen sitivity and more
b inocular s up p ression in Vl neurons than did an earlier e p isode of s trabis mus.
Th ese physiological and perceptual deficits a pp eil!' to be perman ent a nd have
importa nt im p lications fo r the s m gic-a l treatment of infantileesotropia. Alm os t
all s urgeons agree tha t treatmen t should be early, but there has been a lot of
d eba te about just how early. These resul ts su ggest tha t the treabnen t s ho uld
take place before the age a t w hich s tereopsis normally develop s, in ord er to
mi nimize the damage d one by esotropb.
Summary
1. Reconstructing a three-dimensional world from two non-Euclidean, curved,
t\vo-dimensiona l re tinal images is one basic problem fa ced by the brain. Refer to the
2 A nwnber of monOC'Ular cues provide informa tion abo ut three-dimensional Sensation and Perception
space. 1h:se include occl usion, various size and position cues, aerial per- Companion Website
spective, linear perspec tive, motion cues, accommcxlation, and convergen ce. Sltes.Slnauer.comtwoHe4e
3. Having hi.•o eyes is an advantage for a number of reasons, some of w hich for activmes, essays, sl.Jdy
have to d o ....·Hh depth perception. It is important to remember, however, quesl ons, and other study aids.
that it is possible to reconstruct the three-dimensional \Vorld fro m a single
two-dim ension.al image. Two eyes have other advantages over jus t one:
expanding the visual field, permitting binocular summa tion, and p roviding
redundancy if one eye is damaged.
4. Having h'io laterally separated eyes romtected. to a single brain a lso pro-
vides u s with important information about depth throu gh the geometry
of the small differences between the images in each eye. These differences,
known as binocular dispa rities, give rise to stereoscopic depth perception .
5. Random d ot s tereograms show that we don' t need to know what we' re
seeing before \\o"e see it in stereoscopic depth. Binocular disparity a lone can
support shape p erception .
6. Stereopsis has been exploited to add, literally, de pth to entertainment- 0336 AMma App
from nine teenth-century photos to t\venty-first-century movies. It has also
served to enhance the perception of information in military and medical
se ttings.
7. The difficulty of matching an image element in one eye w ith the correct ele-
ment in the other eye is known as the correspondence problem. The brain
uses several strategies to solve the problem. For example, it reduces the ini-
tial complexity of the problem by matching large "blobs"' in the low-sp atial-
h equency in fo rmation before tryi ng to match every high-frequency d etaiL

198 CHAPTER 6
8. Single neurons in the primary visual cortex and beyond have receptive
fields tha t cover a region in three-d imensional space, not just the two-
dimensional image plane. Some neurons Si:'eITl to be con cerned with a crude
in-front /behind judgment. Other neu rons are con cerned ....ith more precise,
me trica l d ep th perception.
9. Whe n the stimuli on corresponding loci in the two eyes are different, we
experience a continual percep tual competition behveen the two eyes knrnvn
as binocular rivalry. Rival ry is part of the effort to make the best g uess
about the current sta te of the world based on the current state of the input.
10. All of the variou s monocular and binocular depth cues are combined
(unconsciously) according to ....·ha t prior knowledge tells us about the prob-
ability of the current event. Making the wron g guess about the cause of
visual input can lead to illusions. Bayes' theorem is the basis of one ty pe of
formal understanding of the rules of combination.
11. Stereopsis emerges suddenly at abou t4 months of age in humans, and
it can be d isrupted through abnormal visual experience during a critical
period early in li fe.
8230336 Aml'la Appa

lq
8230336 Arnrna App.l

CHAPTER 7

Attention and Scene
Perception
IF YOU'Rt Tl-HS, you are probably a s tudent If you' r e a s tudent, you
are probably taking more than one course and are therefore very busy. H ere's
an idea: w hy n ot read t\vo books at the .sa me time? Chapter 2 will ha ve told
you that the lirnit on peri pheral acuity is on e reason tha t this won' t work. The
acuity problern could be overcom e if the size of the print were increased, as in
Figure 7 .1 . Nevertheless, even \Vith nice will be clear that you
ca nnot look at tl'le column of Xs in the figurennd the tvv sent \Ve
just cannot read hvo messages a t the sa me time. Note that you cnt1 read the
words on one side or the other of the Xs while looking a t the Xs. You jus t can' t
read both sides si multaneously. lltis is a specific example of n more general
problen"l-that the retin.11 array oon tains for more infom1.ition th.nn we can
process. Figure 7 .2 shO\vs another example. We canno t possibly recognize a.LI
the objects in this picture at once. That's why 11Where's Waldo?" a nd 111 Spy"
gam es are a di.allenge. This is no t just a visual proble m. All of the senses re-
ceive m ore input than we ca n handle (see Chaptel' 10 fo r a discussion o f at-
tel\tion in hearin g).
Why can' t we process everything at once? Quite literally1 we don' t the
brains for it. Remember from Chapter 4 tha t recognizing a single object like an
elephant requires a sizable chunk of the brain and its processing power, espe-
cially w hen that elephant could be seen in many different orient.'l tions, under
different lighting conditions, a t different reti nal sizes, and so on. Moreover,
in order to tmders tand Figure 7.21 ""'e also need to process the re lationships
beh\.•een objects-like the fact th<Jt the e lephant is dousin g the yellow car '"'rith
its trtmk. If we do the m ath for even a fairly s mall s ubset of all p ossible visual
stimuli, it turn s out that processin g everything, everywh ere, all at once re quires
a brain that will not fit in the human head (fsotsos, 1990).
If it is not possib le to process everything all at once, what shordd be pro-
cessed ? This matter ca n ' t be left todumce. If we 1re crossing a road, we need
to d eterm ine that no car \o\iJl hit us; devoting o ur visual processes exclusively
to the douglmut sh op on the other sid e of the street could be dangerous. To
with this problem, we ''pay attention" to some stimuli and not to others.
As we will see in this chapter, attention is not a si ng le tiring, and it d oes n ot
have a s ingle locus in the nervous system (C hun1 Golomb, and Turk-Brovvne,
2011). Rather, atteution is the name we give to a family of mechanisms th.at
restrict or bias p rocessing in various ways. attention My of the very large set
H ere are som e of the d istincti ons we can make when considering variet- o f selective prooesses in the brain. To
ies of attention: deal with the imposslblllty o f handling
all Inputs a! once, the nervous system
• Attenti on can be intemal or ex ternal. External nt.te11tio11 refers to has evdved mechanisms that are
a ttention to s timuli in the world (our primary concern here), but we able to bias processlru to a subset o f
s hou1d n ot forget internal alteution, our ability to attend. to one line of tt1lngs . places, Ideas, er rnornents In
thought as opposed to another or to select one resp onse over another. time.

202 CHAPTER 7
Is
FIG URE 7. 1 Even though the lett€ts we bi;J t1nough to resolve
These. m while lookin g at the Xs, WQ simply cann ot rE.IEld the left-hand and r1ght-
hand sent9f'IOQS at the same time.
letters x it • Attention can be overt or covert. O;,w1 atten tion usually

refers to directing a sense organ at a s tim.uhts--fixating
the eyes on a sin gle '"''ord, for example. If you point your
are x time eyes a t this page while directing attention to a person of
interest off to the le ft, you are e ngagin g in covert ntte11tio,1.
big x for • Reading this text while continuing to be aware of rnusic

playi ng in the room is an example of dii.•ided at.tentio11.
• Watching the pot to n ote the m oment the wa ter begins to
and x a boil is a ·iigilance task requiring sr:stained attentio11.

In this chapter, we \Vill be m ost co ncerned with selective
tentlon, the ability to pick one (or a few) out of many stimuli.
easy x quick These are not muh1ally exclusive tenns. Staring s traig ht ahead
while attending oaly to an interesting person to the excl us ion
of all else might be described as an act of ex tema t s us taine d,
to x snack cover t, selective atte ntion.
Althou gh we v..ill focus on visual attention. it is importan t
to remember tha t attentional mech anisms operate in all of the
read. x yet? senses. For example, getting a sho t in the doctor's o ffice may
hurt a lot, while an equi valent injury on the playing field m ay
no t seem to hurt much a t all , because of the different am ounts
FIG URE 7. 2 Search fo r the unicorn in this piec8 of a •wh9rB's picture.

You can search for WaJdo. too. but he is a bit sm all in this reprcducUon. (from
W7ltre's Waldo? 0 Martin Handford 2005.)

AITENTlON AND SCENE PERCEPTION 203
of attention we give to ead-i even L We can also u se attentional mechanisms to selective attention The form of
give pri ority to one sense ove.r othe rs . Right now, you are probably selecting attentlm Involved w11en processing Is
visual stimuli over audltory, even if you have music o n in the b ackground . restricted to a subset of the posslble
stimuli.
And you didn't even noti ce the pressure of your posterior on the sea t until
we mentioned it (sorry for disrupting your attention). reaction time <Rn A measure of
the time from the onset of a stlmulus
Jn this chapter, first we discuss eviden ce tha t •ve attend to only one (or to a response.
perhaps a few) objects at any sin gle moment. We consider attentional selec-
cue A stlmulus that might Indicate
tion in space lI am at tending to this object and not that object) and selection
where (or whan a subsa:iuent stimulus
in time (I , ...·as a ttending to that object, but now l'veselected this object). Next will te. Cues can te valid (gMng ca-
we examine the changes that occu r in the brain when \¥e attend to one object rect informatlm). lrnalid Oncarect), or
ra th er than another, and we ·will see what happens w hen the neural subs trate naJtral (uninformative).
of a tte ntion is da m;;;ged . Having con vin ced you, we h ope, tha t we ca n recog- exogenous cue In directing atten-
nize only on e object at a time, \\'e will tum to scenes. If '"'·e're a ttendin g to one tion. an exOgenous cue is located out
object, w hat d oes it m ean to say that we see a w hole scene? (exo) at the desired lnal location of
attentlm.
Selection in Space (a)
'
To begin, let's consid er w ha t it means to a ttend to a St.rt
stimulus. A good place to star t is \.\o·ith a cueing ex- *
periment of the sort pioneered by Michael
(1980). Start wi th thesituati on s hmvn in Figure 7.3a. Te&tprohe x *
The subject in the experimen t fixates on a central
•
point("'). Afte r a v,iriable d elay, a test probe (X) ap-
pea rs in one of the two boxes. All the subject needs
to d o is hit a respon se key as fas t as possible w hen
the probe appears. Thein eas.u re of interest is the av- (b)
erage reaction time (RT)-the 1.unount of time tha t
'
elapses between the point w he n the probe appears Time!
and the point '"'hen the s ubject hi ts the resp on se key.
*
Suppose now th.."'t the situation is s lightly changed:
during the waiting period, the subject is given a cue,
a stimulus that provides a hint about where the target
might appear. In Fig ure 7.3b, the cue is a cha nge in
T1me2
D
the outline color of one of the hvo boxes (ca ll this
36Amm
Tune3 x *
a ..-'periphera l cue"). Becau se the test probe appears
in the cued location, this peripheral cue is sai d to be This rue is valid .
a "va lid" cue. With a valid cue, Posner fmmd, the
RT decreases. Comp ared with the no-cue control (c)
'
situation, th e s ubject generally resp onds faster to
Time1
the probe because she is ''paying attention" to the *
correct location. In Figure 7 .3c, we have a different
kind of cue. 1l-1e red dot is a "sy mbolic" cue, but it
can also direct a ttention. Here the n1le is 11 1f the cue
is g reen, the probe is likely to be on the left. If the
cue is red, the probe is like ly to be on the right.''
In Figure 7.3c, however1 the cu e is misleading, or
Tune3 x *
"i nvalid/ ' because the cued location is on the right
This cu e is i.nl'alid.
but the p robe appea.rson the left. RTs areslmver here
than in the control condition because the subject h as FIGURE 7.3 Th9 Posngr cuoaing paradigm. (a) Thissi rnpl9 proba
been fooled into attending to the wr ong loca tion. detection experiment has two possible p-obe locations. H9re all ycu
have to do is hit one key If the p robe appears on the left and anoth-
The peripheral cue (the outlined box) i s an ex-
er If it appears on the right . (b) A valid cue indicatQS w hefe the ta rgiic>t
ample of an exogenous cue. Exogenous cues seem 'Will be. This time the cue Is valid; we are tellirq the truth . (c) AA
to s ununon a tte nti on by vi rtue of invalid cus points to the wrong side. (fhis is also a different kind of
their ph ysica l salience. Th e symbolic cu e (red box) cue -a symbolic cua. Symbolic c ues direct attention more- slowly.)

204 CHAPTER 7
!8
T woliccu•
FIG URE 7.4 The effGCt of a cue
devebps over time. A peripheral cue
Oika tha t in Figure 7 .ab) beco1Tl9S fulty 20
effective 100- 160 ms after It
A symbciic cue (see Figure 7 .3c) t akes
1
lcf'tgeir to g et started and to rise to full
Giffect. 6 10
_g
"'j 0 100 200 30'.l 400

Stimnlu.s on.set asynch rony (SOA) (m.s)
endogenous c ue In directing atten- is an exa mple of an endogenous cue. Endogeno us cu es can be considered
tion, an endogenous cue Is located In some thing like instruction s tha t can be volunta rily obeyed . The end ogenou s
(endo) or near the current location of cue says go right, so you go right, while the exogenou s cu e could drag you to
attentrn . the right, w h ethe r you w anted to go there or n o t. £11do and exo refer to "insid e"
stlmuus onset asynchrony and ''outside,'' respective ly, as in om endoskeleton and the exoskele ton on the
(SOA) The time between the m set outside of a n insect (Posner, 1980).
of one stlmulus and the onset of
another. Pe ri phera l/ exogen ous and sym bolic/ endogen ous c ues can be valid or
invalid . As in Figure 7.3c, an invalid cue rni ght te U you to go right, w hen
inhibition of return The relative
the ta rget actua ll y ap pears o n the left. 1.n a typical exp e riment, the cue mi ght
ficulty In getting attentlm (or the eyes)
to move back to a recently attendOO be valid on 80% of the t ri<1ls and invalid on the re m ain ing 20%. Since the
(cr fixated) location. observer d oesn' t kn ow if the cu e is valid, it would be in the observer 's in te r-
est to d e ploy attention on the assumption tha t th e cue is v alid. TI1is means
that the exp erimenter can compai e RTs from trials tha t had va lid cues, where
nttention was cued left and the s timulus a ppeared on the le ft, and trials that
had inva lid C\J es, w he re the cue moved a ttention to the left but the s timulus
ap pea red on the rig ht.
How long d oe5 it take for a cu e to redirec t orn a ttention? It d epen ds on the
n..1ture of the cue. At the beginning of a trial in a Posne r cuei ng experiment,
the s ubject is a ttend in g to the fi xation point (time 1 in Figure 7.3b or c). The
rue appears at time 21 a nd the probe a pp ears a t time 3. We Cclll measure the
timing of the attentional s hift by v arying the in terval behveen time 2 and time
3. Th is is called the stimulus onset asynchrony, a psychophysical va ria ble
that typicall y goes by its aa:on}'m , S OA. lf tl>e SOA is 0 millisecond s (ms ),
the cue and probe appear . There is n time for the cue to be
used to direct atte ntion, a nd there is n o d ifference beh Yeen the effects of valid
or in va lid cues. As the SOA increases to a bout 150 ms, the magni tude of the
cu eing effect from a valid peripheral rue increases, as s how n by the red Line
in Figure 7.4. After that, the e ffect o f the cue levels off o r declines a bit.
Symbolic/ end ogeno us cues, sud 1 as the colored do t, ta ke longer to work,
presumably because \ Ve need to d o some work to in terpre t the m (the SOA
is shmvn by the blue line in Fig ure 7.4). In teres ting ly, some rathe r symbolic
cues beha ve like fas t, periphe ral cu es. For examp le, we are very quick to
d ep loy m u· a tte ntion to arrow cues and even faster to resp ond to a pair o f
eyes looking in one direction or anothe l' (Figure 7.5), as if we were built to
get infom1ation frorn the gaze of o thers (Kuhn and Kings tone, 2009). You can
h·y the cueing experit11e nt in some of i ts important va ria tions in Web Activity
7.1 : Attentional Cueing.
FIG URE 7.5 \IVhlch \Naf does your ln terestingly, if you m ove your attention or your eyes to a loca ti on a nd
atttintion shift w hEf'l you lock at this then look away from that location, it is harde r to look back a t it again for a
picture? littl e w hile. This is known as inhibition of retum (Klein, 200J). lnhibitio11 of

ATTENTlON AND SC ENE PERCEPT! ON 205
re turn he lps to keep yo u from gettin g stuck continu ally revisiting one spot.
For ins tance, s uppose yo u were loo king fo r Wa ldo in Figure 7.2 and your
attention was attracted to the S.m'ta in the lower ri ght lf yom a ttention kep t
goin g back to Santa, that would be rathe r useless. Inhibition of rehun helps
your sean:h to m ove fon•lard (Klein Maclnness, 1999).
The "Spotlight" of Attention

In a cueing experirnent, attention star ts at the fi xation point and som ehow ends
up at the cued loca tion. But d oes it actual ly move from on e point to the nex t?
Attention could be deployed from spo t to spot in a ntmlber of ways. It nl ight
tn ove in a rna nn er analogous to the Jnovements of our eyes. \Vhen v.<e s hift
our gaze, our point of fi xation sv1eeps acros.s the in tervening space (alth ough,
as we w ill see in Chapter 8, "saccadic s uppressio n" keeps us from n oticin g,
and being disturbed by, th.is rnovement). Attention might sweep across space
in a si milar rna 1mer, like a spotlight beam (Posner, 1980).
The sp otlight metaph or makes good sense and has become, perhaps, the
1nost common way for cognitive psycho logists to ta lk about attention. But
there other possibilities. For example, attention might expand fro m fixa tion,
growin g to fill the whole region from the fixation spat to the cued loca tion, and
then it migh t shrink to include just the cued loca tion. This would be a vers ion
of a zoom lens model of a ttention (Eriksen and Yeh, 1985). Or, when attention
is \vithdrawn fr om the fixa tion s pot, it mi ght notrn oveatall. ltmight s imply
melt away at that 1ocation and then reappear at the cued location (Sperling and
rt
1995). is hard to say for S lU"e which metaphor is ''correct,.''
because we h ave no direct way to measure the loca tion a nd extent of attention.
H owever, the bes t e·dden ce s uggests that attention is not moving from p oint
to p oint in the brain in the way a physical spotligh t \'\.·mild move across the
world (Cave and Bichot, 1999). Mmeover, there is no guarantee tha t there is
just one "sp otlight." It m ight be split (loo k ahead to Pigure 7.16) (McMains .'.lnd
Somers, 2004), though m ost of the time, visual selective attention is prob.ably
directed to one thing in one loca tion.
Visual Search
Cu ein g experiments provide important ins.igh t into the deployment of attention.
But the sihta tion is rathel' artificial, in that all of the experiln ents involve telling
the observer exactly when and w here to attend. Visual search experiments
provide a closer approxi1nation of some of the actions of attention in the real
world . In a ty pical visual search experirnent, the observer looks for a target
item among dlstractor ite ms. Visual searches are ubiquitous in the real world:
we look fo r faces in a crowd , mngs in a cupboard, books on a shelf, and so
forth. Some sea rd 1es are so easy that '"·e hardly think of the m as searches (e.g.,
finding the cold-w<tter tap on a sink). Others, like looking for the unicorn in
the"\ Vhere 's Waldo?'' puzzle of Figme 7.2, are much m ore d emanding. Some
are, li terall)i matters of life a nd d ea th, as when radio logists look fo r h1mors in
X-rays or airport securi ty officers look fo r threats in luggage.
The quest to und erstand what makes some search and others hard
has proven to be o ne o f the m os t prod uctive and inte res ting lines of cogniti ve visual search Search for a target
psychol ogy resea rch in the past quarter century. Rgure 7 .6 s hows examples In a display containing distracting
of the simplified visua l search tasks often used in the lab . In Figures 7.611 and elements.
b, the target is a red vertical bar. In Fig m e 7.6c, the target is the le tte r T (in any target The goal of a Visual search.
of fo ur possible rotations). Two factors are be ing ,va ried in First, dlstrac tor In visual search. any
tnoving across the figure, the tasks increase lh difficultyfro m left t rig ht. stlrruus othef than the target.

206 CHAPTER 7
(11) Feature search (b) Conjunction seaHh (c) Spatial configuration

ioe.nch
I I
-
find Find Find T-1.l o' t-
L
I I
I
I
I ... , i
I
- I I .J
--- -- - -- , I I L L i
-- -- -
I I
I i
I I T .J .J
I I I I1 -
- - r, ,
- I I L.J L .J
- ·---···
.J
111 I I i
I 111 I I ,i T L
j- Correct target p re&ent - Correct target absent I
] 23U33b AM Tl
I§ <$'.
" -Oms/item
\0 . 11'
"'
Setslze
FIG URE 7.6 Laboratory \'isual search tasks. Each part o f tM figure shows a differ-
cmt task. with diffic ulty inc rQaSing frcm (a) t o (c). Each raw shovvs a diffQr1t1nt
number of items {the set size), w ith difficulty increia.sing as set size increases. Here we
show onl)' .;;ocamples Wth the target present. In o. typical QXPeriment, a target might
00 absent in half o f the trials. The graphs at the bottom depci typk:al patterns of
results for each type of task. The purple line in each graph represents average reac-
tion times for different set siz9s o n target-absent trials: the green line shOW'S rQSults
for target-prtiiSent trials.
Second, the nmnbe1· of items (the set size) increases as we move d own ead1
column. As a genera l (and unsurprising} rnle, it is harder to find a target as
the set size increases.
To m easu re the efficien cy of a visual search, ofte n we ask h o\"' mu ch
time is added (on average) for each item <tdded to the d isplay. To find out,
the experimenter measures the RT required fo r the observe r to say "yes" if
the ta rget is p resent or r:no'' if there is n o targe t in the display an d observers
perform the same type of search over an d over. The hmctions relating RT to
set size The number of Item s in a set size are graphed a t the bottom of Figure 7.6. The efficiency of the search
>Asual display. is described by the slope of these RT x se t size functions OUgher s lopes mean

ATTENTION AND SCENE PERCEPTION 207
lower efficiency). Efficiency describes the ease with which we can work our 1eature search Search for a target
way through a display. It is a way to cornpare search tasks. lf we can direct denned by a slr>gle attnbute, such as a
attention to the target as soon RS the display appears, regardless of the set size, salient color or
then we have an efficient search like the feature searches in Figure 7.6n . Here salience The vMdness of a stimulus
the slope is near zero because adding more blue d istractors or more horizonta l relative to Its neighbors.
distrac tors d oes no t make it a ny harder to fmd a red or a vertica l target. ll paraUel search A eeai:d:l Ii which
we must examine each item in tum until v.;e find the target1 then we have an multlpte stimuli are processed at tre
inefficient searc h. Figure 7.6c shows an ineffi cient search w here each L needs same tme.
to be exa mined until the observer stumbles upon a T. Notice that sa)'ing 11 yes,
the target is present'' is typically faster and m o re e ffi cien t than sayin g 1' no, it
is n ot/ because even in the hardest d lucky o bserver might stumble on
the p resence of the target VY; th her first deployment of attention, but it is n ot
possible to s hunble on the absence of the target in the sarn e way.
Although all the displays in Figu re 7.6 include a target, in a typ ical experi-
ment the target is present in 50°/u of the trials and absent in the other 50%. You
can try a number of different visual search tasks in Web ActMty 7.2: Visual
search. It's much easier to get a feel for the difficulty of these tasks when you
do them yourse lf.
Feature Searches Are Efficient

The task in Figure 7.6n is called a simple feature search. Here the target is
d efined by the p resence of a si ngle fea tu re. Each example contains a n item
1..vi th a unique color or orien tation If the unique item is s ufficiently salient (i f
it sta nds out visually from its neighbors), it really doesn' t ma tter how
d istractors the1·eare. TI1e target seems to "p op out'' of the display. Apparently
we can process the color or orientation of all the items at once (often called
a parallel search). When we measu re the RT, it
does not change with the se t size. The res ul ts wi ll
approxirnate the flat lines plotted a t the bottom of (a)
Figure 7.6n; more tedm ically, the slope of the fun c-
tion relating RT to set size is about 0 m s per item .
Between o ne d ozen a nd two dozen basic a t-
tributes seem able tosuppOTtparallel visual search
(\ Volfe and Hormvitz, 2004). TI1ese include obvious
stimulus properties s u ch as color, size, orientation,
and motion (A. Treisman, 1986a, 1986b) and sorne
less obvious attributes, like lighting d irection (Enns
and Rensink, 1990).
ln Ffgure 7.7a, look for the bar that is oriented
differently in de pth. This task is quite easy; even
though a ll th e item5 have the same orien ta tion in
the two-dimen sional plane of the page. Notice tha t (b) - - - - - - - - - - - - - - - - - - - -
though all the bars h ave the same oriQl'ltation in the

two-dimensional image plar1e. show that the
ease o f the task had to do w ith the threre·dimen ·
simal eff•cts and not just w tth tho relotiw posttions of
diffeirent regions, the expeirlrn9nt was ref)Qated w ith
appareintly two·dim811slonal items composed o f similar
parts In similar positions. This made it hard€4" to fin d
the odd (Mer Enns and Rensink, 1990.)

208 CHAPTER 7
rotation of the distractor objects, and this rotation du=mges its depth orienta-
tion. ln Figure 7.7b, the target is also a 180-degree rotation of the dis tractors,
but you will probably find it harder to locate the target that has light green at
the top rathe r than the bottom. That's interesting because the objects in Figure
7.7a and bare all composed of elongated w hite and dark green regions with
a smaller light green region a t one end . The overall shapes and orientations
of all objects are ttbout the sam e. The critical dmnge in Figure 7.7b is that the
objects no longer look like three-dimensional bricks and no lon,se r differ in
their apparent orien tation in depth. Without that distinguishing fea ture, the
light-green-on-top target is harde r to find.
Many Searches Are Inefficient

serial self-terminating search When the target and dis tradors in a vis ual search tas k contain the same basic
A search Item to ending features, as in Figure 7.6c, seard1 is ine fficient . H ere, we are looking for Ts
w hen a target Is found. among Ls w here bo th targets and d istractors a re composed of a vertical and
a hori zontal bar. It is n ot hard to distin gui sh a T fr01n an L, but we need to at-
tend to eac h item until we stumble on the target. Even il, as in Figure 7.1, the
le tters are big en ough th at \Ve d on ' t h Llve to move our eyes to distinguish T
from L, each additional distrncto r adds about 20-30 ms to a s uccessful search
for a ta rge t, and about h vice that am ount of time (40-60 ms) to a search that
ends with out findin g a target. We call this an inefficie nt search .
\Vhy d o we get the parti cular patte rn o f resul ts seen in Pig u re 7.6c? One
s traightfon1,.·ard propos..'l.1 is that these tasks invol\.·e a serial self-terminating
searchr in which items me examined one a fter a nother {serially) eith er until
the targe t is found or lmtil a ll ite ms have been d-1ecked . Consi der the case in
Fmd:
8230336 Am111a J
"Grace"'
·-
/flll
4-1- 4-1-
FIG URE 7 .8 Search can be muc h
rnore laborious if you 're not furniliar 'Mth
what you 're searching for. Here. the
89arch for the C hinese c haract91'" for
"grace" w ill be muc h easier if you can
read Chinese.
·- 17L
/fill
4-1-

w hid1 a target is p resent Sometimes the observer \vi ll get lucky and find the
target on the first deployment of attention. Sometimes she w ill be unlucky
and have to search all the items befo1-e shunbling on the huget. On average,
she \'ViJI have to sea rd1 through about half the ite1:ns on each tr:i<1l before the
search can be temlinated . \ Vhen n o target is present, our observer will a lways
n eed to search t!U"ou gh all the items. Th ere are o ther accounts that wo uld n ot
involve an exhaustiveexa rni.nation of each item Q. Pa lmer, 1995). The in'lportant
point is tha t sornesearches are ineffi cient, meaning tha t ead1 at.1 ditional item
in the d isp lay irnposes a significant coo t on the searcher.
The ine fficient seard1 for a T a mon g Ls is nothing like the hard est search
we could devise, of course. For s ta rters, if we need to spend a lot of titne wi th
ead1 ite rn, search will be much slower, as it is in the search for a particular
Chinese characte r in Figure 7 .8 (tmless, of course, you happen to read 0-Un ese).
Other sean:::h tasks, like spotting 1nissile s ites in satellite images, could take
days even if, o nce you found one, the missile si te was readily id entifiable. The
T -am ong-Ls search is inte resting because it is a si mple r case w here a target ca n
''hid e in plain sight.1' lhe items are ea::..y- to see and very familiar, but search
is s till ine fficient.
In Real-World Searches, Basic Features Guide Visual Search

ln the real wor ld, it would be very rare to h ave a true feature search for the guided search Search In w hich
only red item a rnong hornogeneous distractors . H ow often d o you have to find attentlm can be restricted to a subset
the strawberry a mong the limes? l t is p robably even rarer to have to search of pooslble Items on tl1e basis of lnfa-
matlon about the target lteYJ 's basic
through a scene containing objects that aJl s h are the sa me bask fea hue:r- features (e.g .. Its color).
perhaps the proverbial needle in the haystack. Usually, bask featurEs can be
used to narrow do\"'11 the sean:h, even if they c.annot eliminate all d istractions.
This is know n as guided search (Wolfe, 1994; Wolfe, Cave, and Fra l\zel. 1989).
[f you're looking for a tonMto in Figure 7 .9, you w ill be a b le to find it quite
quickly, even though it is not the only red thing or the only round thing 01·,
indeed , the only possessor of any unique feature. Tomatoes a re distinguished
from most of th e d istractors by a conjunction of severa l fea tures . If you can
guide yo ur a ttention to red and round ru1d large, you wi ll have eliminated
FIGURE 7.9 A real-wortd conjunction

search. Find the big . round, red toma-
togs arnong things that might b9 big or
round o r red but do not haw all three
basC f00tures.

210 CHAPTER 7
FIGURE 7 .10 SQ arch for arbitrary

objocts is not V9ry efficient. Find the
fauCQt , thQ violin, or the vvindow.
most of the competi tion. In the lab, we use conjunction searches like those
in Figure 7.6b. ln a conjunction search, no sing le fea hue de fin es the ta rget.
lns te:Jd, th e t.nrget is d efined by the conjunction-the co-occurre nre-of tv.•o
82303 OJI rmneJeaturesd n this case \Ve' re loo king for the red, verti cal items. In terrns
of efficiency, conjunction searches tend to lie between the very e ffi cient
searches a nd the inefficient serial search e:.;.
In Real-World Searches, Properties of Scenes Guide

Visual Search
If guidan ce by the basic features of the target object were the whole solution
to findin g objects in th e real world, then search for a fau cet in Figure 7.10
sho uld be efficient . The searche r would think of the basic fe atures of faucehr-
shiny, s ilver, rounde d-and then the search for a fau cet would be a matter of
guiding at tention to a complex conjunction of fecttures. H owever, search for
arbitrary objects is not efficient (Vickery, King, and Jiang, 2COS). Certainly, fea-
ture guidance can help--for example, look for the green paintbrus h in Figure
conji.nction sea-ch Search for a 7.10--but the ease with w h..ich we search in the real world mus t involve more
target defined by the preser>0e of two tha n feahlre g uid an ce.
er more attributes (e .g., a red, vertical It is quite easy to find the faucet in the kitchen scene in Flgure 7 .11, even
target amorxi red horizontal and blue
though it is less: visible tha n th.e faucet in Figure 7.10. You are he lped by va rious
vertical dlstractors) .
forms of scene-based guidance (Castelhano a nd Heaven, 2010). Even though
scene-based glidance lnfolllla- you 've ne ver seen this kitdle n before, you know about kitchen s. Kitchens
tlon ln our underatandlrg of scenes
that helps us fird specmc cbjects In
ha ve sinks in typica l places. Sinks hm.:e faucets in typical places, and the re
scenes (e.g ., objecta. do not noat In air, is the faucet. If you were searching for bowls, you would be gu id ed by the
faucets are near sinks). knowledge that bowls must rest on s urfaces; they do not fl oat. Moreover, you

ATTENllON AND SC ENE PERCEPll ON 211
binding problem The c hallenge of

tylr.;i diff«ent attributes of visual sllmull
(e.g .. oolor, orlentallm, motlofl). which
are handled by different brain clrc ulls,
to the appropriate object so that we
perc eive a unrned ci:Jject (e.g., red, ver-
tical, mo\rjng right).
feature integration theory Anne
Trelsman·s theory of visual attention.
which holds that a llmlted set of b as!:;
features can be pr(X)Sssed In parallel
preatt€<1l lvely, lllrt trat other proper -
ties. lncludlrg the correct binding o f
features to objects. require attention.
preattentive stage The process-
ing o f a stimulus trat occurs before
selective attentlai Is deployed to tl13t
stimulus.
FIG URE 7.1 1 Scene-based gui::tt'11lc.& would help you find the faucet in this scene.
know about the average size of bowls. 1l'lis knowledge would allow you to
figure out that only a few objects in this scene are plausi bly bow l-s ized in the
space d efined by this picture. In that case, that g uida nce would speed yo m
conclu sion that there are n o bowls.
We can think of all of th.ese various for ms o f scene guidtmce as constituting ++++
t++tt
a Bayesia n " prior p robability'' tha t tells you how like ly it is tha t a ny given
object in the scene i.s the targe t (E hinger et al, 2W).
FURTHER DISCUSSION of the Bayesian approach oan be found in Chap-

ters 4 (page 11 0). 5(poge150), and 6 (pages 100-187).
The Binding Problem in Visual Search

++++
++++
We '¥ill return to cornplicated s timuli like scenes later in this chapte r. First,
we n eed to s tep back and cons ide r w hat selective-attention does for us. One
ans wer1 championed by Aime Treisman (1996), is tha t selective attention a l-
++++
lows us to solve the binding problem. Th is problem is illustrated in Figure
7.12. A quick glan ce tells us that we_,re looking a t pluses whose components
are red and blue and vertical an d horizontal. Howeve r, if we need to find the
h \'o red-vertm l,.,blue..horkontal lte1 / w will need to searc h, cllrecting a t-
tention to the individual items (Wolfe and Bennett, 1997). In the language o f
Treisman's 1eab.Jre Integration theory {Treisman and Gelade, 1980), the bas ic
features of color and o rientation are avai lable in a preattentlve stage of p ro- ++++
cessing, but we d on 't know how those fe..i.tures are bound together until 've
attend to a specific object. Loo king back a t Figure 7.6c, the problem '"ith Ts
and Ls is tha t they both have vertical and horizontal fea tures. It is the rela-
tionship of those features to each other that's cri ti ca l. To recognize an object,
+++
its features mus t be 1'bow1d.''
FIG URE 7. 12 A oonjunction S9arch w ith a binding pro bhilrn. In this imag9 it is hard
to find the red vgtlcals (thge are two o f thetn) . because all of the items have the fea-
tures red and blue and vertbal and horizontal.

212 CHAPTER 7
ll'J.e idea of guided searchr m entioned earlierr fits into this frainework. We
ca n use some preattentive featu re information to guide the choice of w h at
F L y
sh ould be attended and botmd next. lf we re searching for a Iit tie, brown rnouse,
we s hould guide our a ttention to Jittle s tuff and brovro s tuff and n ot d evote
attention to shiny, red or large, square things. Even before attention can enable
binding, the mere pre:ience of the right collection o f unbound fea tures might,
K p in some cases, very quickly gj.ve u s a pretty good idea that th ere is a m ouse
(or a t leas t an animal} present (Li et a l., 2002), but full object recognitio n is
w H goi ng to require attention.

lf attention is needed to bind feah.ues correctl)i what happens if we don' t
have en ough tirn.e to complete the job? \Ve'll see somethin g, but what? If it
works for you, the demon s tra tion in Figure 7.13 (on the next page-but d on' t
ollJt. yet) will p rovi de pa rt o f the a ns wer. What you mus t d o is look quickly
to rec.:.'1.11 the colors of the letters. You at the figure, and then loo k ah·ay and \'\rrite d mvn as many of the lette rs and
may know what cciors W91'& thst-9 and their colors ns you can . Try that n o\\'.
you may rern9fT"lber many o f the letters,
but you are q uite llkaty to pair a color
lf you compare your list to the ach1al figure, yo u may find that you matched
and a lette< inoorrectly. the wrong color to a letter. For example, you mi ght be quite s ure tha t you
s..1w a green H. If so, you ha ve e.xperie.nced an Illusory conjunction, a false
combinati on of the fea tu res from two or m o re different objects (A_ lvl Treis-
man and Schmidt, 1982). You are lllilikely to think that you have seen a blue
Hor a green T, beca use neither the color blue n o r the letter Tis p resent in
the figure. We conjo in only those features tha t are actuall y in the display.
When we can' t complete the tas k o f binding, we d o the best we can wi th the
infor mati on we h ave.
Attending in Time:
RSVP and the Attentional Blink
So far in this chapter1 we've con c:entrated on atte ntional selection in space.
Let's consider what hap pens if you sea rd1 in time rnther than in space. Imagin e
the fo llow ing s itua tion : You 're looking at a s trea m of le tters that all appea r at
thes..1 me location in space. Showi ng the s timuli in th is way is kn own as rapid
serial visual presentation (RSVP}. You're trying to d ecid e w hether the re's an
X in the strearn of letters. How fast can the characters be presen ted and s till
permit you to d o the tas k vdth high accuracy? Wi th fairly large, clearly visible
s titnuli, we ca n reliably pick an X out of letters when the characters are appear-
ing a t a rate of 8-10 items per second. The task does not need to be lim ited to
Illusory conjunction An errcneous simple characters. It we're wa td1ing a stream of photographs flash by at this
comblna:ttn of two features In a visual rate, we can m oni tor that stre...9.m for the appearan ce of a picnic photograph.
scene-for example, seeing a red X 'Ale d on 't even need to know w h ich particular pia Uc we're looking fo r. The
when the display contains red lettern general idea of 11piaUc" is adequate for the task (Potter, 1976). You can view
andXs but no red Xs.
scenes in an RSVP sequence yourself in We b Activity 7.3: The RS VP Paradign_
rapid serial visual presentaUon Returnin g to s imple letters, let's chan ge the. task a bit lnste:ld of looking
(RSVP) An ""P•rimental procedure fo r one target in a stream o f le tters, n o w we're looking for two. \Ve can ca ll the
In whlcl1 stimuli appear In a stream
at one location (typically 1he pcint of first "Tl " (for "targe t mm1ber l ") and call the second "TI." In this
fixation) at a rapid rate (t)'plcally about Tl is a white letter in a s tream of black letters, and T2 is an X (Figure 7 .14a).
eight per second) . 1l1e X w ill be p resen t in 50% of the trials. The cri tical is the interva l
attenUonal blink The tender>OY not behveen Tl a nd 12, and the somewhat s urprisin g result of the experi ment is
to perceive cr reepord to the second sh own in Figure 7.14b. lfT2 appears 200--300 m s afte r Tl and if Tl is correctly
of two different target stlmull amid a reported 1 \oVe are very likely to m iss T2. (Note that each frame in Figure 7.14n
rapid stream of distracting stimuli If the and each position on the x-axis in Figure 7.14b represents 100 ms of time, sin ce
OOserver has responded to the first
target stimulus within 200-000 mll- le tters appea r every 100 ms.) This phenomenon is kn own as attentional blink
llseccnds before the second stimulus (Shapiro, 1994); it is as if om ability to visually attend to thed1aracters in the
Is presentOO. RSVP seque n1."e is temporarily knocked out, even tho ugh ou r eyes remain

AITENTJON AND SCENE PERCEPTION 213
w ide open. Ymf'll find more on this topic in Web Activity 7.4 : The Attentional
Blink and Repetition Blindness.
To show that missing T2 in this s·c ena.rio is an attentional problem , we can
d o a cont rol experiment in which subjects report only TI. If subjects are asked
to m onitor the RSVP strea rn for the T2 X, they pay no particular attention to
the n ml\' irrelevant white letter, and T2 perfonn.'l.nce
. is unifonnly good at a ll
delays between Tl and T2.
Somehow the act of attending to Tl makes it very hard to attend to T2
if T2 appears 200-300 m s later in the stream . What is h.appening? Marvin
Clum suggests the following metaphor: Imagine you' re fishing \'lith a net in
(a)
+
Letter on, D
15ms
Letter off,
85 ..,,;
Tl
Tl, a white letter
Position2
x
T2, the letter X (.ifter Tl)
(b)
100
FIG URE 7.14 T healtrantional blink(AB)in

\'idea game and n on-v ideo game players.
(a) In this AB ol::lsefvers try to report
- Video g.;wne pl.a •ers two targgts: a white leitter (T1), and an X
ga:m pli.yer.s (T2) c oming In the stream after that white
lett9r. Letters appear eivsy 100 ms. T2
can appear in various positicn s in tl"'l9 lis t
after Tl . Here T2 Is In p osition 2. (b} If too X
a ppears w ithin 500 m s (positions 1-5 after
Tl ), detection Is impaired. That im pairment
Is known as th9 b llnk. Video
Position o fT 2 in list reL1tive to Tl game play.a.rs show IQ!S.S o f a blink.

214 CHAPTER 7
(a) (b) (c)
-
Time
FIGURE 7.1 5 Mal"\lin a-iun 's fishing m9taphor for att9ntional blink. (a} You can SQQ
all the stuff In the river as it drifts by (e.g .. the boot and the fi sh}. fP) You commit to
fishing o ut fish numbGf 1 (Fl ). The boot drifts by, and fish 2 (F2) appears. {c) & cooS'9
you're tied up w ith Fl . you wi ll no t 00 able to fish out F2. and that seccnd flsh swi ms
by- det ect ed perhaps. but uncollected .
a less-than-pristine s t rea m, wi th boots and tires and the occasional fish fl oat-
ing o r S\vi mmin g by. You can m onito r the s tream a nd identify each item as it
p.1sses-boot, tire, boot, fish, boot, and soon. You can also d ip your ne t in and
catc h a fish (Figure 7.15a). Once you have a fish in the net, however, it takes
some time before yo u ei:i.n get the net brick into the water to ca tch fis h numbe r
2. As a result, you n1ight miss fis h 2 ifit s,,,,· ims by too soon after you 'vernught
fish l (Figure 7.15b,c) (Chun and Po tter, 1995).
Looking back at Figure 7.14b, notice that performance is quite good if T2.
appears imme diately after Tl . In Chun ' s met.'l phorical accoLmt, this would be a
case in which on e scoop ne ts tw'o fis h. 1ilis metapho r illus tra tes the idea tha t h vo
p rocesses a.re a t work. Notice the interesting difference beh veen the role a Hen tion
p lays here and the role it pbys in the spatial ex.ample of visual search. In search,
there is a capacity limit you ju.st don ' t seem to be able to recognize m ore than
on e object at a time. In attentional blink, though, the problem is probably not
the size of the metaphorical net. The p roblern is that, on ce attention has dipped
tha.tnetinto the river, there is a tem porary inhibition (Olivers and Meeter, 2008)
or a tem porary loss of control (Di Lallo et al., 2005) that ma kes it irnpos.sible to
coordinate a second dip fo r several hundred miJliseconds.
Figure 7.l4b sh ows d<tta from h-vo groups of subjects w ho pe rfor med differ-
en.tly n thea ttentiomll blink task. Jnteres ting ly, the group that shows a s maller
8230336 a ttenti al blink consis ts of playe rs o f 11 first-person s hooter '' video ga mes.
Indeed, those g..1 mers seem to d o better on a ra nge of a ttentional tasks (C. S.
Green and Bavelier, 2003). Why? It cOLtld be that people who d 1oose to play
these v ideo ga mes have better attentional resources. Alte rnatively, the gam es
themselves might e nhan ce attention. To test this idea, Green and Bavelie r (2003)
conducted a s tudy with two groups of n on-video g ame players. One group
got exp e rience with a fi rst-person shooter game, and the othe r group p layed
tile ga me Tetris . 1lle first-person shooter g ro up improved on a ttentional tas ks;
the Te tris group did not. Of course, this is n ot an end orsemen t of violent video
games, but it does su ggest tha t a ttentional abilities can be changed by training
nnd that those firs t-person sh ooter video ga mes produced chan ge for reasons
that remain to be worked out. L, Web Ac tivity 7.4: The Atte ntional Blink and
Repe tition Blindness, you can experien ce a related but not identical effect.
Effects like the attentionaJ blink a re important b eca use they tell us some--
thing about the tim e required for acts of a ttention and / or selection. From these

ATTENllON AND SC ENE PERCEPll ON 215
RSVP experiments alone, we can see that there are several "speed HnUts" in
om mental life. The s ingle-- target RSVP experiments tell us th at if we rue just
watching a s tream of stimuli over time, we can speed a long at a rate of many
irn.ages per second (perh.:ips .JS many as 75 per second; Potter et al., 201 4). A
mud1 sl ower rate describes the s peed with whid1 we can act on items, even
if we are just grabbin g them to rep ort about their p resence. Here, the RSVP
data suggest a speed !Unit of 2-5 item s per second.
The Physiological Basis of Attention

In an oft-qu oted passage, William James d ecla red, "Everyone knows what
attention is." TIUs great nin eteenth-century psychologist went on to say, ;' It is
the taking possession by the mind, in clear and vivid form, of one o ut of w hat
seem several simulta neously p ossible objects or trains of tho ught" Qames,
1890, Vol. 1, pp. 400-404). Th.:'lt seems fair enough, but w hat does it mean for
the mind to take possession o f a possible object? lf \Ve try to get specific, we
discover that thi s attention th..'l.t everyone ''knov."S'' can be rather difficult to
pin dovvn. In part, it is difficult to know wh at attention does because attention
performs a variety o f tasks, as already discussed. But we also need to d eal
wi th the physiological ques tion of w hat the brain is achtaUy doing w hen it
selects one location in space, or one object, or one moment in time, for further
processing (Carrasco, 2011 ). Let's consid er some of the neural possibilities.
Attention Could Enhance Neural Activity

lf we are asked to attend to one loca tion in the visual field, as in a Posner
cuein g task tsee Figure 7.3), the neurons th..-it respond to stim uli in that part
of the field w ill become more active. ltused to be thought tha t this response
dld not apply to s tria te cortex, the first part of visual cortex to process visual
input {see01apte.r 3). However, fun ctional magn etic resonance im aging (t1v1RI )
expe riments in humans, as well as electrophysiologicaJ studies of m on keys,
have shO\vn that even the first stages of cortical processin g are influenced
b y attention (Figu re 7.16 ) (Brefczynski and DeYoe, 1999; Gandhi , Heeger,
and Boynton, 1998; Hoeru1)'«1nd Schiller, 1988; McMain s a nd Somers, 2004).
As we progress further into the visua l areas of the cortex, even larger
attention al effects a re seen (Reynold s a nd 01elazzi, 2004). In fact, the effects
seen at the early s ta ges in the cor tex are quite possibl y the resu lts of feedback Subject B
from these late r stages of processing (Martinez e t al., 1999). 11,at feedback
m.ay be a very important part of visual processing (Ahissar and H ochstei n,
2004i Di Lollo, Enns, and Rensi.nk, 2000).
Attention Could Enhance the Processing of a Specific Type
of Stimulus
The m echan isms of attention d escribed in the previous section mlght s uf-
fice if we wa nt to keep watch "out of the corner o f the eye'' by enhancin g
rictivity in a specific region of visual cortex. But imagine picking pennies
out o f a change bowl (a rea l-wor ld visual search task). Somehow the wish to
FIGURE 7 .16 Spotlights of attention
find pennies m akes the pennies more salient. [f we sv.ri tched to quarters, the human brain. This fMRt irnage of
pennies wo uld recede in salience and the quarters would rise. In th is case, human visual oortQX (striateo and someo
we are a ttending not on the basis of spatial loca ti on, but on the basis of a extrastriate) shows activity in the brain
stimulus p rope rty. w hile subjects p aid attention to
stimulus (Att 1) o r t\11.io stirnLJli (Att 2).
You can experience this attentional shift in the s.:-tlience of differentstimulus
The stimuli were out side tM fovea. so
attributes in Figure 7 .17. \'\o'ithout mov ing your eyes, select the red items and thei rQgions of activity don't include the
notice h ow an oval emerges with no chan ge in the s timul us. Si milarly, you fovoo. (From M cMains and Scmers,
can select the blue items or the h orizontal items. Each act of selection seems 2004.)

216 CHAPTER 7
8230336 Aml"la Appa

fusiform lace area (FFA) A region
of extrastr1ate visual cortex In humans
that Is specmraly and reiably acttvated 1
······ ·- ....
by hUman faces.
pa-ahlppocampal place area 11.!J • • , ,• • • •••
(PPA) A reglcn o f axtrastrlate >Asual
cortex ITT humans that Is specll cally
and r<>lably activated more by Images
I I - . I1 I I
of places than by other stimuli.
, •• 11 · ·· · ··· ·- ·
FIGURE 7.17 Attentional S919ctlon. Rrst, attend to the r9d itQffiSand n oticQ the
rough oval that they form. Next, w ithout moW'g your eyes, switch atterrtion t o the
blue Items or thQ horizontal Items. Diffenmt aspects of the pcture appear more proml·
nent as you shift the property that you setect to attend to.
(•) (b) to alter what you see, even as the stimulus remains physically unchanged .
ln Figure 7 .18, you can see attention change the apparent brightness a nd / or
color of a stimulus (Tse, 2005).
Attentional selection can also activate parts of the brain that are specialized
for the processing for one o r another type of stimulus. Recall from Chapter
4 that fMRI has shown tha t different p arts of the human brain are especially
important in the p rocessing of faces (the fusnorm lace area [FFA]; Kan wisher,
McDermott, and C hun, 1997) a nd places (the parahlppocampal place area
[PPA]; Epstein et al., 1999) (Figure 7.19). If s ubjects view an image of a fa ce
s uperimposed on an image of a house ( Figure 7 .20), the FFA becomes mo.re ac-
x
Fusiform face area Parahippcc.unpal place aN:a
FIGURE7.16 Attentional modula-

tion of appearance. '8) Lock at the X
and notice that when you attend to
one rectangle or the oth91', that one
appears Ip} Here, if you attend
t o the pink r9Ctangle, the owrlapping
area looks pink. It changeis to blue If
you attend to the tfoe rQCf:anglB. (Aft91"
Tse, 2005.)
FIGURE 7.19 F\Jnctloni> MRI roveals that different plee<>s

o f the cortex are activated by faces and by places. This is true
QV9n wi"l9n both s timuli are present at the same tlme (as in
Figure 7.20) and the obS9fV91' roorety attGrS his mrantal set from
facas to places. (M Rls oourtesy of Nancy Kan'if\risher .}

AITENTlON AND SCENE PERCEPTION 217
FIGURE 7.20 ThQ:s.Q imag9s com ·

b lne faces and houses. In both cases
you c an use attentio n to enhance thie
PQrception of on9 or the o thg. As ycu
s'Witch your attention f rom one type of
object to th9 o ther , you also c hange
the activation of different p arts of the
bro.i n . as shown in Figure 7.1 9. (Left
image from Downing, Liu, and K.an-
w Bher, 2001 .)
tive when the s ubject is attendin g to the face, a nd the PPA becom es more active
w hen the s ubject is atten din g to the ho use (O'Craven a nd Kan wisher, 2000).
FURlH ER DISCUSSION of FFA and PPA can be found in Chapter 4 on

page 11 3.
Attention Could Coordinate the Acti vity of Different Brain Areas

How might the brain irnplement a solution to the binding problem'? Morespe-
a Appa
if di ffe:l'ent brn in areas perform different tasks li ke processing f.:ices
or places or color or orien ta tion, how rnigh t those
areas be coordi nated if, fo r instan ce, we wan t to
think about h ow the \Vind ows and door of some
red house look like n-..·o eyes and a m outh? On e
possibility is tha t th is bind in g a nd coordiMtion
o f areas synchronizing the tempor,'\l pa t-
terns of dctivity in those are<1s (Singer, 1999). One
role of atten tion ap pears to be to con trol w hat is
synchronized with wh at. For instance, Baldauf and
Desi m on e (201 4) s howed tha t attention to faces
or places could chan ge w hether FFA or PPA we.re
sy nd u onized wi th a third brain a rea.
''"''....
Attention and Single Cells
We have been talkin g ab out areas of the brai n.
O n a finer sc<Jle, v•e c-an ask how a ttenti on n1ight
chan ge the resp onses o f a sin gle ne u ron. Figure
7 2 1 shows one set of possibilities. Supp05e a cell
respon ded to a range of different orientati ons but
was maxi m ally resp ons ive to vertical lines (see
C hapter 3). A tte nti on m ight make the cell m ore
res ponsive across the boar d (Figure 7.21n). This
I I _
u&_
(c)
FIGURE 7.21 Thre9 ways that thQ resp onse o f a 001

coukl be changed as a rEO\Sult o f attention: (a) enhance·
m.ant, (b) sharper tuni ng, (c) alterecl tuning. In each
L1l_ Aftm dhmmg
row. the red linei shows the respon SQ of a single ooll to

tines of d iffertint or1i8f"ltatlons, w ithout attention (left ool-
umn) and with attention (right column).

218 CHAPTER 7
FIGURE 7.22. Each o f thQS.Q images (a) (b)

is a map of the respons-
es of one c i;:ill. In this QXPQ"irnQ'lt, the
lnwstigatCfs tlashed little spots of light
at differ&nt locations and measured th9
ro91SPonses o f the cells.. Brighteir, whiter
locations are areas of greater resp:mse.
The only b9'twQQO (a) and (b)
is that in (a) the mookay was attending
t o the bcation marked by the diamond,
white in (b) it attended to the location
o f the cirde. This c hange in attention
caLJSQd the reCQptrve fiekj o f the CQll to
shrink-wrap around the attended
s timulus location. (From Womelsdorf et
al .. 2006.)
.sort of response enhancement has been seen in visual cortex: cells, but simple
response enhancement seems like a rnther d1ange. Alterna-
tive ly, a cell might becom e mo re p recisely hmed. In the example in Figure
7.21b, sharper tuning would mean that a tte ntion could make it easier for the
n euron to find a weak \o-ertical s.ignal am.id the noise of o the r orientations (as
.seems to happen \·vhe n we a ttend to the h orizontal bars in. Figure 7.17). Lu
and Dosher (1998) carried o ut e legant psychophysical experiments showing
tha t a ttenti on does m a ke it possible to exclud e n oise in this way, thou gh som e
efforts to fmd physiologica l evidence for sharper ttming have fo und only re-
sponse enhancem ent (Tre ue and Trujillo, 1999).
A m ore ra dical p ossibility is thM attention changes the prefe:rences of a
n euron. As illu strated in Figure 7.21c, a cell tha t was initially tuned to vertica l
lines might come to respond better to a different orientation under the influence
of a ttention . The best eviden ce for a d1ange in the fund am ental preferen ces
of a n eu ron comes from studies of its preferences in s pace-that is, the size
and shape o fa neuron's recepfrve field and Desimone, 1985). Fig ure
7.22 s h ows just such a cha nge in a specific of mo nkey cortex. When the
monkey a ttends to one s timulus, the cell 's sensitivi ty is enhanced around tha t
location . \'Vhen attention is shifted to another location, the receptive field shifts
too (Womelsdorf et al., 2006).
If cells are restricting their processing to the object of attention, then sen si-
tiv ity to neighboring items might be reduced as resources a re wi thdravvn from
the m . This prediction is bom e out: mapping the e ffects of a ttention reveals
inhibit.ion sturounding the object of attention (M o unts, 2000).
TI1e three hypotheses about the rnicrodrcuitry of a ttention d iscussed he re
are not mutually exclusive. Different neurons p e rform different parts of the
response ertlancement An effect task of attending, and each s ubta s k m:Jy requi re a different type of change
of attention en the response of a neu-
in ne uronal responses. As a hypothe tical exa n1ple, shnple response e nh.'\nce-
ron In 1'1111ch tlie rieuron respxidlng to
an attended stimulus gtves a bigger rnent frorn o ne neuron m.ight be the 1'conunand " that cau ses a nother ne u ron
res1x:nse. to s harpen its tuning and / or to c hange the shape of its rece ptive fie ld . In
sharper tuning An effect of atten- fact, these may n ot even be different functions, say John Reynolds and David
tion on tt1e response of a neuron In Heeger (2009). Reyn old s a nd H eeger 's 11 no rma lization " theory says tha t the
wl1ich th3 neurai respondl ng to an current response of a ne uron is the product of that ne uron's built-in receptive
attended stimulus resfX)nds more field and the effects of attention. This p roduct mus t then be ''n ormalized # by
precisely. For example. a neuron that neural s uppression . Otherwise the numbers or the level of excitation of the
respcnds to lines with ortentatlons
fraTI - 20 degreoo to +20 degroos cell) can get too big. The results of this single pr sof multiplying and divid-
might ocme to respond to ±10 degree 4 ing c1 n loo k like response enhancement, like sharper tuning, or like ch anges
lines. in receptive-field size and s hape--all d epending on the specific stimuli tha t

FIGURE 7.23 Five "slices" through

are used in the experiment. For present purposes, the main message is th at the brain of a neglect (magntetk::
resonance Imaging view-00 as though
attention can change the activity of s ingle cells. from above). The damag9. shown her9
in indudes light palietol and
fronh\I lobes.. Th9 patient neglects the
Disorders of Visual Attention left side of space. (Image courtesy of
What wou ld h appen if you could n o lon ger prty attention? A comple te in- Ly nn Rcber1son and Krista Schendtll.)
ability to attend would be d evastating. Because we cannot recognize o bjects
o r find \vhat we're looking for witliout atten tion, comple te inattention would
be som.ething near to fun c tional blindness. Brain damage that produces a
defici t this severe is very rare. -f\.1ore common is the attentional equivalent of
a defect. As you may recall, a person w ho is unfortunate enough
to lose primary visual cortex in the right hemisphere will be blind on the op-
posite (left) sid e of visual space. Suppose, h owever, tha t the lesion is in the
right parietal lobe. Patients \vi th this sort of lesion (Figure 7 .23) are not blind
on th eir left sid e but have problems directing a ttention to objects and pJaces
there. These problen1s manifest them selves in a cu rious set of clinical symp-
visual- fteld defect Aportlcn o f
tom s, including neglect and exti11ct.ion. tha visual field with ro vlslm or 1..;th
abnormal vision. typlC8Jly resulting l'om
Neglect damage to the visual nervcus system.
Neglect patients behave as if part of the world were not th ere. Asked to de- parietal lobe In each oerebral hemi-
scribe w hat he is seeing, a patient exp eriencing neglect o f the left vis ual field sphere. a lobe that lies toward the tcp
will tend to name objects to the right of fixa ti on and ign ore objects on the left of the brain bet1-veen the frontal and
A " line ca ncellation test" isa m ore sys te matic way to assess this proble m. 1l-1e ooc/pltal lobes.
patient is given a piece o f paper full of lin es and asked to draw an intersect- neglect As a neurdo;ilC8J S)rnptcm ,
ing lin e thro ugh ench one. A neglect patient might produce something like In vll'•.Jal atteritbn: (1I T11e Inability to
the drawing in Figure 7 the lines on one sid e (th e right, in this case) are attend or respond to stlmuH In the
contraleslonal visual field (typically, the
cros.sed out, but those on the other side (here, the left ) are overlooked. Figure lett neld attar nght parietal damage) . (2)
7 25 s hows what rnight result if a neglect patient, again experiencing neglect lgrl:f/ng half of the boct1 or half of an
of the left visua l field, were <-11sked to copy a picture. Again, the right side is cbJoot.
FIG URE 7.24 A neglect patient might produc9 this sort

of result if re..k.00 to c ross out all the lines. The patient might
neiglect the lines in the left side of the image, following damage
to the right par1"1tal lob9.

220 CHAPTER 7
(b)
D D 0
D D oCJ
FIGURE 7.25 In copying a drawing like the one in (B}, a neglect patient often omits
one side of the obj€1ct, as in (b).
reproduced fairly faitllfu lly, but the left side is missing. Such a neglect patient
might also eat only what's on the right side of his di.imer plate or shave only
the right side of his face.
It is not always clear whether neglect affects one side of the visual world
or one side of objects. Mostly likely, neglect can be either or both. In one
elegant illus!Tation of this point, Steve Tipper and Marlene Behrmann (1996)
asked neglect patients to detect changes on one or the other side of a barbell.
When the barbell had one ball in the left Held and one in the right, patients
paid better attention to the right. Upper and Behrmann's clever trick was to
rotate the barbell, while the patient was watching. so that the left ball moved
into the right field and the right ball moved into the left (Figure 7 .26). The
patients paid better attention than they should have to the piece of the object
in the normally neglected, contraleslonal neld (the field on the side opposite
the lesion). Apparently they were neglecting half of tlte object, and the neglect
somehow moved ¥.ith the object.
Extinction
The phenomenon of extinction is related to neglect; it might even be neglect
in a milder form (Driver, 1998). A neurologist might test for neglect by having
a patient fixate on her (the neurologist's) nose. She might then hold up a fork
in the patient's good (in this case, right) visual field and ask, "What do you
see?" "A fork," would come the reply. Next the neurologist might present a
spoon in the bad Qeft) field. A patienr-rth left-visual-Held neglect would be
nonplussed, but an extinction patient would be able to report the presence of
cootraleslonal ne1d The Visual fleld

ai the side owoslte a brain leslon. For
example, points to the left of fixation
are oontraleslonal to damage In the
nght hemisphere of the brain.
extinction In Visual attention. the
lnablllty to perceive a stimulus to one
side of the po'1t of fixation (e.g .. to the
right) In the presence of another stlmu- FIGURE 7 .26 Tipper and Behrmann's (1996) experiment. step 1: The patient
<is. twk:8Jly In a comparable position neglgcted the left side of the barbell. Step 2: The barbeU was rotated through 180
In the other Visual field (e.g., on the left degrees. Step 3: The neglect rotated with th9 object, indlcatlng that, In this case, thg
side). neglect was r91atlw to the object, not to th9 whole SCQne.

the spoon. If the ne urologis t were to hold up a fork in one ha nd and a spoon ipsileslonal field The "1sual field on
in the other, however, the extinction patient would report only the objec t in the sarne side as a brain leslon.
the good fie ld . The other object would be pe rcep tually "extinguished ." attention deficit
In neglect, the patie nt may be entirely unable to deploy attention to the disorder (ADHD) a quite common
cont r.-tlesional s ide of the world o r the object. In extinction, the patient may be childhood disorders that can oontlnue
able to dep loy attention to an object in the contralesional field if it is the only Into adulthood. Symptoms Include
d ifficulty focusing attention as well as
sa lient object. However, if the re is competition fr01n another similar object in problems controlling behavior.
the lpslleslonal fteld (the same s ide as the lesion), then the hvo objects com pete
for attention and the ips ilesional object \•; ins. Objects are always compe ting
for our attention, but as a genera l mle it is no 1na t ter for us to redeploy
attention from one obje ct to the next. In extinction, the object tha t loses the
com.petitio n is far less likely to attract atte ntion than. is the c.ase norma lly. To
leam w hat happens w hen the parietal lobe is dam aged on both sides of the
brain, see Web Essay 7.1: Balint Syndrome.
Selective Attention and Attention Deficit

Hyperactivity Disorder (ADHD)
Neglect and extinction are relattvety rare consequences of damage to
the nervcus system. There are much more common disorders of at-
tention. Attention deficit hyperactivity disorder (ADHDJ is the best
known of these. Recall that . ear1y on in this chapter. we said that at-
tention was not a single thing but a name for a famHy of processes that
restric t o r bias our mental activities. lhat Is d emonstrably true, but at
the same time , we have the Intuition that there Is some general atten-
tional resource that is being summoned when our mother or our teacher
or our significant other says, MPay attention!" Some seem to
hwe m ore d ifficulty ' paying attention" in this sense than others . If that
difficulty Is severe eno ugh, they may be diagnosed with ADHD , w hich is
diagnosed in North American ct1ildren more than any other behavioral
disorder. It is characterized by three kind s of symptom s: impulslvlty (an
Inability to contro l behavlot1. hyperactMty, and of most interest to us, in-
attentiveness. An Inattentive child or adult might fail to carry out a set of
instructi ons or tasks Oi ke homework). He might have problems organiz-
ing his activities and be easily di stracted or forgetful. He migllt seem not
to li sten w hen spoken to (Feldman and Rein, 2014; Volkow and SWan-
son. 2013).
How does the sort of attention that is disrupted in ADHD relate to
the attentional processes that 'Ne have been discussing here? For in-
stance, if you have trouble organizing a task. might you have trouble o r-
ganizing a visual search? Mullane and Klein (20C8) analyzed the results
of a number of different studies of this question. They found that there
·were differences between cbsetver.s \.vlth and "vithout a diagnosis of
ADHD but that these differences were fairly subtle . Doing search tasks
like those sh::>w n in Figure 7 .6, :ADHD 0 bseivers prpduced a R"ttern of
results that w as qualitatr;ely simil ar to t he pattern shown at the bottom
of Figure 7 .6 . Feature soorches were lig,!y entclent; this was basically
the same in AOHO and non-ADI-ID ct>servers. Inefficient searches were
a little mo re Inefficient In ADHD observers , but the difference was not
dramatic . ADHD and non-ADHD observers seem to search In pretty
muc h the same manner. ADH D Is a disorder wft h significant attentional
comporents, but the major components of ADHD seem to tie o utside of
the realm of visual spatial attention.

222 CHAPTER 7
bottlMe<:k
FIGURE 7.27 Tl/llO pathways fro m the world to our perception o f the vvorld. (a)
Early stages of the visual system feed two pathways: Ip) a s.eltJctive patlT-Nay, with a
bottl9n9Ck govi:orn9d by si94Qciiw attQfltlon, and (c) a nonS9lect tve pathway that Is no t
subject to this attentional bottlenQCk. The selective pathway permits the recognition of
one or a vgy few objraicts at one time. The nonsraloctive pathway contributes inform a·
tion about the distribution of features across the SC919 , as wc:-11 as informatlro about
the rough gist ofth.e scene. (Aftel'Wcifei et al., 2011 .)
Perceiving and Understanding Scenes

Two Pathways to Scene Perception
This chap ter has devo ted considerable space to the argumen t that you need
to attend to specific objects if you ore going to fully recognize them. However,
we certainly seem to be able to "see" more one object at n time. More-
over, we can use information about the whole scene to guide our attention to
a par ticular object in that scene, s uch as the faucet in Figure 7.11. Guidance
by scene properties raises a question: How d o we know tha t we're lookin.g at
a kitchen before we find the faucet or the sink?
As a framework to think abo ut this and about the percep ti on o f scenes,
we w ill use a two-path way approach tha t is somewh a t similar to the w1m t
and where pathways o f Chaprer 4 (see Figme 4.2) (Wolfe e t aL, 2011). Here,
the path ways are labeled "selective" and "nonselec tive." Early vision (Figure
7.27aJ processes bask feahue in.formation a nd uses grouping and other ntles
(C hapter 4) to create w ha t we might call "protcrobjects" (Ren sink, 2000) tl1a t
ca n be selected a nd recognized in the selective pathway (figure 7.27b). The
selective pathway has been the implicit subject of the chapter to this poin t. Now
le r's consid er w h.."l t can be d one by the nonselecti ve pathway (Figure 7.27c).
The Nonse/ective Pathway Computes Ensemble Statistics

Look at Figure 7 .28. You very rapidly kn mv several things about this sch ool
of fish. You know that the sch ool is oriented to the left, perhaps s lightly dmvn
from hori zonta l. You know the average size and color of the fish. You also know
something about the distri buti on of oden tations? size:s, and colo rs. That is,,
you know that a range of fish orientations is present. They are not all pointed

AITENTJON AND SCENE PERCEPTION 223
FIGURE 7.28 Some "fishy' "insernble statis tbs. In a glance you can tell that these
fsh ar9 oriented roughty horizontally as a group , though you cannot t911 , in that
glance. IM"lich specific fish are oriented exactly horizontally. Similarly, you know In
a glanCQ tho.t some fish ar.e more tilted and som e are more blu9 in color. S91ective
attention Is requiri:id to determine 'Whether ony one fish is both strongly tilted and
bluish.
in exactly the same direction. To know these things, you are computing en-
semble statistics (Alvarez, 2011). Si!!."a.tistics rep rese nt knowledge
about the properties o f a group (ensemb1e) b f obj ts r, perhap ve s ho uld
say, an e nsemble of proto-objects. Om estimates of these statistics are s urpris-
ing ly ac-curate (Cho ng and Treis m.in, 2003; Chubb and Landy, l()(J;I).
FURTHER DISCUSSION of our ability to perc8ve the ch<racteristics of a

group of objects can be found in Chapter- 4 on page 102 (Figure 4.17),
11'1ese are ensemble s tatistics beca use we knmv thern \vithout knmving the
properties of the individual fis h. For example, you would have to search to find
a fish that was pointed exactly horizontally, even though that fish's orientation
contributes to the es timate of the ensemble 01ientation. Features are n ot bound
in ensemble .sta tistics. You wou ld n o idea, ·with out attenti on al scrutiny,
whether any of the .small fish were bluish fis h. You know there are s ma ller fish
and bluer fish, but appreciating conjunctions requires the selective pathway.
The Nonselective Pathway Computes Scene Gist

and Layout - Vety Quickly
As '"as mentioned earlier in our discussio n of attending in time, people can
monitor a s tream of images for one particular image or type of image at rates
of one image every 100 ms (see Figure 7.14)_In foct, i n som e cases you d on' t
need as much as 100 ms. Holle Kirchner and Simon Thorpe (2006) presented
ensemble statistics The average
hvo si d e- by-sid e scenes to observers and fo tmd tha t the observers could and dlstrlbutlrn o f properties like orien-
reliably move their eyes to fixate whichever scen e include d a nimal (any tation or color over a set of ctijects or
animal) in as little as 120 ms. The fact tha t m ost o t that 120 ms was taken up over a region In a scene.

224 CHAPTER 7
336 Arnrna Appa
FIGURE 7.29 A simple visual
composed of t\o\lo sine
wave gratir)gs.
in generatin g the eye movement implies that the processing of the scene \.•:as
really fast. Michelle Green e and A udeOliva (2009) presented single scenes very
fas t, followed by m asking s timuH. They fo und that people could differenti-
ate between natural and nrban scenes with just 19 ms of e.xposu re to a scene.
Global properties th.."1t described the spatial layout of the scen e {e.g., was it
a navigable or non-rui vigable space?) could be extracted a bit m ore quickly
than could basic categories like "desert" or nlake/' but all of these could be
appreciated w ithin about 50 ms. It probably takes 20-50 ms to recognize just
on e object, so observers are certain.I y not recognizing scenes by recognizing a
strin g of objects and putting the m together. \.\•11at are they d oing?
An interesting potentiaJ answer comes from the work of Aude O liva and
Antonio Torra lba (2001). Consider the very boring "scene'' in Hgure 7.29. It is
com posed of a vertica lly oriented, low-spatial-frequency component (a b rood,
upright sine wave) combined with a diagonally oriented, high -spatial-frequency
component (a narrmver sine that's tilted to the right). As we learned in
C hapter 3, any image can be broken down like this into its spa tial-frequency
components, and in fact, the visual system to process images in exactly
this via a set of spa tial frequency-hmed cha nnels.
If we further analyze the spa tial-frequency components of scenes, we find
tha t certain patterns of frequencies cor respond to C'eftain types of scenes. To
take a fairly obvious example, wide-open scenes (beaches, prairies, pa rking
lots, and so on ) tend to h ave a lot of strong horizontal comp onents (corre-
sponding to the h orizon). Oliva and Torralba railed this dimension" openn ess''
and d efined a number of other dimensions, including 11 naturalness" (versus
"m an-mad e'' ) and "rou ghness." l11ey could m easure the a mount of each of
these dim ensions by examining the sine waves that made up the stimulus.
\ Vhen Oliva and Torralba measured the properties of thousa nds of scenes,
they observed something very useful : scenes with the sa me mea ning tended
to be neighbors in the space d efined by their scene dimensions. Suppose we
d efine a si mple two-dimensiona l space us ing the dimensions of openness
and expansion . Figure 7 .30 gr.lphs a set of man-m ade scenes using these h-'lo
dimensions. Look at how the scenes organize themselves. The fronta l vie\vs
o f buildings fal l in the uppe r right (low open ness, low expansion). Highways
fall in the lower left QUgh openness, high expa nsion ). The rules tha t m ake
spaUal layout The doocn pt Ion of the th is nrrangement d on 't include anything about buildings or high ways, just
structure of a scene (e.g., enclosed,
qien. rough. smootti without refer- sin e waves. Neverthe less, scenes M th the same gist tend to clus ter togeth er.
ence to the Identity of specific objects Remember that these dimensions are defined on the basis of a spatial-frequency
In the scene. analysis of the entire scene. To gain an lmderstanding of the scenes, would

ATTENTlON AND SC ENE PERCEPTl ON 225
0.5
•fl-
"'
--05
-1
'5
!,
ct 5
c..
[
1i
:;: -2
-2 - 1.5 -1 -p.s 0.5

Mo re open Less open
Openness
FIG URE 7.30 Spatial layout from global ink>rm:itlon. In this coll9ction o f urban
scenQ:S, th9 x -axrs rep rQSQnts a measure of •open n9Ss, • from panoramic SCQnes to
vertically structur9d scenes. deriVQd from spatial tequency infoonation. T he y-axis
represents expansion (captu ring a SMse of d9pth from pgrsp9Ctlw). SC9MS vvlth the
satnQ mi9afling ciust9r togeth91'" in this sp ac9. {Courtesy o f Aud9 Oliva)
be no need fo r us to shift ou r gaze or even our a ttentional focus, no need to

b ind features o r recognize objects, n o n eed even for p roto"bjects.
Oliva and Torrnlba's work shows how we might achieve a basic tmders tand-
ing of the meaning of a scene very quickly by c:u utiyzing the s pa tial-freq uency
components of the image. This a nalysis migh t te ll us th..'lt w h at we're looking
at might be a highway, a building. or a farmy ard. The selective path way might
contribute the identities and rela tive loca tions of a few objects. TI1en we would
have a scen e descripti on a long the lines of "a farmyard w ith this ch icken in
this p ose n e ar this cow and this tractor." This combination of globa l and local
info rma tion might create a r id1 enou gh to specify a pa rticular
scene and to gi ve us the impression of seei ng a wh ole scen e.

226 CHAPTER 7
FIG URE 7.31 Spsnd a second or

two looking at 83.ch of thQSQ p ic tures.
Then m ove on to Agure 7.32. Memory for Objects and Scenes is Amazingly Gooo
The representation of scenes by information 6"om the selective and
no nselective pathways is very powerfol. First, it gives r ise to our perception
of a world full of cohe rent o bjects in a co herent scene, even before we have a
chance to attend to m ost of those objects. Second, it is easily stored in m em ory.
Have a look at Figure 7.31. It cont:Jins photographs of 16scenes. Spend about

o ne second o n ead1 scene, and then flip the page to view Figure 7.32_ TI1ere you
will find anoth e r set o f 16 scenes. Your jo b is to d e termine w hich 8 of the 16 in
Figure 7.32 you saw in Figure 7.31. Yml probably won't n eed the answer key
(i t's in the figure) . [t is likely that you will correctly classify 14 or more of the
im.ages as "old" or 11new." 1hat's pretty quick learning of fairly complicated
stimuli. Yet 1t is a mere fraction of what you could do if you had the tinle and
we had the pictures. Roger Shepard (1967) conducted the original version
of the test 'Vlith 612 pictures and found that his observers were 98% correct.
They were 90°/o correct \V hen quizzed a '"'·ee k later. Standing, Conezio, and
Haber (1970) got 85% accuracy after s howing their observers 2500 pictures
for 2 seconds apiece, a nd Standing (1973) later obtained similar results with
l0,000(!) images.
In his s tud ies, Standing used a diverse collection of scenes dipped from
magazines o f the time. Consi der what might happen if we res tricte d the
''scenes " to single objects (a n apple, a backpack), and what might happen
if '"·e included o the r examples of these objects in the test set (did I see that
app le?) or, worse yet, the sam e objects in diffe rent poses (\\"as that backpack
lying d own before?). Aude Oliva and her s tude nts (Brady et aL, 2008) tri ed
this \•1.rith 2500 objects. Whe n as ked to pick betvveen an old item and a ne\v
one, their observers got 92o/o correct if the n ew ite m \Vas an object of a type
that had not been seen before in the experiment. Amazingly, observers were
still 88% correct at remembering whidl e.xample of a catego ry they had seen
and 87% correct at rememhering which s tate of a specific object they had seen .
In our di scu ssion of a ttentional blink earlier in this chapter, we mentioned
another set o f e xperiments with pictures, conducted by Molly Potte r .:md he r
colleagues. They presented pictures of scenes very rapidly and asked observers
to say whethe r, for example, a pimic scene was presentin the s trea m o f im ages
(Potter, 1975, 1976). Once again, ob_servers were remarkably good at this task.
Helene lntr,nib (1981) took the parndigm a s tep further and showed that people
could m onitor a stream of stimuli for the one thc1t Wl1s not an anima l a t rates
of e ight images per second. Th.:1 t mean s th..'l t every image in the strea m could
be classified as "an_imal" or "not animal " after a mere 125 m s of e>.'Posure. By
the wa)'i at that rote we cannot conunit images to mem ory very effectively, but
we do have a pretty clear idea o f what we 're seeing as it flas hes by.
But, Memory for Objects and Scenes Can Be Amazingly Bad:

Change Blindness
From the findings just described, we could conclude that pktmes can be tm-
derstood very rapidly and that, given enough time-perhaps a seocond-\.ve
can code the m into memory in sufficient d etail to be able to recognize them
days later. But d on' t endorse tha t conclusion too quickly. In the 1990s, Ron
Ren.sink and his colleagues introduced a diffe rent sort of picture rne mory ex-
periment (Rensink, O'Regan, and C lark, 1997). 1l1ey just
one picture at a time. The observers would look a t the pic ture tor a while, and ppa
then it \'Vould vanish for 80 ms and would be replaced by a si milar irn.age.
The observer 1s task was to determine what had changed. The two versions
of the same image would flip back and forth, separated by a brief interva l in
which a blank screen was presented, until the observer spotted the change
or time ran o ut. A s tatic version of the task is illustrated in Figure 7.33. Try to
find the differences be t\.veen the two pictures in this figure. TI1e answers a re
in Figure 7 .34. You can also try detecting chan ges in some flicker movies in
Web Activity 7.5: Change Blindness.
It may have taken y·ou a while to discover all four changes in Figure 7.33.
That's what happened in Rens ink's experiment:: participants took several seconds

228 CHAPTER 7
FIGURE 7. 32 \l\lithout refQtring back to Figure 7.3 1, tty t o identlff \Nhic h o f these
pictur9s you afrQady saw. Decide wh9ther 9ach picture is or "new.• You c an
check your ane.wers on the grid to the left. n1e shaded squarf!iS show the locations of
ok:t p ictures.
8230336 Arlma Appa

ATTENTlON AND SC ENE PERCEPTl ON 229
FIGURE 7.33 There are four d ifferenoes between thase two lma;;1es. Can you find
them ?
on aver age to find the changes, and some never num aged the task at a ll in the
time they were given. This phenomenon is known as change blindness. Jf
the blank screen between the two itnages is rem oved, the cha nges are obvious
because observers experience a kind of apparent-motion effect (see Chapter
8) w h en a n object d um ges position, d isap pears, or changes color. But on ce
we1 ve h'tken care o f tha t lo w - level facto r by inserting the blank screen , view ers
ca n be quite obliv ious to jet engin es tha t vanish from airplan es, b ra nches tha t
jump from tree to tree, and boats tha t cha n ge co lor.
Actua lly, we can elimin a te the need for a bla n k screen if the cha nge is ma d e
w hi le the eyes are m oving. We are virtuall y blind during a n abrupt (saccadic)
eye movement McConkie and Currie (1996) took ad vanh.1ge of this fact to m ake
dramatic ch an ges in a scene while an observer was jus t looking around . If they
made the chan ges w hile the eye was in moti on, observers often foiled to notice.
FURTI-IER DISCUSSION of ete movements can be found in Chapter 8

starting on pa;Je 250.
It is not immed ia te ly obvious how these various facts can all be true of the
sam e visual system. This is a topic of current controversy in. the field. How
can we remember thous.an ds of objects after a mere second or two of v iew-
ing? And if we can d o that, h ow can we fail to n otice s ubstantive change in a
picture that is right in front of us? Does it m ake any sense that we remeinber
the p ich.ue but seem oblivious to important details?
What Do We Actually See?

In the 1990s, Dan Simons and Dan Levin (1997) conducted a wei rd experiment
on the s h·eets of Ithaca, New York. One experim enter asked an uns us pecting
passierby fo r d irec tions. \'\fhile the passerby was g iving those directions, a couple change blindness The failure to
notice a change betvJeen Thvo scenes.
of othe r p eople carried a door d own the street and bet\.veen the p asserby and If the gist. or meaning , of the scene Is
the experimenter. While the d oor was m O\.;ng by, the experi menter ducked not altered , quite large chan:Jes can
d own, s neaked away, and was replaced by a second experimen ter. 1l1en the pass unnoticed.

230 CHAPTER 7
FIG URE 7.34 HQre are the locations

o f the differences between the two
irnageis in Rgure 7.33. \NhQn you go
back to figure 7.33 with this informa-
tion. you will hav9 no troubl9 s99ing
what was alttfOO.
d oor was gone d own the street, and the critkaJ ques tion '"'hethe r the pass-
erby wou ld return to giving instn1ctions. In many c,,15es, the passerby did just
tha t, OlS if the d1ange in experimenter had gone u1mohced . How could that be?
Perhaps this change was undetected because it didn't ch ange the gist of the
scene. The passerby might think, ''l' m giving ins tructions toa guy. Now there
is this weird d oor. Now I'rn giving ins tn1ctions to d gu y again," and simply
n ever register that the gl1y chan ged. See 11 The Colour Changin g Card Trick''
in Web Activity 7.5: Change Blindness for anoth er exa mple.
To make this into a more con v inci ng theory, it would help H we really
unders tood w ha t the o f a scene nlight be. Gist is clearly m ore tha n the
brief verba l d escripti on \Ve ml ght give if asked to "d escribe the gist. " Look
at Figure 7 .35. Now look a t Figure 7 .36 and decided w hich one of these two
photographs is the same as Figure 7.35. You ca n p robably d o this quite eas ily,
even though the gist descriptions of Figures 7..36a and b \Vould be very similar: a
skier on a fairly s teep slope on a s unny d ay \vi th mountains in the backgrotmd.
TI1ere is some thing a.bou t 11gist" that we cannot trivia lly put into '"1ords .
You can experien ce a much simpler version of c hange blindness in Web
Activity 7.6: The Attentional Bottleneck. Participa nts ob served a collection
of red and green spots. W hile they watch e d, one of the sp ots s uddenly got
brig hte r, a nd on half the trial s the brightened spo t also changed color. T he
ch an ge in brightness inv:ariably a ttracted a t tention to the critical s pot. All the
observers had to do was determine wh e the r the sp ot had changed color w hen
it changed brightness. l11ey performed horribly; tl1ey man aged to get onJy 52%
correct in a task where they couJd get 50% correct just by guessing. It seemed
tha t unless the o bservers happened to be a ttending to the critical spot before
the d1an ge occurred , they had no idea \vhether the color ha d cha nged (\ Volfe,
Re inec ke, and Bra\oVJ.11 2006).
Th is finding is consis tent \vith our change blindness sto ry. The gist of
the disp lay di d n ot cha nge wh e n on e sp ot changed, so the ch an ge was no t
8230 deteeted t'm1ess tha t spot h appened to be attended a t the time of the d1ange.
At any moment, then, we are percei\o;ng the gist of the scene, even if we can 't
qui te verbaliz.e w ha t that gist might be; and we a re perceiving the currently
attended, bound, a nd recognized products of the seledive path way. The ad -

FIG URE 7.35 Look at this pciure for a couple of seconds. Describe it to yourself in
a sentence. Then go find Agure 7 .36.
va ntage of this very simple display is th at we can estimate how many objects
""'ere attend ed at the mornent of the d1e1 nge. TI1e estimate is in the range of
:zero to fo ur items, indicatin g, again, that selective attention selects very few
objects at any on e time.
The problem with this view is that we tlrink \¥e see so much more. As we
look around the world, we experience a full-color, in-focus, three-.dimensional
scene fill ed with dearly identifiable objects. \ftle d o n ot h ave a pe rcep tua l
expe rience of zero to four ob}ects, fl oatin g in a sea o f ensemble \Ve
know, from the research described in other chapters, that this experience mus t
be a constructi on of the m ind, not an exact copy of the s timulus. ·we know
from Chap ter 2 that the world is decently focused only at the fovea and that
high -resol ution processing occurs only in the central 1 degree or so of the visual
field_We wi ll see in 8 that the scene before us is s m eared across the
retina three or fo ur times ead\ second as we m ove our eyes. We know from
C hapter 6 th at \o\>'e1 re actu a ll y getting h.voslightly different cop ies of the .scene
from our hvo retinas, and that both copi es are rea lly rn.·o-dimensional.
Over and over in this book, we say thata particular aspect of perception is
an inference a guess abo ut the world . Our co1 s:dous experience of the world
is the moth of.::ttll Th i!.)eaJization is hard for us to aa::ept beca use
perception doesn' t/eel like a n inference about wha t is. It feels like it must be
sho'"ring us what's really out there. It feels like Truth. Pheno mena like change
b lindness are important beca use they s how us the gap between percepti on
and rea li ty. Experiments investigating ch ange blindness s how t LS that we d on ' t
see all of wha t is there. Rather, in some spots, we see \vhnt wns there wh en we
last paid attention, and even then we may not remembe r it perfectl y (lrwin 1
Zacks, a nd Brown, 1990). In othe r s pots \·Ve perceive w h at we think sh o uld
be there {see Web Essay 7.2: Boundary Extension). Moreover, we may n ot
perceive what we d o n ot expect to see--whn t d oes not fit with the current
inference. This fail m e to notice the unexpected lead s to the phenomenon of
inattentional blindness. TI1e great example of inattentional blindness comes
inattentional blindness A failure to
from a n other experime nt of Da n Sim ons, this time in colla boration w ith not be -cr at least to report-a stimu-
C hristopher 01abris. Th ey had observers keeping track of ba ll m ovements in lus that would be easily reportable If it
a basketball passin g gam e. Many of these subjects failed to no tice an actor inn 'Were atterded.

232 CHAPTER 7
(a) (b)
FIG URE 7.36 Which of thes9 two lrnag9s is the on9 you saw in Figur& 7.35? You
probably know, even tt1ough your wrbal d'9SCt1ptlon o f th9 new Image would be very
similar to the d95crlptbn you gave for Figure 7 .36. ApparQfltly, the gist o f a Sc.QOQ
contains more than Is captured by our usual verbal descriptions.
gorilla into the middle o f the scene, waved, and wandered

off (Simons and Chabris, 1999).
Outside the lab, the bottleneck be tween the world and. our perception is
not u su ally mud1 of a problem, because the world is a pretty stable pre-
dictable place-at leas t o n a m o m en t-to-mo m e nt basis. U you put your coffee
mug down on the desk and turn your '1ttention to the computer screen, the
mug will be there w hen you choose to attend to it again. Only in the lab does
the coffee mug vanish during an eye m ovement. This means tha t the physical
\\:orld can serve as a n 11externa.1 memoryu tha tb.:lcks up the perceptual world
we create in our m inds (01Regan, 1992).
\ Vhen the wor ld does cha nge, it usually changes in very predic table ways.
U we' ve ju st driven past Second Avenue a nd then Third Avenue, we probably
d on ' t need to look a t the next street sign to know that it says "Fourth.'' At a
rate of about 20 objects per second, we can moni tor the relevan t ite ms in th is
relatively s table world and be reasonably sure that we are up-to-date. And
if some thing smprising does occur (a gorilla leaps into the middle o f Pourth
Avenue, for exampl e), the event will probably be marked by a visual transient
that grabs our a ttention so that we can update our internal pretty
qttickly and maintain o ur grasp o n reality (Yantis, 1993). (See Web Essay 7.3:
Attentional Capture.)
A certain amount of work needs to be d one in order to induce us to m iss
gorillas. Still, we sh o uld n ot fee l t oo complacent a bout o ur inference of the
8230336 Amma A perceptual world. Con.sid er, for example, tha t eyewitness testimony is based
on the ass umption that you can report w ha t was there, not '"'hat yo u in ferred,
gues.sed, hoped, or feared was there. Sadly, if unsurprisingly, we know that
eyewitness testimony is subject to the sa me sorts of effec ts discussed here a nd
th..1t these factors are p robably the cause of errors with real conseque nces (G.
L. Wells and Olson, 2oro). \Ve need m ore data before the courts and the public
v1,i ll understa nd this problem. As a step in th at directi on, C hnbris and Simons
(2011) have extended their work on real-word inattentional bl indness to show
that people w ho were g iven a task of following a jogger down a track could

AITENllON AND SCENE PERCEPTION 233
eas ily fai l to notice a very visi ble apparent bea ting occurring along their ro ute.
The researche rs picked this particu la1· scena rio because of a re<ll case in w h ich
a polireman rep orted that1 as h e chased a StlS}-"'>ect,. he neve r saw his colleagues
beating a nother man . People d idn't be lieve tha t he could have been so blind .
Now you kn ow that such a scenario is, at least, p ossible.
Summary
1. Attention is a vital aspec t of perception because w e cannot process al l o f
the input from our senses. Tite term attm tio11 refers to a large set of se lective Refer to the
mechan isms tha t ena ble us to focus on some stimuli at the expense of oth- Sensation and Perception
ers. Though this chapter talked almost exclusively about visua l attention, ComP'!nbn Website
a ttentional mechanisms exist in all sensory domains. sltes.slnauer.com/wolfe4e
2 In vision, it is poss ible to direct a ttention to one location or one object. If for aotMtm. essayo, study
some thing happens at an attended location, we will be fas ter to respond to
it. It ca n be useful to refer to the "'spotlight" o f attention, though deploy- questlrns. am other stu:ty ak:ls.
ments of a tten tion differ in important ways from movements of a spotlight.
3. In visual search tasks, observ ers typica lly look for a target item among a
nu mber of distractor items. If the target is defined by a salient basic feature,
such as its color or orienta tion 1 search is very effi cient and the number of
d.istractors has little influence on the reaction time (the time required to
find the target). It no bask featuJE info rmation can guide the deployment o f
a ttention, then search is ineffi cien t1 as if each item n eeded to be examined.
one after the other. Search ca n be of intermediate efficiency if some fea ture
info rmation is avai lable (e.g., if we're looking for a red car, we don ' t need to
ex:amine the blue objects in the parking lot).
4. Search fo r objects in real scenes is guided by the known features of tlle
objects, by the salien t fe atures in the scenes, and by a variety of scene-based
forms of guidance. For example, if you're looking fo r your can of scxla,. you
t,'/ill guide your a ttention to physically p lausible locations (horizontal surfac-
es) and logically sensib le places (the desk or counter, p robably not the floor).
5. Attention varies over time as well as sp ace. In the a tten tional blink p ara-
digm, ob servers sea rch fo r two items in a rapid s tream of stimuli that
awear a t fixation . Attention to the fits t target makes it hard to find the sec-
ond if the second appears \...-itlUn 200-500 ms of the first. When two iden ti-
ca l items ap pear in the stream of s timuli, a diffe rent phenomenon makes it
hard to de tec t the second instance.
6. The effects of attention manifest themselves in several different \vays in the
brain. Jn .s ome cases, attention is marked by a general increase in ne ural
activi ty or by a greater correlation between activity in d ifferent brain areas.
In other cases, a ttention to a particular a ttribute tunes cells more s harply
for tha t a ttribute. And in still other cases, attention to a s timulus or loca-
tion causes recepti ve fields to shrink so as to exclude una ttended stimuli.
All of these effects might be the result of a s ingle, nnderlying physiological
mechan ism of attention.
7, Da mage to the parietal lobe of the brain prOOuces deficits in visual a ttention.
Damage to the right parie tal lobe can lead to neglect, a d isorder in \vhk h it
is hard to direct attention into the contralesional (in this case, the left) vis ual
field. Neglect of an oli t or half of their own body.
8. Scene perception involves both selective and non__c:,elective processing. Tasks
like visual search make ex tensive use of selective processing. Nonselective
processing allows observers to appreciate the mean and variance of features
across man y objects (or p roto-objects). Tims1 you know the average orienta-
tion of trees in the woods (vertical) before knowing w he ther any par ticular
tree is o riented perfec tly vertically. Using spatial-frequency infom'\ation,
even ·without segmenting the scene into regions and objec ts, the nonselec-
tive can provide informa tion about the na ture of a scene (e.g.,
whether it's na tural or man-mad e).

234 CHAPTER 7
9. Picture memory ex'Periments show tha t people can remember thousan ds of

images after only a second or two of exposu re to each . In contrast, chan ge
blindness experiments show tha t people can miss large changes in scenes if
those changes d o no t markedly alter the meaning of the scene.
10. Our perceptu al experience of scenes consists of nonseJective processing
of the layout and ensemble sta tistics of the scene, combined "'fi th selective
p rocessing of a very few o bj ects at each mom en t. Hmvever, the fi n.._"ll experi-
enc:e is an inference based on a ll of the p receding p rocessing. not merely the
sum of that processing. Usually th.is inference is ad equ ate because we can
rapidly check the world to determine whether the chair, the book, and the
d esk are still there. In the lab, however, we can use phenomena like inatten-
tional blindness and ch ange blindness to reveal the limits of o ur perception,
and it is becomi ng increasing ly clea r tha t those limits can have real-world
consequences.
8230336 Anma Appa

lq
8230336 Amma Appa

CHAPTER 8

Visual M otion Perception
LIKE ANY SELF-RESPECTING LADYBUG, the on e in Figure 8.1 Hits around from
leaf to leaf. At time 1 it is on the lowermost lea f; at time 2 it has moved to the
middle leaf. Otu visual systern d istingui s hes the bug by a numbe r of features,
sud1 as its s hape, its locatio n in s pace, and its color. Previou s chapte rs have
established these features as fundall"1ental perceptual dimensions: character-
istics of visual s tirnuli that are directly encoded by neurons fairly early in the
visual system.
ls the ladybug's m otion als o a fundamental perceptual dimension? At
first glance, we might think not. After all, mo ti on is really just a d1ange in an
o bject's loca tion over time. If we alread y have neural mechanisms set up to
d e te rmine p osition., why go to the ex tra expense o f addin,s more low-level
machinery to process mofi on ?
Tho ugh the added in ves tment rn.ight see m unnecessary, consider this
question from the position of the primordial vertebrates that "inven ted " the
\o;sual syste1n we even tually inherited. Many bugs move pretty quickly, so if
we depend on ca tching them for ou1· supperr fast detection of their d.irection of
m otion will be important to our s urviva l. Similarly, if o ther animals are trying
to catch us for their s upper, we need to be adept at detectin g the m o tion of
these predators too. ln Chapter6 we saw that mo tion is an irnportant
cue for depth perception. And from a m ore modern perspective, it would be
hard to believe that hockey goalies, baseball batters fucing knuckleba ll pitch-
ers, and boxers trying to avoid be ing pummeled by their oppo nents could
d o their jobs without ha ving a m ec hanis m for very quickly d etermining the
m ovem en ts of pucks, balls, and fists.
In fact, werve already seen some s tro ng indica tions that rnotion is a low-
level perceptual phenomenon-in Chapter 3, where we learned that many cells
in the primary vis ual cortex selecti vely respond to m otion in one particular
directi on _ A phenomenon ca lled the "v,:aterfall il lusion'' provides ano ther
piece of e viden ce tha t there is som eth ing special about motion. Here' s h ow
Robert Addams (1834) d escribed this illusion after a visit to the waterfall of
Foyers (Figure 8.2):
Having steadfastly looked for a fe\Y seconds at a particular part of the

cascade, admiri ng the confluence and decussation of the currents forming
the liquid drapery of and then suddenly directed my eyes to
the left, to obseive the face of the sombre age-wom rocks inunediately
contiguous to the water-fall, I saw the rocky surface as ii in motion
Upl-v ards, and \vi th an apparent velocity eq ual to that of the descending
water, which the moment before had prepared my eyes to behold that
singular deception.

238 CHAPTER 8
Time"! 1lle r'deception" th...'lt Addan'l.s d escribed-which had also been noted by
Aristotle (384-322 BCE)-was later dubbed the motion altereffect (MAE).
After v.ia,q.ng motion in a cons tant direction for a susta:ir1ed period of time (at
least 15 seconds or so)r \Ve see any sta tionary objects that we view subsequently
(like the rocks around the waterfall) as movin g in the opposite d irection. TI1is
phenomenon may seem a lo t like the color aftereffects \Ve s tudied in Ch.:1pter
5, and that's n o coincidence. Jus t as color aftereffects are caused by opponent
prOC'eSses for color v ision, MAEs are caused by opponent processes for mo-
tion d etection . We'll discuss these p rocesses in detail later in the chapter. Ler's
back up first and take a broader look at how the visual system mig ht go about
detecting motion.
The wate rfall illusion is jus t on e o f many m oti on illus ions. Akiyoshi
Kitaoka's art sh ows other powerful illusion s of mo tion (see , . . . W\v.ritsumei.
ac.jp/-akitaoka/index-e.html and Web Essay B.1: Perceiving Motion In
Time2
Static Images). Kitaoka's pictu res a re s tation ary on the page, but they can
produce a .s trong illusion of mo tion if you all ow your eyes to drift across
them. H ow d oes this work? \Ve d on' t yet have a consensus, but recent wo rk
s uggests that patterns like Kitaoka's Rollers elici t directional responses from
n eurons in m onkey rortex that correspond closely to the directio ns perceived
by huma ns (Conway et al., 2005).
Computation of Visual Motion

H ow would you build a m otion detector? Because m otion involves a change
in p osition over time, a logical place to .s tart is with ti..vo adjacent rerep tors
(call them neu rons A :md B) separated by a fixed d istance. A bug (or a spot
of light) moving &om le ft to right would first pass through neuron A's recep-
FIGURE 8. 1 Motion is a change in

position over time, as the m ovQfTlents
o f this ladyt:ug demonstrate.
motion aftereffect (MAE) The lllu-

sim of motion of a statlonar; object
that occurs after prolonged expcsure FIGURE 8.2 ThQ lowef falls
to a moving object. of Foyers.

VISUAL MOTION PERCEPTION 239
{a) ReceptivefuJds (b) (c) (d)

/\
:
' . '' '' .. '.
vv 8 mmaA
tive field, and then a sh ort time later it would enter neuron B's receptive fie]d
(Figure B.3a). hi. theory, a third cell that '1istensH to neurons A and B s hould
be able to detect this m ovem ent.
FIG UR E 8.3 Constructing a n9Ural
c ircuit for the detection of rlghtvYard
motion. See the t9XI: for details.
However, our m o tio n d etection cell (call it M) cannot sim,ply add up

excitatory inputs from A and B. Given such a neural circuit, M would fi re in
response to the moving bug, bu t it would also respond to two stationary bugs,
one in each recep tive fie ld (Figure 8.3b). To solve this problem, we need tw'o
additional components in our neural circuit, as shown in Figure 8.3c. The first
new cell, labele d '' D" in the figure, receives input from neuron A a nd d elays
transmission of this input for a short period of time. Cell D also has a fas t
ada ptatio n rate. That is, it fires when cell A initially detects light, but quickly
s tops firing if the light remains shining on A:s receptive field. Cells Band D
are then corUlected to neuron X, a multiplication cell. This multiplication cell
will fire only wh en both cells Band D are active. By delayi ng receptor Ns
response (D) and then multiplying it by recep tor B's response (X), we can
create a mechanism th.at is sensitive to motion (M).
This mechanis m would be direction-selective: it would respond well to
mo tion from left to right, but no t from right to le ft. A bug m oving from cell
B's receptive fie ld into cell Xs receptive field would cause Band D to fire in
the wrong order, so X would not receive its two inputs s imultaneously, and M
would not fire. The mechanism would also be h.lned to velocity because w hen
the bug is moving at just the right speed, the delayed response from receptor
A and the direct response from receptor B occur at the same time and therefore
reinforce each other. If the bug m oves too fast or too slow, the outputs from
Band D w ill be out of sync. This simple "bug" m otion detector is based o n a
model developed by Werner Reichardt in the 1950s to explain how flies detec t
motion (Reidlardt, 1986). Almost all models of hwnan motion detection are, at
their core, an elaboration of Reic:hardt's model adapted to the spatial-
frequency filtering properties of the human visual system (see Chapter 3).
One elaboration, developed by Barlow and Levick (1965), uses w hat electrical
engineers would call an" AND gate." Cell X fires ii and only if both its inputs (B
and D) are frring simultaneously, and it passes this message on to the motion
detection cell M. The AND concept is almost certainly n ot correct, a nd a m ore
elaborate version-developed by Adelson and Bergen (1985) a nd based on
linear filters that delay, sum, and then are followed by nonlinearities-seems
closer to the " tn1th. " In Web Activity B.1: Motion Detection Circuit, which
presents a m ore elaborate ve.rsion of this neural circuit, you can interactively
move spots of light around and see how it works.

240 CHAPTER 8
A m ore realistic circuit would indudeadditional receptors to detect longer-

range m otion , as shO\--\'T\ in Figure 8.3d. H e 1'E", the Iv[ cell fires continually as
the bug moves across the fields of the five receptors at the top of the circ uit.
If you \>Vere the "circuit designer/' how would you change the speed tunin g
of this circu it?
Apparent Motion
apparent motion The illusory One possible objection to this neural circuit is that it does n ot, in fact, require
Impression of smooth motlcn resultlng continuous motion in order to fire. An image of a bug that appears in A's re-
frcm the rapid of objects ceptive field, then disappears, and then reappears in B's recep tive field a short
that appear In different locations In
rapid succession. time later ,.,•ill drive M to resp ond just as strongly as if the image had m oved
smoothly acros.s the two recep ti ve fields. Although it raises a va lid concern,
this observa tion turns out to be a virh1erather than a liability for the Reichardt
rnodel, because it prov ides an excellent explanation for a vis ual ill usion., called
apparent motion, that modem humans experience on a daily basis.
Appa'rent motion wa £1rst<8lt\itonstrated by Sigmund Exner in 1875.
Exner set up a contrapti on that would generate e lectrical sparks sep ara ted
from e..1ch o ther by a very s hort distance in space and a very s hort period
of tin.1e. Even though there were tivo separate sparks--that is, two d iffe rent
perceptual objects-observers swore that. they saw a single spark moving from
one p osition to another. (Apparent motion is d emonstrnted in Web Activity
8.2: Types of Motion.)
\ Ve in the h venty-first century experience apparen t m otion every time we
wa td 1 te levision , go to a movie, or use a computer. You are p robably aware
that an animated cartoon is really a series of still drawings. Objects sud1 as
Daffy Duck (Figure 8.4) ch an ge positions each frame, and wh en the frames
are sh o\Nl"l to us a t a s ufficiently fast speed (e.g., 60 frames per second), we
perceive these position ch.:1.nges over time as motion. Web Activity 8.2: Types
of Motion shows the hvo frames in Figure 8.1 at a speed that m..'1kes it easy to
Frame l Fr.ame2
Frame3 Fr.ame4
FIG URE 8. 4 Appar£int

motion on the screen: A
series of still Images, played
one after the o thg,r at thie
correct rate, produces the
apparent motion of cartoons
(like Daffy Duck impersonat -
ing Carmen Mkanda in these
four stills from the Loongy
Tunoo animation Yankl/113
Doodle DatfY).

(•)g VISUAL MOTION PERCEPTION 241
, _, lEJ p,,,,,. 2
FIGURE 8.5 Th& corr9Spondgnce probl9m. (a) The squar9 mOVQS down and to th9
right. For a dietGCtor that "sees" just the oont9lts o f the white square. the motion Is
ambiguous. ¥=} What the detector sees Is consistent w ith both rrotion straight down
and motion down and to the right. Sgg Wab Activity 8.2: Types of Motion for anl-
matad wrslcns.
infer motion. Live-action m ovies and television programs work in exactly the
same way, except that the frames are still photographs rather than drawings.
The Correspondence Problem

Although apparent motion turns out to be a win for our motion detection
circuit, one of the classic perceptual ''problems" that we've discussed over
and over in this book remains an issue for the Reichardt model. We can il-
lustrate this problem with the movies diagrammed in Figure 8 .5. Each movie
has two frames that alternate back and forth. The only difference between the
two frames is that the only object in the movie, the red square sh1dded with
small circles, has been shifted diagonally by a short distance. (These movies
are animated in Web Activity 8.2: Types of Motion. Viewing the animations
as you read this section will make the following discussion mud1 clearer.)
If you view the first version of the movie (depicted in Figure 85a) on the
website (click on "The Correspondence Problem" and select "Movie 1 ''),
you will dearly detect diagonal motion. This scenario appears to present no
problem to our motion detector: the square moves down and to the right, and
then back up and to the left, and detectors sensitive to these directions pick. up
and signal this movement. Now consider movie 2 (depicted in Figure 8.Sb),
where we cover most of the square with a black "mask," leaving three of the
circles viewable through a small window (in the figure the mask is illustrated
as transparent so that you can see what's behind it, but ·i n the movie the mask
is opaque). Beneath the mask, the square moves exactly as before, but if you
view the movie, you will quite dearly perceive up--and-<lown, not diagonal,
motiont (You will need to view the animation to appreciate the difference in
the motion perceived in the two movies.)
The larger issue that these movies bring up is called the COITespondence
problem for motion detection. Consider the close-up view of the sit uation
in Figure 8.Sb, shown in Figure B.Sc. Here we superimposed the two frames,
coloring the circles blue in their frame 1 positions and green in their frame
2 positions. The difficulty for our motion detection system is this: how does
it know which circles in frame 2 correspond to which circles in frame 1? correspondence problem n
Because we have motion detectors for all directions, one detector will sense motion detection, the prrolem faced
the diagonal motion implied by matching the circle labeled "A" in Figure
8.5c with the circle labeled r'C." But another detector will sense the vertica l
by tte motion detection system of
knowing whK:hfeature n frame 2
respcnds to a particular feature In
=-
motion implied by matching circle A with circle B. l11ese detectors compete to frame1 .

242 CHAPTER 8
d eter min e o ur overall perception . (Yo u rn n see some o ther d em o nstrations of

the corresponden ce problem in Web Activity 8.3: Motion CorresJX)ndence.)
FURTHER DISOJSSION of correspondence as it relates to stereoocopic

depth perception can be fc:urd in Chapter 6 starting on page 182.
The Aperture Problem

aperture problem The fact that The aperture problem is closely related to the corresp ondence problem . It
when a moving coJect Is viewed gets its name fro m. the fact that a different de tector may win this com petition
thrcugh an aperture (cr a receptive
w h en an object is viewed thro u gh an apertu re than would \Vin if we could
field), the direction of motlm of a kJcal
feature or part of tlie ob)OCt may be see the w hole o bject. Consider the d em ons tration illus tra te d in Figure8.6and
amblgu:us. web movie 2 (Web Ac tivity 8.2:Types of Motion, "1l1e Aperture Pm blem").
aperture An opening that allows
Here the obj ect is a grating moving behind a window, vvh k h is referred to as
mly a partial view of an object. the aperture. The motion d irection of the grating is ambiguous-the grating
could be moving up and to the left {perpendicular to the s tripes, but diagonally
overall ) (Figure 8.fu), but it could also be m oving just up (Figure 8.6b) or jus t
to the left (Figure 8.6c). The motion com poneT1t parallel to the gm ting cannot
be inferred fro m tl'1e vis ual inpu t. TI1is means that a varie ty of contours of dif-
fere11 t orient.;1tions movin g at different speeds can cause identical responses in
a mo tion-sens itive neuron in the visu al system . 'Witho ut the aperhu-e, there's
n o ambiguity and no problem. Bu t when \Ve view the grating through the
apertu re, the system appea rs to impose some kind of sh ortes t-distance con -
s traint, a nd thus the vertical- mo tion detector \viJ1S .
At this p oint you rnay be thinking it's rather s illy todaim tlt.."l. t an artificial
.si tuation such a s the one we' ve set up here poses a broad cha llen ge to the
visual sys tem. After all, how often is our view of a moving object limited to a
small wind ow as in the examples presented here? To unde rstand the broader
irnplica tion.s of the correspondence and apertu re problems, consider the fact
tha t every neuron in Vl (the primary visual cortex) has a limited receptiYe field.
8230336 AmMa A Jn o ther words, every \11 cell sees the world througli a small aperture. The refore,
as Rgure 8 .7 illustrates, non e o f the Vl cells (represented in the figme as gray
tri an gles) ca n tell wi th ce rtnin ty which visu al e lements corresp ond to on e
un other when an object moves, even when no m ilSk is presenL
TI1e solution to this proble m is to have an other set of neurons listen to the
Vl neur ons and integrate the potentially conflicting signals. As Figure 8.7
shows, only one d irection-d m,,.,... and to the rig ht-is consistent V\>ith wha t
all fo m Vl cells are seein g here, and this is the direction \'\"e perceive w hen we
see the object as a w hole. If the neuron represented by the orange trian gle in
the figure has access to all the Vl cells d etecting local-motion directions, it w ill
be in a position to compare their outputs and find this common den ominator.
(o7) (I>) (c)
FIGURE 8.6 Tl'ltl aperture problem: n re oont9r"lts of the circle are the same In (a),
(b), and (c), eVQn though the physic.al motbn Is quite d ifferent.

FIGURE 8 .7 BuikJing a global-motion detector. See

the too fer detalls.
Global-motion V1 local-motion
dete.:::to r detector
.... .·
Of course, we haven 1t specified how this global-motion detector performs this

comparison, but tha t question would lead us beyond the scope of this book.
As Duje Tadin fr om Vand erbilt University poin ts out, the aperture problem
is like the fnd ia n parable about the blind men and the elephant. In one version
of this par(1ble the king askssi.x blind men to deten:nine what an elephant looks
Bke by feeling different parts of the elephant's body. The firs t blind man feels
a leg and says the elephant is like a pillar; the second feels the talJ and the
elephant is like a rope; the third feels the trunk and says the elephant is like a
branch; the fourth feels the ear a nd likens the to a fan; the fifth feels
the belly and says the elephant is like a \o\-a ll; and the sixth feels the h1sk and
says the elep ha nt is like a so lid pipe. The king explains to them , "You are all
correct. ·n-1e reason each of you is tellin g it differen tly is because ead1 one of
you tou ched a different part of the e lephant. But, in fact the elephan t has all
th e features you mentioned." In the aperture problem, eac h of th e Vl cells is
a blind m an, and the correct answer comes from combinin g their responses .
Detection of Global Motion in Area MT

Thou gh a discussion of ho'"' g loba l-motion detectors work wou ld be too middle temporal area (MT) An
b road for this text, we ca n say something a bout where the g lobal-m otion d e- area of the brain thought to be Impor-
tectors are. We saw in Chapter 3 th at lesions to the magnocellular layers of tant In the perception or motion.
the lateral genicul a te nucleus (LGN) impair the perception of large, rapidly
moving o bjects. lnfom1a tion from magnocellular nemons feed s in to Vl and is
then passed on to {among oth er places) the middle temporal area of the cor-
tex, an area conunonly refe rred to as MT (Fig ure 8.8) in nonhwnan primates.
The human equi valent of MT has been localized using hmctional magn e tic
reson..1.nce imaging (fMRI) and vmiou sly labeled as MT+ or VS. Recent work
suggests tha t this m otion -sensitive are.a may h ave at least hvo separate maps
located on the lateral surface a t the temp ora l-occipital (TO) boundary. TI1e vast
majority of neurons in the MT are selective for rnoti on in one particular direc-

244 CHAPTER 8
FIG URE 8.8 Mo tion-sensitive areas in the hurnan

brain. The middle temporal ar&a (MT , gr99n) and medial
superior temporal area {M ST. turquoise) ar.e sh()l/$n
In rGlatlon to other visual areas (V1, V2. Y.3, and\.'&$.
(After Heeger, 2006.)
tion, but th ey show little selectivity for fo m1 or color. But do these MT cells
correspond to the orange ne uron in Figure 8.7, which responds to large-scale
motion of w hole objects, or are they more like the lowalevel motion detectors
re presented by the grny triangles in Figure 8.7?
To find out, Newsome and Pare (19E8) trained a group of mon keys to respond
tocorreL,ted-dotmotion dispki ys(FigureB.9). ln Figure8.9n,all the dots (100%)
are m oving to the right. In Figure 8.9b, SOo/..., of the d ots have corre l'1ted m otion
(they are moving in the same direction), w hile the motion of the res t o f the dots
is uncorrela ted (these dots are moving in directions). In Figure 8.9c,
onl y 20% of the d ots have correlate d motion . Jus t as in the aperture- proble m
deinons tra tion, n o s ing le d ot in these displays is s ufficient to determine the
overall d irection of correlate d motion. So, to d etect the correlated directi on, a
neuron must in tegrate info m1a tion from many loca l-mo tion d etectors. (Note
th.a t in the actual stimuli, all dots a re the same color a nd no a rrows me p resent.
You rn n get a much better feel for w ha t Newsome nnd Pare's m onkeys were
expeiiencing by visi ting th e "Corre lated-Dot Motion" section o f WebAcUvtty
8.2: Types of Motion.)
Once they \.'/ere fully trained, the monkeys in N ewsom e and Pa.re's study
could recognize the correla te d-motion direction w hen on ly 2--:3% of the dots
were moving in th is direction . The researchers the n lesion ed the monkeys'
MT areas. Foll owing the s urgery, the m onkeys needed about ten times as
ma ny corre la te d d ots in order to correctly id entify the direction of mo ti on.
H owever, the monkeys' ability to discriminate the orientation of s tationary
patte rns was generally wi.impaired. Interestingly, the m onkeys' performance
in the correla te d-dot-m otion task improved marked ly during th e weeks fo l-
lowin g the lesion , presumably because they learned to u se othe r brain areas
to d iscriminate motion.
FIG URE 8.9 Thlili NeV11Som 9 and Par9 paradigm . Th9

obSf\f'Vlfs t ask is to identify th9 direction o f rnotion o f
the correlated dots. shown in purple here. On the actual
s timuli thQ dots are not cobred and are no arrows.) (n) 100% (b) 50% (c) 2C1l'o

8230336 AmMa Appa
Lesion s tudies have been central to o ur understa nding of the specificity of

brain areas. H owever, s uch studies are often less than completely compelling,
because lesion s (even nice 11cle an '' chemical lesions, like those used by New-
som e and Pare) may be incomple te o r may influe nce other s truchtres. To test
the involve1nent of lvIT ne urons in g loba l-mo tion perce ptio n m ore
et al. (Salzman, Britten, and Newsome, 1990) trained a ne\v g roup of
1nonkeys to discriminate correlated-motion directions and then poked around
in the monkeys' MT areas to find groups of neurons that responded to one
particular direction. Once they had found a group of neuro ns that responded ,
for example, to motion" they showed the monkey a new set of
s timuli and e lectrical.ly s timulated the identified MT neurons. Re markably,
the m onkeys s howed a s trong te ndency to report m oti on in the stimulated
neurons' preferred direc tio n, even when the d ots they were seeing were actu-
ally moving in the opposite direction[ These results make a very stron g case
that the MT is the si te of global-mo ti on de tection n eurons in the visual system .
Motion Aftereffects Revisited

We know that the m onkey brain is very s imilar to the htunan brain, but it is
always nice to es tablish convergmg evidence su gges ting tha t results fr om
m onkey labs apply to human vision as well. ln the case of mo tion perception,
researchers h ave looked for th is eviden ce us ing the moti on aftereffect. As
noted earlier, the existence o f the implies a n opponent-process sys te m
mud\ like the one that plays a role in color vision (discussed in Chapter 5).
When we view a .stationary object, the responses of neurons tuned to differen t
directions of m oti on are n orma lly balan ced . lhlt is, neurons sensitive to up-
ward m otion fire at the same rate as ne urons sens itive to d ownward m o tion ,
so the signals can ce l out and no m otion is perceived . But w hen we look a t a
waterfall for a p rolonged period, the d etectors sens itive to d ov\fnwaid motion
becom e adapted . When we then S\vitch our gaze to a s tationary object, s ud"\
as the rocks next to the waterfr1ll, the neurons sensitive to upward motion fire
faster than th e h1ckered-out d ownwa rd-sens itive neurons, and we therefore
perceive the rocks as drifting up.
You might wonde r w hy any motion d e tectors would fire in response to
a s tationary rock, but as we will see in a bit, mu eyes are constantly drifting
around, so th ere is always a s mall amount o f retinal motion to s timulate mo-
tion detectors at least s lightly. You might a lso wonder wha t h appens when
we adapt to d ovvnward motion and the n see something moving h orizontally.
Make a predic tion, and the n te.s t your h yp othesis us ing Web Ac tivity 8 .4:
Motion Attereffects.
H ere's another experinlent to try wit h this web activity: adapt for 15
seconds or so to rightward motion with your r ight eye ope n but your left
eye clreed, and then quickly switch eyes, dosing your right eye and op ening
your left. You should exp erience an aftereffect that is only slightly reduced
in magnitude compared with the aftereffect you get with b oth eyes open the
w hole time. What doe.s this interocular transfer tell us a bout the locu s o f the
MAE in the visual syste m? Could m otion aftereffects besubserved by ne m ons
in the retinas?\ Vhat abo ut ne urons in the LGN?
The fact that a strong MAE is obtained when one eye is adapted and the
other tes ted mea ns tha t the effect mus t be reflecting the acti\·ities of ne m ons
in a part of the visual system v...!i.ere information collected fr orn the two eye.s
is combined (Ray mond, 1993). As we learned in C hapter 3, input from bo th
eyes is not combined until Vl, where neurons sh ow a p refe1en.ce for input from Interocular transfer The transfer of
one eye or another but respond to some exte nt to stimuli in both eyes. Recent an effect (such as adaptation) from one
advances in functional imaging techniques may make it possible to loca te "fe to the other.

246 CHAPTER 8
first-order motion The mo tion of the site of m.o tio n a fte reffects even m o re precisely. The e m ergi ng e,.;dence
an object that Is denned by c hanges In s uggests tha t the MAE in lnm1ans is ca used by the same brain region shown
lumlnanoe. to be resp onsible for global-motion detection in monkeys: cortical area MT.
luminance-Oeftned object An For ex.ample, David Heeger and colleagu es (Huk, Ress, and Heeger, 2001),
cbject that Is delineated by dWferences after controlling for the important effects of attention, demonstrated that the
In reflected light.
direc ti on-selec tive a d aptation produced a selective imb alance in the fMRI
motion The motion s ignal in human MT, providing evid ence that MAEs are due to a population
of an object that Is defined by changes
In contrast or texture1 but not by
imbalance in area MT. Remarkably, recent work sh o\11/S that motion aftereffects
luminance. can occur even after very brief exposures-as little as 25 millisecond s, and that
these can be explained by direction-selecth-e responses of neurons in MT to
texture-defined object or contrast-
deftned object An object that Is
subsequently presented stati onary stimuli (Glasser et al., 2011).
defined by dlffererces In contrast. or
texture, but not by lumlnanoe. FURTHER DISOJSSION of the role of the striate cortex (V1) in combining
input from both eyes can be found in Chapter 3 on page 72.
Up to this point our discussion has focuse d on motion-the

change in position of lt..ndnance.-deftned objects over time . In the next section
we describe anoth er interesting m otion phen omenon: second-order motion,
in w hich texture-defined objects (also ca lled contrast-defined objects)
change posi tion over time.
Second-Orderlv1oVon
Second-order mo tion, illustrated in Figure 8.1 0, s an interesting phenom-
enon. TI1e three frames in t11e figure look like collections of rand om black and
white d ots, which is w hat they are. But if you go to the \l\>'ebsi te and look at
the sequence of frames p layed as a movie, you \l\·i ll dearly pe rceive a set of
leftward-mo\oing stripes. (Web Activity 8 .2: Types of Motion includes some
additional frames that lead to the perception of an infinite loop in w hich the
s tripes move leftward off the edge of the movie and reappear on the right side.)
TI-US mov ie is constructed from a combination of two patterns: a random
collection of s mall whi te and black d ots, a nd a series of "vide w hite and black
stripes. Tile patten-.s are overlaid and combined in such a w<Jy tha t d ots covered
by w hite bars are inverted (black d ots turn w hite and vice versa), and d ots
co\o-ered by black ba rs are left alone. TI1e frames o f the movie are made by the
process of shifting the bars to the left while the d ots re main s tatio na ry. T11e
resulting perception of m ovem ent is called "second-order m otion."
A.s in firs t-orde r a pparent-motion disp lays, n othing actuall y m oves in
second -order mo tion. Even more in credibly, there is nothing t o m ove in
second-order m otion. As Figure 8.11 s hows, the only thing that changes in
our second-order-moti on movie is tha t strips of d ots are inverted from o ne
frame to another: fro m frame 1 to frame 2 the d ots in the oran ge s trip me
inve rted; then fr om frame 2 to &ame3 the d ots in the blue s trip are inverted.
Frnme t Frami22 Frame 3
FIGURE 8.10 Second-ordEf motion,

If you try Web Activity 8.2: Types of
Motion you w ill qu ite clear1y Se€! stripes
moving fro m rig ht to left across the field
o f dots wen though you cannot see
the stripes in thooe static images. See
the t9xt for details.

VISUAL MOTION PERCEPTION 24 7
FIGURE 8. 11 Close-ups of ong section

of the fram9s in Flgur9 8. 10, illustrating the
chang9S from fram& to tram&.
Just as random dot stereograms prove that matching discrete objects across
the two eyes is not necessary for s tereoscopic depth perception (see Chapter
6), second-order motion proves that matching discrete objects across m ovie
frames is not necessary for motion perception.
Several lines of evidence'<>uggestthat the visctll'!)IS!em includes specialized
mechanisms for second-order motion. For example, Lucia Vaina et al. {Vaina
and Cowey, 1996; Vaina et a l., 1998) described one neurological patient who
suffered brain damage that impaired the perception of first-order motion but
not of second-order motion. A second patient showed the opposite path:m!
impaired second-order but spared first-order motion perception. These two
patients had lesions in different brain areas, and together they demonstrate a
double dlssoclatJon of function, which wouJ d not be possible if there were a
single motion mechanism. Tun Ledgeway demonstrated a motion aftereffect
for second-order m otion and found that the second-order MAE transfers even
more com pletely between eyes than does the first-order MAE (Ledgev...·ay,
1994; Ledgeway and Smith, 1994).
Why did we evolve a motion detection system for something as esoteric as
second-order motion? It turns out that second-order-like motion does occur
in the real world, especially when an object is effectively cam ouflaged . This
chapter's home page o n the website shows a situation in which something
like second-order motion makes an otherwise invisible object spring to Ufe.
(See Web Essay B.2: Beyond Second- Order Motion.)
Using Motion Information

Now that we know something about how the motion perception system works
under the hood, let's ronsider some ways we might use motion information
to interpret the world around us.
Going with the Flow: Using Motion Information to Navigate

For sighted people, getting around is easy. Our vision effortlessly guides our double dlssoclaUon The phenom-
motion thro ugh space, whether we're walking or driving, so that we usu- enon In which one of two funcUons,
ally read1 a destination quite safely and efficiently. Indeed, safe navigation such as first- and seocnd- order
is one of the primary functions of the visual system. What infomlation does motion, can be damaged without
the visual system use to help us navigate our way across a busy intersection? harm to the othE<, am vice versa.
See also Chapter 4.
Or ronsider the challenge that confronts a pilot landing an airplane at high
speed. In an attempt to improve World War II pilots' performance in such opUc array The ocllectlon of light
rays that Interact with objects In the
situations, J. J. Gibson (1957), who was working for the US Army at the time,
world that are In front of a "1ewer.
developed an influential theory about the optic an-ay, the col1ection of light Term coned by J . J. Gibson.
rays that intern.ct with objects in the world in front of a viewer. Some of these
opUc flow The changing angular
rays strike our retinas, enabling us to see. Gibson argued that when we move positions of points In a perspective
tluough our environment, we experience pa ttems of optic flow that our \·isual Image that we experience as we move
systems use to detenuine where we're going. through the world.

248 CHAPTER 8
Consider a pilot corn.Ing in to land her plane. As the plane approaches the
ground, the optic array VI.i ll expa nd in a pattern known as " radial
expans ion" 8.12). The pilo t's h ea ding-the specific point at which
she is ai ming her plane-will always be the center, or focus of expansion, of
the o ptic array. As ind icated in the fig ure, the focus of expansion is the on e
place in the vis ual field tha t '"'; II be stationa ry, so if the pilot's visua l system
ca n locate this sta tionary p o int, it can de te m1i.ne the heading. (Fo r a releva nt
d em ons trati on, see Web Activity 8.2: Types of Motion.)
G ibson was able to derive a numhe r o f optic flow he uristics that the visual
system might use to na\dgate arOLmd the world. At the m ost basic level, the
mere presence of optic flow indicates locom otio n, a nd a lack of fl mv is a sig-
n al tha t you are s tationary. If you have ever been sitting in a station a ry train
a t a busy railroad s ta tion w he n the train beside you moves, you may have
FIGURE 8.12 The optic flow field experienced a situ ati on in which optic flow alone gives you the illusion. tha t
produC9d by rrovement forward in you (ra ther than the neighboring train ) are in m otion . Outflow (flow toward
space. FOE = foc:us of expansbn.
the periphery, as depicted in Figure 8.12) indicates tha t you are app roaching
(After W. H. \"larrern and Saunders,
1005.) a particular destina ti on.; inflow ind icates re trea t (ass wning that you r head is
facing forward). And as mentioned alread y, the focus o f expan sion, or ufocu s
of constriction 11 if you 1re lookin g forvvard \vhile driving in reverse, tells you
""·here you 're going to or coming from.
8230336 Am111 Are humnns actua lly a ble to make use o f op ti c flow information? Labo-
rolories like Bill Warren's have used elaborate computer-generated displays
of moving d ots and lines to simul a te optic flow information, and they have
made a good de.a] of progress in understa nding both the utility and complexity
o f optic flow in.formation.. For example, Warren et a l. (W. H . \o\farren, Morris,
and Kalish, 1988) de monstrated tha t huma ns could e..sti mate their direction o f
heading to \vi thin about 1 or 2 degrees, us ing as their sole g u ide the pattern
of op tic flow simulated by the m o\·ing d ots, even w hen the display contain ed
only a very small number of d ots.
O f cou rse, the nice clean opti c flow pattern diagrammed in Fi gure 8.12
occurs only if the head a nd eyes re main fixed and p ointed straight a head . As
soon as gaze s hifts to on e side, a new "radial" component is introduced to the
optic flow. H owever, Warren showed th.at observers were <lble to discount these
radial components, both when the observers m oved their Ot'Vll eyes and w hen
the compu ter-generated display mimicked the radial fl ot'\' caused by an eye
movernent QN. H . Warren and Hann on, 1990). If the radial shift is relative ly
s l0tv, observers can compen sate for si mula ted eye m ovem ents jus t as re.:1di ly
as they d o for reaJ eye m ovements; but with faste r simulated eye m ovement
speeds, performance breaks d ow n (Royden 1 Banks, a nd Crowell 1 1992). This
result implies that the visual system can make use of the copies of eye muse.le
s ignals w he n it is processing optic fl ow information. We1 1l discuss this notion
la ter, in the section titled ''Sacca dic Suppress:ion and the Compara tor."
Something in the Way You Move:

Using Motion Information to Identify Objects
focus of expansion The point In The fact tha t we use m otion information to guide us as we move through our
the oenter of the horizon from which, environment is n ot aH that s urprising. Wha t m ay be less obvious is that rno-
when we're In motion (e.g .1 drlvirq en tion can also h elp inform us about the nattu·e of objects. About 40 years ago,
the hlgl1way). all points In t11e pernpec-
tlve Image seem to emanate. The
G unnar Johansson (1975) recogni zed tha t the.re m ight be some thing specia l
focus of expansion is one aspect of about the m otion of animals a nd motion-that h elps us
q:itlcftow. identify bo th the m ovin g object and its actions.
biological motion Tlie pattern of Consider the rennis pl ayer in Figure 8 .1 3a. lt's clear from the contours of
movement o f living belrgs l}turnans her body tha t sh e's just sm ashed a n innocent little tennis ball in the directi on
and animals). of her opponen t Figure 8.13b sh ows the same tennis p layer in the dark. All

(b)
•
• •
• • •
•
• •
•
FIGURE 8.13 Biok:glco.I motion can bQ SQQn comp9111ngly whQl'l lights attach9d to a
moving hLHnan are 'Yiev..red in totaJ darkness. You can expefi€>11ce blologicaJ rrotion in
Web ActMty 8.2: Types ot Motton.
we can see are the little lig h ts a ttached to h er ankles, knees, hips, elbows,
w rists, and sh oulders. There's n ot very much in the s tatic pattem of the lights
to inform us that the contour is a human (le t a lone a woman ), o r that s he/
it is engaged in athletic activi ty. \Vhat Johansson d iscovered, tho ugh, is th.at
w hen the lights move, their motion gives the viewer immedia te and very
compellin g impression of a live hmn an in action. O nce agai n, you really need
to see this to apprecia te it, so watch the classic " d ot wa Lker'' m ovie, Lmage 3
in Web Activity 8.2: Types of Motion.
There is even evidence that observers can use biological m otion to identify
w hether a set of movin g lights is attached to a male or fe male walker. How
d o th ey do that?,A5 we '\valk "·hen the right leg is in hont of the left leg,
th e riglif shot td.eris behind, ·cE versa. If we draw one line connect-
ing the left s hould er with the righ t hip and another connecting the rig ht
sh oulder with the left hip, the intersection o f the two lines is the center o f
the wa lker's motion. Because males typic...1 ll y have broader s houlders and
narrower hips than females, the average male's center of m oti on is h igher
than tha t of fem.:i les. James Cutting e t a l. (Barclay, Cutting, and Kozlowski,
1978) s uggested tha t observers use estima tes of the center of m otion as a
cue to the wa lker's gender. O ther s tudies (e.g., Mather a nd Murdoch, 1994)
suggest that, in certain views, the a m ount of body sway might provide an
even more salien t gender cue; and other work (Neri, Morrone, a nd Burr, 1998)
shows tha t the m echanisms that analyze biological motion obey differen t
rules for integrating m oti on over space and time th an d o mech anis ms for
other forms o f complex m otion.
Biological motion appears to play an important role in how we t.mderstand
hmnan actions. Recent stud ies of biological motion with hvo ''actors" either
dan cing or figh ting show that we are much 11"1.ore efficient in discrinU.nating
biological m otion of a human when h vo humans are acting in synch rony (e.g.,
fighting o r d ancing) than when they are out of sync. This finding suggests
that when we wa tch h\·o people in temctin g, knowing w hat onE"is doi ng helps

250 CHAPTER 8
time to collision (TTq The time us unders tand the actions of the othe r. \Ve c.an think of this phe n omenon as
required for a moving object (such as proof of the old adage tha t ''i t takes h \.·o to tango" {Neri, Luu, and Levi, 2006).
a cncket balQ to hit a stationary object
(SUch as a batsman 's head). TIC = Avoiding Imminent Collision: The Tao of Tau
distance/rate.
In cricket, as in so irtuch of life, the object is to keep yom eye on the bal l and to
tau (1) Information In the optic flow
that could signal time to collision (ITC)
s tay out of trouble. Indeed>much of ou.r visua'l may be designed to
without the necooslty of estimating do just that. TI1e bats man s how n in Figure 8.14 eviden tl y did no t keep his eye
either absolute distances or rates. The on the ball. Luckil y, acc ide nts like th is are not too common, beca use cricke t-
ratio of the retinal Image size at any ers are extremely good a t judging precisely w hen a tiny ball, hurtling toward
moment to the rate at which the Image the m at s peed s appro..1ch ing 100 miles per ho ur, is about to collide with thei r
Is expanding ls tau, and TTC ls propor·
head (Regan, 1991). \Vha t visuaJ informa tion do we use to avoid imminent
tlonaJ to tau.
collision s, or to achieve collision when catching or batting a ball? To rephrase
somewhat m ore precisely, how d o we estima te the time to eolllslon (TTC) of
an approaching object?
Consid er a small, red cricket ball, thrown by a large, ra the r an gry-looking
man (o n the o ther team), that bounces off the ground i.'1.bout 10 feet \Vay from
you. At this distance, if th e ball hurtles tmvard the bridge of your n ose a t a
constant rate of SO feet per second, it vvill collide with your face in 0.20 second
(TTC = dis tance / rate= 10/ 50). The m ost d irect way to estimate ITC would
therefore be to estimate the d_is tance and s peed of the ball. Ho v.'E:ver, determin-
ing absolute distances in depth is a tricky propositio n, as w e saw in C ha p ter
6, and humans are far better a t jud ging ITC than would be p1·ed icted on the
basis of their ability to judge di stance.
ln a n a tte mpt to reconcile this apparent discrepa.11cyr D. N. Lee (1976) and
o thers have poin ted o ut that there is an alte rnative source of infonn.ation in
the optic fl ow that co uld signa l TIC '"'; thout the need for a bsolute dis tances
or rates to be estimated . Lee called this inforrnation source tau (r). Here's h ow
tau works. As it approach es yo ur nose, the image of the ball on yotu retina will
grow larger (if you don' t be Beve this, have someone throw a ball- preferably
a so_ft one-at your face and see for yourself). The ratio of the retina l image
size a t a ny m omen t to the rate at w hich the image is expanding is tau , and
TIC is proporti on..,t l to tau . The grea t advantage o f us ing tau to estimate TIC
is that it relies solely on informa tion available di rectly from the retina l image;
all you need to do is track the visual angle subtended. by the cricket ball as it
a pproach es your eye.
Do we actually make use of ta u ? 1l1e jury is still out. ft is clear tlwt estimating
the time to immine nt collision is critically important to animals and human s,
and almos t e very species tested will a tte mpt to avoid a .simulated collision.
There is also evidence that certain n eurons in the visual systems of pigeons
and locusts respond to objects on a collis ion cou rse with the m and can signal
a particu la r time to collision (Rind and Simmons, 1999; Wang and Frost, 1992).
Interestingly, a looming object on a collis ion path with an observer ca ptures his
attention w hereas a looud.ng object on a near-miss pa th d oes n ot, even w hen
the observer is not aware of an y difference between collision and n ear-miss
objects (Lin, Murray, and Boynton, 2009). H owever, Tres ilian (1999) concluded
th..1.t ta u is jus t one of a number of different sou rces of visu al infonna tion tha t
ca n be used to judge the time to coll ision .
Eye Movements
Ch.u eyes are con sta ntly m oving. When they m ove, the image on the retina
moves too. So h ow d oes our brain fig ure out whid1 motions on the retina be--
FIGURE 8. 14 This batsrnan has just long to rea l moving objects: a nd w hid1 are ca u sed by our m-vn eye an d head
been hit by a very llard cricket ball. rnove.m ents? Let's try a simple exp eriment. Pla ce a blank piece o f paper on

(a) (b)
•8Z30'>'>'-' pp.:i
•
FIGURE 8. 15 Stucfying gye movernents. (a) Axing our gaze on the dot whlle the
pencH moves to the left: causes the penc;ll to generate motion across the retina. and
we pgrcg,lve mOVQ'TIQnt o f th& pgrlCil In this dlr9Ction. {b) Rxating our gazg on th9 pgi-
al while it mows to the right caJS9S the dot to gen9rate motion In the same direction
across the retina. but we do not peroalw movement o f the dot.
the desk in front of you, and draw a small black d ot right in the center o f the
paper. Close your le ft eye a nd foc us your right eye's gaze on the dot; the n
p osition a pencil so that it is n ear the bottom right com er of the paper. Keep-
ing your eye trained on the dot, m ove the pencil slowly across the s heet to the
le ft side, as sh own in Figure 8 .15a. 'What did you perceive? The image of the
pencil jus t s""-ept across your retina from right to left, so asswning tha t your
rightward-motion detectors are functioning correctly, you should have perceived
movement in this direction. (You may have thought tha t the word righhvard
in the previous sentence was a typo, but technically the image sweeps across
the retina in the op posite direction from the actual m oveme nt, s ince the right
side of the world p rojects to the left s ide of the retina and vice versa. To avoid
confusion , we'll ignore this inconvenient bit o f physics for n ow and pretend
that images move on the retina in the same direction that the objects in the
world are m ovin g. Web Activity 6.& Eye Movements provides illustrations
of these phenom ena tha t honor physics.)
Now try a slightly different demonstration . Start with the pencil near the
bottom le ft corner of the paper, fixate the eraser, and track it with your eye as
you m ove it back across the sheet of paper to the right corner (Figure 8 .15b).
Congratulations! You have justexe.:uted a type of eye movement called smooth
pursuit, w hich kept the image of the pencil stationary on the retina w hile it was
in motion. But think about what happened to the image o f the dot just above
your pencil. When the pencil was on the left side of the page, the dot was to
the right of your fixation point. As you tracked the pencil, the dotshifred to
the center of your retina, and then it s lid to the left of your fixation point once
the pencil read1ed the right s ide of the paper. However, you sho uld not have
perceived the do t to be moving in this case, even though the image of the dot
made essen tially the same journey across your retina that the image o f the
pencil did in Figure 8.15a.
We hope you 've convinced yourself that the retinal image movement of
the pencil when you keep your gaze centered on the d ot {see Figure 8.15a)
is essentially the same as the retinal image m ovement of the d ot when you
keep your gaze centered on the pencil (see Figure 8.15b). The question is, smooth pursuit A type of voluntary
Why do we perceive motion of the pencil in the first case, but perceive that eyemovement in which the eyes move
the d ot is sta tio nary in the second case? TI1e reason is that in o ne case there's smoothly to follow a moving dlject.

252 CHAPTER 8
superior colllculus A structure In an eye m ovement. Think about how the visual system might accomplish this
the mldbfalnthat Is Important In Initiat- balancing act while we back up a nd describe eye 1uovements in a bit more
ing and guldklg eye movements. detail (By the way, you may want to save the piece of paper with the do t for
another exercise later in this chap ter.)
Physiology and Types of Eye Movements

8230336 Ar As Figure 8 .16a shows,six muscles are attached: to each eye, arranged in three
pairs. These muscles are contro Ued by an extensive network of s truch1res in
the brain. One way to get some inkling of the role of these brain structures is
to stimulate them with small electrical s ignals a nd observe the movements
of the eyes. For example, if a cell in the supertorcolllculus (Rgure8.16b) of
a monkey is stimulated, the m onkey's eyes will move by a specific a mo unt
in a specific direction. Every time that cell is stimulated., the same eye move-
ment wiU res ult. Stimulating a neighboring cell will produce a different eye
movement (Stryker and Schiller, 1975). (The s uperior colli culus also gets some
input directly from retinal l'f'nglion cells; this input presumably helps with the
planning of eye movements.) By contrast, in response to stimulation of some of
tl1e cells in the frontal eye fields (Figure 18.16b) (and, indeed, certain superior
colliculus cells too), the monkey will move its eyes to fixate a specific spot in
space. Depending on where the eyes start, this adjustment may require an eye
movement up, d own, left, or right. In this case, it is the d estination and no t the
movement that is coded (Mays and Sparks, 1980; Schiller and Sandell, 1983).
(a)
Latera1 view Top view
./Superior
/ obliqu<
•
rectus
I
Inferior oblique lnferiorreictus
FIGURE 8.16 Muscles oftheg.;e. {!!}Six musckis-

arranged in thr8'1 pairs (superior and inferior
sup91'ior and inferior roctus, and medial and lateral
rectus)-are attached to each !J'ffJ· '1:;i) Brain c irc uits for
visualty guided saccades. {Part b after Wurtz, 2000.)

This d escription merely scratd1es the s urface of a motor system that is not mlcrosaccade An Involuntary,
only very complex1 but very active. For example, even when we try to h old small, Jerkllke eye movement.
our eyes completely s tationary, they continue to execute srnall but important vergence A type of eye movement
move ments. Specifically, there are involuntary eye drifts and small jerks (mic- In whlcl1 the two eyoo move in oppo-
rosaccades); if the eye muscles are te mpora rily paralyzed-say, as a result of site directions; for 9X81llple. both eyes
turn toward the nose (convergerce) or
taking curare (it's amazing what some people will s ubject them se lves to in the
" /'lay l'om the nooe (diVefgenoeJ.
name of science]) (Matin e t al., 1982}-the en tire visual world gradually fades
saccade A fype of aye movement,
from viei,.v, Of course, in n onnal viewing. image velocities are fast enough to
made both Vduntarlly and lnvolun-
prevent fading \vithout the need for microsaccad es (Kowle r and Collewijn, tarly. In v.tilch the eyes rapidly c hange
2010). So what are microsaccades good for? Recent work suggests tha t they fixation from cne cbject or location to
m:ay be important for very fine s patial judgments, s u ch as threa din g a needle, another.
because they precisely move the eye to nearby regions of interest (Ko, Poletti,
and Rucci, 2010), and for compensating fo r the very rapid falloff of acuity even
a fe\.v minutes outs ide the fovea (Poletti, Llstorti a nd Rucci, 2013).
In additio n to invo luntary eye m ovements, the re are three types of vol-
untary eye movements. Most ob vious, perhaps, are the previously d iscussed
sm ooth-pursuit 1novemen ts that we make w hen tracking a moving object, TileSe
sm ooth-pursuit eye movements are often used by d octors as a simp le screenin g
for neurological impairments,and they can he lp distinguish schizophrenic
patients from others {Benson e t al ., 2012). Vergence eye m ovements occur w h en
we rotate our eyes in\'\:ard (con verging the eyes) o r o utward (divergin g the
eyes) to focus on a near or far object. The third type of voluntary movement
is the saccade:a fast jump (up to 1000 degrees persecond)(lla hill and Stark,
1979) of the eye that shifts our fi xatio n p oin t fro m one spot to another. We can
d ecide to make a saccad e d elibe rately, but w he ther we' re thinking about it
or n ot we w ill ma ke three or four saccades every second of every minute of
every wa king ho ur of the day. Tha t's some thing like 3 saccad es x 60 seconds
x 60 minutes x 16 h ours= 172,800 saocades per day- and tha t doesn' t include
the s::iccad es we make during our dream s in rapid-eye-movement (REM) s leep
(Moquet et., l., 1996).
\'\ihen we view a scene, our saccades are n ot ra.ndwn. \Ve tend to fi xate the
"interesting" pL.1.ces in the image. Thus, theeyes are more li kely to make sac-
cades in response to contours than to broad fea tureless areas of an image (Figure
8 .17). Tnterest also h as a richer sem antic m eaning: We m ake eye movements
tha t a re based on the content o f a scene and on our specific interests in tha t
F IGURE 8. 17 A classic scan path (.righ1;1

showing the pattern o f eye movements dur-
ing Inspection of the picture of the girl on
th19 li9ft. (From Yarbus, 1Q67 .)

254 CHAPTER 8
reflexive eye movement A rnove- scene (Yarbus, 1967). O ur pattern of eye movements as we en ter the cafeteria
ment of the fJ'fe that Is automatic ard will be different if we' re looking fo r lunch than if we're looking for love.
lnvolu ntary. Tilere are also reflexive eye movements-for example, when. the eyes rnove
optoklnetlc nystagmus (OKN) A to compensate for head and body movement w hile maintaining fi xa tion on a
reflexive eye movement In which the particu.lar target. These a re known as vestibular eye movements and operate
eyes will Involuntarily track a continually
via the vestibttlo-octtlar reflex (V OR) (see Chap ter 12). In optoklnetlc nys-
moving objoct.
tagmus {OKN) th e eyes wi ll in voltmtarily track a continually m oving object,
saccadc suppression The movin g s moothly in one direction (e.g., right) in pursuit of the object moving
reduction of visual sensltMty that
occurs when we make saccadlc eye in that sa me direction , and the n s nap back. T11e presence of OKN in response
movements. Sacca.die suppression to moving stripes has often been used as a measure of visual acuity in infan ts.
eliminates tt'<l smear rrom retina Image
motion during an eye movement. Eye Movements and Reading
1f yo u are reetding this chap ter, then your eyes are d o ing a lot of m ovin g!
Reading English involves fi xa ting for roughly a quarter of a second, and then
making a s.a.ccade a( about 7- 9 letter spaces--over a nd over again. We make
.saccades in o rde r to bring the tex t onto our foveas, because print that's too
far &om our fixation cannot be read, in p art because of vis ual crow ding (see
C hap te r 3 a nd Levi, 2008}. Interestingly, readers of English are able to gain
infonnation from u p to 15 characters to the right of fixation, but on ly 3--4
charac ters to the left. Thus, the percep tual span is asymmetrical. Readers of
Hebrew (whid1 is read fro m right to left) have th e reverse asymmetry. Readers
of both Hebrew and English can svvitc.h this asynunetry depending on w hid1
lan guage they' re reading (Rayner, 1978), so the asymm ehy is a ttentional, no t
a p roduct of limitations imposed by the visual system .
As we d iscuss next, there is no in.formation prOC'E'Ssin g during saccades, so
while we're reading, all of the information processing mu.st take place d\uing
the fixations . Interestingly, such processing occurs during on ly a s mall frac-
tion of each fixation. "Disappearin g text" experin1ents (in which the words
achtally d.isappe:u off of the computer screen while participa nts a re reading)
reveal tha t if a word rem ain s on the screen for onl y 50 milliseconds after it is
first fixated, reading proceeds normally (Rayne r et a l., 2003). TI1ere me mnny
othe r interesting aspects of eye moveme nts in scene pe rception and in visual
.search and attention (see Chnp ter 7).
Saccadic Suppression and the Comparator

Now let's return to the tricky problem of d iscrimina tin g 1notion across the
retina that is due to eye movements versus object m ovements. Let 's do one
more d emon stration using tha t white piece of paper w ith the d ot in the mid-
dle1Close i:;o ey and &'"'just to the left of the d ot; then execute
so-:ne saccades, shifting yotu eyes back and for th to the rigl1t and then to the
left of the dot. The d ot wi.11 be moving across yom retina,. but you should no t
experience any perception of movement. Now,. vvi th your left eye still dosed,
fix..'lte the dot, p lace your right index finger on the right side of your right eye
socket, •md gently "jiggle" yo ur eyeball. Now the d ot (as well as the paper
and the desk the paper is sitting on ) s110111d appea r to move back a nd for th!
\'\1hat's going on here?
Pa rt of the answer is thought to be saccadic suppression. When we make
a saccade, the visual system essentially shuts d own for the durati on of the
eye movement (visual activity is suspended in a sin-Ul1r way '"'hen we blink).
To be a bit more precise, the visual system d oes not s hut dmvn altogether.
Saccad k s uppression acts rnainly to suppress information carri ed by the
magnocellulaJ pathway.

FIGURE a 18 Th9 comparator. The

braJn stt1ds two copies (red lines) of
each commard to m ove the Q'ffiJIS.
One cop y {the mo tor signa.O g oes
to the extraocular muscles; the
othier (known as the effJQOC9 oopy
or 'corollary discharge signan goes
to the oomparator, w hic h oornparas
the image rnotion signal with U1e eye
rrotion signal and can compensate for
image changes pro duc9d by the eya
VI rrovement .
• Corollary disch.irge
.sign,11
._...
Image movement
si,gn.tl
J
Compar,1tor
To observesaccadic suppression, you need to find a mirror. As you look a t

yourself in the mirror, fi xate first o ne eye a nd the n the o ther. You M ll n o tice
tha t you d o n ot see the sa ocadic eye moveme n ts th.i t yo u must be rn.aking. If
you ' re concerned that the saccades might be too s1na.ll to see, find a friend to
help you _H ave this person s tan d in front of you and move his o r her fi x'1ti on
frorn one of your eyes to the other. You v.ill have no tro ubl e seeing your fri end ·s
saccades, even though you are quite blind to you r ow n .
Although saccadic suppression eliminates the smear of the m oving world
during a it seems as if \<Ve should s till be disturbed by the s udden
d isplaceme nt of the objects in fron t of us. In any case, no .suppression takes
place w he n \'\-'e execute smooth-pursuit eye m oveme nts, as in the earlier exercise
wi th the pencil. Althou g h ma ny scientis ts believe s uppression to be an active
process, others arg ue that s uppression is little more than maski ng
and that the magnoce llularpathway (see Cha pter 3) is no t s uppressed during
saccades (Castet, Jean jean, and Masson , 2001).
By sending ou t two copies of e'1 ch order to m ove the eyes, the motor sys-
tem is thought to solve the "problem" of w hy an object in motion m ay appear
stationary. O ne copy goes to the eye muscles; a nother (often referred to as the
''efference copy") goes to an area of the visual syste m tha t has been dubbed the
comparator (Figure 8 .18 ). 1}1e comparator can the n compensate for the image
dlanges cau sed by the eye m ovement, inhibiting any atternpts by other parts of
the visual syste m to interpret the changes as object rnotion. \.Yh en we jiggle the
comparator An area of the visual
eyeball with a fin ger1 no s ignal is sent from the eye muscles to the comparator system that receives cne copy o1the
(the eye mus d es are no t w hat m ove the eye), so the vis ual input is interpreted as cornrnand Issued by the mote< sys-
our world being rocked . (For inte rn.ctive de mons trati ons of the s imilarities a nd tem when the eyes move (the otrer
differe nces beho\,-een eye movemen ts and object movements, see W eb Activity copy goes to the ff;e muscles). The
8.5: Eye M ovements.) While a lot of work has been dedicated to the analysis comparator oornparoo the Image
motion slgnal the eye rnotlon
of e:ctraretinal s ignals during sm ooth pursuit, visual stability during s riccades sgnaJ am can comperoiate for the
re n41. ins a matter of active researc h (Cavanagh et al., 2010; Poletti, l istorti, and Image changes caused by the eye
Ruo:i, 2010), and attention seems to play an important role. movernent.

256 CHAPTER 8
FIGURE 8. 19 RflC QPti\1'9 field updat- (a ) RF.s oolative to a point o f fix,1tion

ing, Rgcepttve fiel ds are generally oon-
sidGr9d to be flX9d in SpaCG, relative
t o the fD<atlon point (a), so if the gyet::;
rnow to a rravot fixation point, the rec0p-
tl\'9 fields should shift In ·1ockstep" so
that they maintain the same positions
relatiV9 to th9 foV9a (/:;J). Shifting fixation
Ollus tratoo by the star) from the sculp-
ture on the OOw o f the boat (.::1) to the
roof of the boat f/J) results in a lockstep
shift Of the receptive fie ld3. (ellipses in
the figure}. Recwit 1NOrk suggests that
r.:iceptlvg fiek:l s of neurons In the frontal
f1Ye fi6'ds (FEF) shift toward the target
location (c}.
(b) RFs shift with th e point o f fixation
(c) RF.s trnns iently re!Tulp tov.·,ud the new point of fix.11ion
Updating the Neural Mechanisms for Eye Movement

Compensation
H ow does the brai n cornpensa te for eye m ove1nents in order to p reserve the
stability of our visual world ? Well over a century ago_, Hermann von Helmholtz
s uggested that sin ce the brain gen erates the neural signals fo r saccadic eye
movernents, it can perceph1ally compensa.te for them tby using the efference
copy described above). On e way this compensa tion could occur is through
the "remapping " of visual receptive fields. \Ve've discussed recepti ve fields
(RFs}-the region in space in w hich a v is ual s timulu s causes a n e m on to
change it's firing rate-at many points in the b ook (see Chapters 2-7). RPs a re
generally consid ered to be fixed in space_, relative to the fixa tio n point (star in
Figure 8.19a)_, so if the eyes move to a new fixation point_, the RFs (ellipses in
8231

Figure 8.1 9) sho uld s hift in "lockste p'' so that they maintain the same posi-
tions relative to the fovea (Figure 8 .19b). Shifting fi xation from the sculpture
on the bow (see Figu re8.19n) to the roof of th e boat (Figure 8J 9b) Tesults in
a lockstep shift of the RFs. However, the RFs of some neurons in the parietal
cortex actually shift to the new locations before the saccad e. Duha met Colby,
and Goldberg (1992) referred to this anticipatory shift as predicti ve remapping
or updating. A key ass umption abo ut th.is spatial-updating hypothesis is that
the n eural representation of the visual fie ld is rigidly translated just p rior to
the eye m ovem ent. H owever a very recent s h1dy su ggests a different view,
illus trated in Figure 8 .19c. Zirnsak and colleagues (201 4) found that the RFs
o f neurons in the frontal eye fie lds (FEF) shift transiently toward the tar get
location. In this vie\V, the RF shifts d o not p redict the retin.:'11 displacements
prod uced by saccades, but rather reflect the perceptual compression o f space
that occurs before s.accad es (Ross et al., 2001).
Development of Motion Perception

Sensitivity to visu al m otion d oes no t d evelop a ll a t once. Some aspects o f
m otion perception are a li·eady evi dent a t birth. For example , reflexive eye
m ovemen ts to m ovin g targets (OKN) are present in newborns (as lon g as the
targets suffidently large), and physiological s h1dies s hm"· tha t neurons in
V1 have adultlike sensitivity to visual d irection. On the other hand, sensitiv-
ity to global motion, which is thought to reflect processing in the MT, appears
to develop more s lowly, readlln g rna h.uity at about 3--4 years of age, while
sensi ti vity to nw tion-defin ed fom1 and biological motion takes even longer
(Freire et al., 2006; Parrish et a l., 2005).
The M an Who Couldn't See Motion

Recall what it's like to experience a motion aftereffoct. You stare at a
waterfall and then shift your gaze t o the cliff next to the falls. You know
that the reeks are not moving. and you can see that they aren't act u-
ally going anyv;here, yet you still experience an otherworldly :sense of
"disembodied" motion. (Review the phenomenon in Web Activity 8.4:
Motion Aflerettects if you haven't tr1ed ii for a while.)
Now imagine the opposite exi:etience: objects change position.
and you are fully aware of these location shifts but you experience no
perception of motion. As bizarre as it seems, this is exactly what hap-
pens in a rare neuropsychologlc.al diso rder known as aklnetopsla, de-
scribed in the fo llai.'Vi ng case repcrt {Horton ard Trobe,
A47-year-cid man rep<>ted soong streams of muruple. frozen Images
traJllng In the wake of m<Mng objects. As soon as motlm ceased. the
images collapsed into each other: He compared his vision to a scene lit by
a l ashing strobe, except that statlcrary elements were perceived normally.
helfl
In fact. If not11lng was In motlm 1"1d he pert!";tlY, still. h5 vision was
entirety rormal. The momenl anYlHlng mdVed, tiOWeVer. It lett a stream of
static coploo In Its path. For example, whHe rut for an evenlr(I strol. he saw
a pack of identical dogs lined up behh:l ris West Hghland ten1er. Driving
was Impossible because he was confused t1f multiple .snapshots of cars, aklnetopsia A rare neuropsycho-
streets, ard signs. M<Nlng l{)'lts were foRowed by a long comet trail. loglcal disorder In which tl1e affected
lndtvldual has no perception of motion.

258 CHAPTER 8
Another patient reportec that wh en she watched her own ann

m ovi r"Q . "passage of the limb would be reduplicatec by multip e, fuzzy
images, the way a cartoonist might draw motion. H
Not surprisingly, a!Onetopsia appears to be caused by disruptions

to visual area MT For the t\vo patients described here. the disruptions
were side effects of a prescription antidepressant drug, and their motion
p erceptio n problems o nce they were taken off the dn.1g. In
other cases , akinetopsia is brought on by direct trauma to area MT that
is due to stroke or elective brain surgery (e.g .. surgery to alleviate epi -
leptic seizures). Patients in the latter cat egory sometimes reQaln nonnal
m otion perception abilities several weeks after surgery, indicating that,
as in Newsorne's m onkeys, the human brain can som etimes rearrange
its connect ions so that different areas take over the MT's moti on-pro-
cessing fun::tlons.
Summary
Refer to the
Sensation and Perception
i.
mine where objects are going and when they' re likely to ge t there, and to
is
Ccmpanbn Website he1p us move through our env ironment witho ut being hit in the head by
sltes.slnauer.comlwolfe4e flying objects.
for aotMtm. essa\'-l. study 2. We can build a s imple motio n-detecting circuit by using linear filters that
delay and sum infomlation (and are fo llowed by nonlinearities).
questions. and other study aids.
3. Vl neurons vie\..• the world through a s mall window, leading to the well-
known aperture problem (tha t is, a Vl neuron is unable to teu , ..111.ich ele-
ments correspond V-·i th one another when an object moves through its
receptive field).
4. Strong phys iological and behavioral evidence sugges ts that the nUddle tem-
poral area (MT) is involved in the perception o f global motion.
5. Aftereffects for motion, like those for orientation or can provide
important insights into the nnderlying mechanisms of perception in
humans.
6. Luminance-defined (first-order) motion and contra.s t- or texture-defined
motion appear to be ana lyzed by separate systems.
7. TI1e brain has to figure out v"'11.ich retin..11 motion arises in the world, and
whid1 arises because of eye movements. Moreover, the brain must suppress
the motion signals generated by our eye movements, or the world will be
pretty "smeared."
8. Motion infomIBtion is critically important to us for navigating around our
world, avoiding imminent collision, and recog11.izing the movement of ani-
mals and people.

lq
8230336 Aml"la Appa

CHAPTER 9
Qns- Ccjci!o...,.c , M9m:xiBs Can? Wa1 w-2. 20:)2

Traditional oil technique on medium msdiJTJ tgxrure linen can\.o'aS
39 hch9s hcrizcntal x 40 inches vertical

Hearing: Physiology
and Psychoacoustics
IT IS OFTEN SAID THAT HUMANS ARE VISUAL ANIMALS . We are r eminded of the
importance of vision in our lives w hen ever we close o m eyes or av1i.-aken in the
aight, because so much of w hat we sense a nd know about o ur en viro nment
is s uddenly gone. In contrast, m ost people never get s u ch reminders of the
importance of heMing. Our ears are a lways open, and we can hear perfectly
well in the dark. \'\ihen we e nter a theater just before the movie starts,. we
d on' t n eed to wai t half an h our to be able to hear the soft s trains of the open-
ing theme. We can hear things when our nose is not pointed at the source of
the sound, and \'\'e can even he.:u around obstacles and corners. For better or
worse, we can often hear thro ugh barriers that ligh t canno t penetrate. For a ll
these ·t to hearing for granted.
Try to tmrtgme a world, though, where no thing makes a sound . You watch
a movie or online video, and the sound is always off. You cannot talk on
the phone. Deafness deprh-es you of the m os t fundamental of human abili-
ties: communicati on through speech. A person cries out, a nd you don' t help
because you cannot hear the cry. Without heari ng, the world is a more dan-
gerous ploce. You are relatively isolated by day and nearly totally isolated by
night when your sense of vision is corn promised by darkness.
The Func tion of Hearing

For the next three chapters, yo u will be hearing aU abou t heming. In th.is
chapter we cover the basics: the nature o f sound, the ana tom y and physiol-
ogy of the auditory system, and how \Ve perceive th e two fund am.ental sound
qualities-loudn ess and pitch. We condude this by looking a t some
of the ways in which hearing can be impaired and w hat we can do to avoid
and overcome these impairm ents. In Chapter 10 \""e will move on to discuss
some of the ways we use acoustic in fonnation to learn about our s urrolmd-
ings. Then, in Chap ter 111 we wi ll cover the h igher-level auditory fun ctions
that we use w hen we're listening to speech and music.
Many fundamental principles in h e(u-ingapply to all of the senses. However,
each sense developed at different periods in our evolutionary h istory and
in response to different en\oi ronmental cha llen ges. So, although you sh ould
find that much of what you've lean1ed thu s far wi ll help you to understand
hearing, you will also be impressed by ways biology h as p rovid ed some
very different (and very clever) solutions to the challen ges of sensing and
interpreting sound .

262 CHAPTER
(•) What Is Sound?

Sounds are crea ted when objects v ibrate. The vibrations of an object {the
Speaker
sound source) cause molecules in the object's s urrounding m edium (for
humans, us ually the Earth' s atmosp here) to vibrate as well. 1lUs vibration
causes press ure changes in the medium (F1gure 9.1 ). TI1ese p ressure changes
are best d escribed as waves, a nd they are s imilar to the waves on a p ond
caused by droppin g a rock into the wa ter. Water m olecu les d isp laced by
(b) the rock do not themselves trave l very far, but the pattern of d isplacement
Sinusoi.d."ll wave
will m ove ouhvard from the source until som ething (the sh ore, a boat, a
I s\.vimming duck, or anything else) gets in the way. Altho ugh the patterns
of p ond a nd sound waves d o n o t change as they s pread o ut, the initial
amount of pressu re change is dispersed over a large r and larger area as the
wave moves away, so the wave becom es less p rominent as it 1noves farther
from its SQ u rce.
Distance ---+-
So und waves tra vel at a par ti cular speed d ependin g on the medium,
FIG URE 9.1 The pattern of pressure moving faster through d enser s ubstances. For exampl e, the speed of sound
fluctuations of a sound stays thG sam9 through air is about 340 meters per second, depend in g on the humidity level
as t11e sound wcwe moves a.Nay from (sounds travel a bit faster on muggy d ays), but the speed of sound th rough
the source (a), but the amount of pres-
sure change decreasi;is 'v'Vith incr98Slng water is about 1500 meters per second. Light \Vave.s move throug h air almost
distance fb), a million times fas te r than sound waves d o. This is wh y you see lightnin g
before hearing thund er- abo ut 1 second pe r mile .
Basic Qualities of Sound Waves: Frequency and Amplitude

amplitude or Intensity The mag- As we' ve seen, sound waves tha t we hear are si mply fluctua ti ons in air
nitude of dlsplacementfinaease or pressure across tilne. The 1nagnitude of the pl'essure change in a sound
decrease) of a sound pressure W'a\16. wave-the difference the hig hest pressure a rea and the lowest
Arn plrtude Is perceived as loudness.
pressure a rea- is called the amplitude, or Intensity, o f the wave (Figure
frequency For srund. the nurnbef 9.2). Pressure fluctuations may be very cl ose togethe r or spread apart ove r
of times per second that a pattern of
pressure change repeiats. Frequency Is longer periods. For light waves, we usllillly d escribe the pattern of fluctua-
perceived as pitch. tions by measuring th e distance between peaks in the waves--that is, the
''wavelength.' 1 Sound vvaves also have Wll.velengths, b ut we desc ribe th eir
pa tte rns by no tin g how quickly the p ressure fluctuates; th is of fluc-
tua tion is knmlv'n as the frequency of the wave (see F'igiue 9.2). To see an
example of freque n cy, dang le a thread in fr ont of a s tereo. As the speaker
creates fluctuations in air p ressure, the thread waves back and forth. The
tempo of this waving is the threa d 's frequency. Sound wave frequencies are
1neasu red by these back-and-forth cycles, and the unit o f measure is ca lled
(•) (b)
l}I _ _ _JI G"''."

amplin1de '
i!l\Vl{llf\JV\1 AmpLtude d oubled;
frequency same
(tuning Y by the ti.ming fork' s vibration of movemen t original
,\.:S
fork)
!l)lli(V\ A 1\----F Amp litud.,.Sinewm

y V V V ...1..repre:sentatlo n of the
(c)
G reater li\mil\\J.!\v:{\v{\-}'-{\v{\---:'f .original,;
..\1:rrplit11de same as
pn:!s.su re above
\VdY...S
y)Ul
Wave length,
fn?quency
of mo•..ement H doubled
one cycle
\\'.l\'el.ength,
one cycle
FIGURE 9.2 A mplitude and frequ€<lcy. (a) Sound W81/f!!IS are d9scrlb9d by th9
freq uency and amplitude o f pressure fluc tuations. Changes in ampfitude (b) Md
frequency are shown for sine W3'/9S, the slmplQS.t kind of sound waw..

HEARING: PHYSIOLOGY AND PSYCHOACOUSTICS 263
FIG URE 9.3 Humans can hear frequMCies that range from about 20 to lffi
20,000 Hz across a very wide range o1 intensities, or sound preissure
(dB SPL, vvt1ere dB= IJJ
a hertz (Hz), w here 1 cycle pel' second equals 1 Hz. For example, the . 110
pressure in a 500-Hz wave goes from its highest point d mvn to its
lowest point and back up to its highest point 500 times every secon d .
Ju.st as the amplitude and wavelength of light waves correspond to
.. 00
perceptual qualities in vision (brightness and color, respectively), the • 70
amplitude and frequency of sOlm d waves are doselyrel.:1ted to auditory

characteristics. Amplitude is associated \.....-ith the perceptual quality of
•
0. ffi
loudness: the more intense a sot.md wave is, the louder it will sotmd . § .)J
Frequency is associated \.vi th pitch: low-frequency sotmds correspond
JI
to low pitd1es {e.g., low notes played by a tuba), and high-frequency
10
sounds correspond to high pitches (e.g., the high notes from a piccolo). \ Ve
will have much more to say about the relationships behveen amplitude
and loudness and between frequency and pitch later in this d1apter. -100
20 XlO 20,0CO
In Chapter 2, you learned how visible light makes up onl y a s ma11 Fn>quency {HzJ
portion of the much broade r range of electromagnetic energy; simi-
larly, human heari ng uses a Limited range of the frequencies presen t
in enviromnental sounds. If you are relatively young and you've hertz (Hz) A unit of measure for frequency.
been. care.fol about your exposu re to loud sounds, you may be able One hertz equals one cyde per sooond.
to detect sounds that vary from about 20 to 20,000 H z (Figure 9.3). loudness The psychoklglcal aspect of sound
Some animals sound s that have lower and higher frequencies relatoo to percelvoo Intensity (amplitude).
than those heard by hw11ans. In general, i<'lrger anirn.a ls are better at pitc h The psychological OEj'.)OOt of sound
hearing low frequencies, and smaller animals are better at using h igh relatoo mainly to the Fundamental frequency.
frequencies. Elephants hear vibra tions at very low frequencies that decibel (dB) A unit o f measure for the phy\jcal
help then1 detect the presence of large animals, such as other elephants. Intensity of soun::l. Decibels define the difference
Dogs be called with whistles that emit sounds Lit frequencies too between rwo sounds as the ratio betvJeen tw'O
high for hwnans to hear, and the sonar system s used by some bats soum pressures. Eact1 i O: i ro.Jnd pressure
ratio equals 20 dB, and a 100:1 ratio equals 40
use sotmd frequencies above 60,CXXJ Hz. dB.
Humans hem across a very \'I.ride range of sound intensities. The
intensity ratio behveen the fain test sound humans can d e tect and
the loudest sounds that do not cause serious damage to hmn an ears TABLE 9.1
is more than on e to a million. To d escribe differences in ampli tud e Decibel levels that correspond to differ-
across s ud1 a broad range, sound levels are measured on a logaritlunic ent sound pressure ratios
scale usin g units called decibels (dB). Decibels define the difference Ratio relative to
beh veen hvo sounds in tem1s of the ratio betv.'eei1 sotmd press ures.
1
0.0002 dyne/cm' (p,) dB
Each 10:1 sound p ressure ratio is eq ual to 20 dB, so a 100:1 ratio is
0.0
equa l to 40 dB. The equation for defining decibels is
6.0
20 log(p/ p,) 12.0
The variable p corresponds to the pressure (intensity) of the sotmd 16.0
being described. 11,e constant term Po is a reference pressure and 10 20.0
is typically defined in auditory research contexts to be 0.0002 dyne 20 26.0
per square centimeter (dyne/an 2), and Levels are defined as dB SPL
(sound pressure level). This level (0.0002 d yne / cm 2) is d ose to the 50 54.0
minimum pressme that can be detected at freq uencies for which hea r- 100 40,0
ing is m ost sensitive, and decibel val ues g reater tl1an zero describe 1000 60.0
the ratio between a sound being measured an d 0J)002 dyne/ an 2• 10,000 80.0
The range of hmnan hearing extends from 0 to over 120 dB SPL,and 100,000 100.0
as s hm'\o'n in Table 9 .1, th is decibel range corresponds to a ratio of
1.000,000 120.0
grea ter than 1,000,000:1.

264 CHAPTER 9
Leaves rustling Library EhWne;s offic.e Heavy truck Jad<lu:unmer Jet takeoff
ofheoring 0 10 20
FIGURE 9. 4 S ounds that 'W0 hear In

our daily gn\lironrmints vary gr9atly in
3-0 40 50 60 70
Loudne!S (dB}
9J 90 100
I .L
110 120 13-0 140 of paln
int911slty. Using a referen ce value s uch ilS p0 is common to many m easuring .sys tems.
For examp le, O°C is d efine d as the temp erature a t w hich water freezes, and
l OO°C is the te mperature a t whidi water boils. H the p ressure o f the sound
that you're measuring l.p) is equal to 0.0002 dyne/cm ', then 20 log(l ).
Because the log of 1 is zero, a sound p ressure that low would be equal toO dB
SPL, but 0 d BSPL is not silence. Sounds with a mplitudes e \ 'ffi smaller than Po
have negative decibel levels, just as substan ces colder than the freezing p oint
o f water have negative centigrad e temperatures.
An important thing to remember about loga rithmic scales such as decibels is
Iha t relatively small decibel changes am correspond to large physical changes.
For example, an increase of 6 dB corresponds to a d oubling of the am ount of
p ressure. Flgure 9 .4 sh cnvs the d ecibel levels of some common sound .sou rces.
Sine Waves and Complex Sounds

In Chapter 1 you were introd uced to one of the .simplest kinds of .sounds: a
sine wave, which is often called a pure tone. All sound s, even as complex as
the sounds p roduced by musical ins truments, human speech, and city traf-
fic, can be described as a combination of sine waves. (See Web Activity 9.1:
sine wave or pure tone The W@l0· What We Hear.)
foITTI for which variation as a furctlon Complex sounds are best d escribed in a spectrum (plural s-pectrn) that
o f time Is a sine function. displays h ow much energy, or amplitude, is present at multiple frequen cies,as
spec1rum A repreoontatkln of the shown in F1gure 9.5. Many common soWlds have hannonlc spectra, iJlustrared
relative energy (lntenslt)? present at in Figure 9 .6. TI1ese are typically caused by a s imple vibra ting source, su ch as
each freq uency. the s tring of a guitar or the reed of a saxophone. Ead1 freque_ncy component in
harmonic spectrum The spectrum su d1 a sound is called a "harmonic'' The first h armon ic, c:Uled the fundamental
of a complex sound In which energy Is frequency, is the lo west-frequency component of the sound. AH the o ther
a! Integer multiples of the fundamental harmonics have freq uencies that are integer multiples of the fw,d am ental.
frequency.
TI1e shape of the spectrum (spectral shape) is one of the m ost important
fundamental frequency The qualities that distinguish different sounds. The p roperties of sound sou rces
ffl"IPO"El(ll of a determine the spectral sh apes of sounds, and these sh apes help us identify
complex periodic sound.
sound sources. For example, Fig ure 9.6 illustrates spectra from three musical
timbre The ps)ffiologlcal sensa- instruments. Each instnm1ent is producin g a n ote with the same fundamental
tlon by which a Mstener can judge that
&equency (262 H z, which cor responds to the n o te C 4, or middle C) and the
two sounds with the Slm0 loudness
and pitch are dlsslml..-. Timbre quality same harmonics (524 Hz, 786 Hz, 1048 H z, and so on ). H owever, the shapes
Is ccnveyed by harmcnlcs and other of the spectra {the pattern o f amplitudes for each harmonic) vary. llmbre
high frequercloo. (p ron ounced ''tamber," like "amber") is a term used to d escribe the qua li ty of

(a) Waveform (b) Spo<trum
A E\/\/\/\;, JI
A=dB
-1
D1 D Tune (ms)
soco
Frequency (H z)
10,llXJ
FIG URE 9. 5 A spectrum dSplays the amplitude for each

pr'11Sent In a sound wave. Each signal Is sho'M1 as a
waveform (a) and as a spectrum fb),
a sound that depends, in part, on the relative energy levels

of h.:1rmonic compon ents. !'me IDn• (262 117, C4)
We will return to ha l'mon.ics, timbre, a nd other
of complex sound s in Ch apter 10. In this chapter we'll s tick
mainly to the s tory of how the audito ry syste m processes
si mple sound s s ud1 as si ne wave tone,s.
------+-------
Basic Structure of the
Mammalian Auditory System
Now that you know what sound is, we ca n examine how
sounds are detected and recognized by the auditory system .
The sense of hearing has evolved over rnillions of years to be
able to do some pretty amazing things. We are about to de-
scribe quite a few anatomical struch ues that are essential to
:ffi
Trombone
111111 ,,,
Lmderstanding how seq uen ces of tiny air pres5ure changes are
htmed into meaningful sound percep tion. TI1e discussion m ay
occasiona ll y be a bit confusin g, but if you consult the figures
:tl111 I W lli q
and Web Activity 9.2: Structure of the Auditory System
often, you will soon know tQe pa rts 1md how th fit ex Piano
FIG URE 9.6 Harmonic sounds with the SEmQ fun<:iM)Qnt.al

frequEf'lo; can sound different because amplitudes of lndi\liduat
frequGrncy cornp::ments are restitlng 1n different spec-
tral shapes. For example . d ifferent music al instruments playing
1111
the same note (the same frndarrumtal frQQUQncy. abbrnvi.at9d I) 0 0000 10,llXJ
sound diff.erent. C,. ::::: middle C. Frequency (H z)

266 CHAPTER 9
8230336 Amma Appa
Outer Ear
So unds are firs t coll ected fro m the en vironment by the pinna (plura l pinuae),
the curly s tructure on the s ide of the head that we typically ca ll an ear. Only
ma rnmals have pinnae, an d they vary wildly in shape a nd size across species
and vary less dramatically across individuals within species (Figure 9.7). As
we will see in Chapter 10, the s hape of the pinnae plays an important role in
our ability to loca lize sound sources.
Sound vvaves are funn eled by the pinna into and th rou gh the ear canal,
pinna The outer, funnB-like part of which extends about 25 millim e ters (nm1 ) into the head (Figure 9.8). The
the ear. length and s hape of the ear can al e nhance sound frequ encies bern.·een about
ear canal The canal that conducts 2000and 6000 Hz, but the m ain purp ose of the ca n.al is to protect thestrudure
scund vibrations fran the pinna to the at its e nd, the tympanlc membrane (eardnlm), fr om damage. The tympanic
zympanlc membrane and prevents rne rnbran e is a thin sheet of skin tha t nloves in a nd out in response to the
damage to the t)mpanl: membrane. pressure changes of sound waves.
tympanlc membrane The ear- It is a cornm011 myth that puncturing your eardrum will leave yo u deaf.
drum; a thin sheet o f skin at the end While a ruptured eardntm can be excruciating, in most cases a damrtged tym-
of the outer ear c anal . The t)mpanlc panic me mbrane w ill heal itself, just as other parts o f the s kin do. However, it
membran e vibrates In respcnse to
scund.
is p ossible to da mage the ty mpanic membrane beyond repa ir1 so it's a good
idea to fo llow yorn m other 's advice to n ot s tick things in your ear.
outer ear The externaJ &YJum-gath-
erlng pcrtlon of the ear, consisting o f
the pinna and the ear canal.
Middle Ear
Toge the r, the piima and ear canf'l make up a division of the auditory system
mld<le ear An air-filled chamber
the middle bcnes, or called the outer ear (see Figure 9.8). The tympank me mbrane is the borde r be-
osslcles. The mlddle ear conveys ard tween the outer ear a nd the middle ear, which consists of three tiny bones, the
amplifies vibration frcm the tympanlc osslcles, that amplify sound waves. TI\e first ossicle, the malleus, is connected
membrane to the oval window. to the ty1npanic membrane on on e sid e and to the second ossicle, the lncus,
osslcle Any of three tiny bones of on the othe r. ll\e incus is connected in turn to the third ossicle, the stapes1
tha middle ear: rnalleus , lrcus. and w hfch transmits the v ibrati ons of sound waves to the oval window1 another
stapes. me mbrane, \•vhich forms the borde r behveen the middl e ear and the Inner ear.
malleus One of the three ossldes. TI1e ossicles are the s mallest bones in the htm1<.'1 l1 body, and they amplify
The rnalleus receives Vibration from sound vibrations in two ways. First, the joints between the bones are hinged in
the tympanlc membrane and Is
ways that make the m work like levers: a modest a mount of energy on one side
attached to the lncus.
o f the fulcrum ijoint) becom es l::irger on the o ther. Titls lever actio n increases
incus The mlddle of the three the am ount of press ure change by about a third. TI1e second way the ossides
ooslc les, connecting the malleus and
the stapes. increase the e nergy transmitted to the inner ea r is by concentrating ene rgy
from a la rger to a s malle r surface area. The tympanic membrane, w hich m oves
stapes one of the three ooslc les.
Connecte::1 to the lncus on one end, the malleus, is a bout 18 times as la rge as the oval \'Vi nd. ow, w hich is m oved by
the stapes presses against the oval the stapes (see Figure 9.8). Therefore, pressure on the oval wind ow is magni-
window of the cochlea en the other fied 18 times relative to the pressure on the ty mpanic membra ne. Thi s is the
erd. same principle that makes snowshoes keep feet on top of the s now and makes
oval window The flexible open- s tile tto heels a danger to wood fl oors (think of the tympanic membrane as the
ing to the oochlea through w l1loh the heel of the foot and the oval vvindow as the tip of the s til etto)_
stapes transmits vlbratloo to the fiutl Amplification provided by the ossides is essential to our ability to hear
Inside. fai nt sotmds because the inner ear, as we will see in a moment, is made up
Inner ear A hollow cavity In the tem- of a collection of fluid-filled cha mbers . Becau se it takes more ene rgy to move
pcral bone of the skull, and the struc- liquid tha n it d OES to move air, this fluid creates a mismatch. If sound '""'aves
tures v.;thln ths cavil)!; the oochlea
and the semicircular canals of the were tran smitted to the ova l window directly, ma ny would simply bounce
vestibular system canas. back w ithou t movin g the oval window and the liquid behind it a t alL
tensor tympani The muscle Ossicles p lay an important role for loud sounds too. 111e middle ear has
attac hed to the m<"leus; tensing the two mu scles: the tensor tympani (attached to the mallet.is) and the stapedius
tensor tympani decreases Vlbrat.lcn. (att.:td\ed to the stapes) (see Figure 9.8). As might be expected for the s mall-
stapedlus The muscle attached est bones in the body, the tenso r tympa ni and the s tap edius are the smallest
to the stapes; tensing the stapedlus muscles in the body. Their main purpose is to tense vvhen sounds are very
decreases vlbratro. loud . They restrict m ovem e nt of the ossid es and thus muffle pressure changes

Malleus lncus
st.1p es
incn·al
win dow
I
I
?\fi ddle ear
FIGURE 9.6 Structur13s of th€l that thetyrnpan\c rnernbrane h.:s

about 18 times as much surface area as the ova.I window beneath the sto.pes.

268 CHAPTER ()
acoustic reflex A retlex that pro- that m ight be large enou gh to damage the d elicate s tructures in the inner ear.
tects the ear from Intense ooums. via Unfortunately, this acoustic reflex follows the onset of loud sounds by about
contraction of the staped ius and ten-
scr tympani muscloo.
one-fifth of a second. So, while JT1uscles he lp in en\·i ronments th..1t are loud for
sustained periods, the acoustic reflexca1motprotect agains t abmpt loud sotmds,
cochlea A spiral structure of the su ch as the firin g o f a gun. Muscle:; of the middl e ear are also tensed during
Inner ear containing the organ of Conl.
s,,..,·allowing, talking, and genera l body rnovement, helping to keep the auditory
tympanlc canal One of three system from being overwhelrned by sounds generated by our own bodies.
nul:l-llled passages In 1110 cochlea.
The tympanlc canal extends from
the round window at tha base of the Inner Ear
cochlea to the hellootrerna at the apex. The itme r ear is an impressive feat of evolution. ft is here that the minu te
Also called sea/a t}mPaf>I. changes in sound pressu re available in the em·i ronme nt are trans lated into
vestibular canal One of three nuid- n eurn. l signals tha t inform the listener about the world. TI1e hmction of the in-
ftlled passages In the occhlea. The ner e ar \'\rith respect to sound waves is ro ughly analogous to that of the retina
vestibular canal extends trom the oval with respect to light waves in vision: it translates the information carried by
window at the base of the oochlea the waves into neural sign als.
to t11e helioctrema at the apex. Also
called sea/a vest/bu#.
COCHLEAR CANALS ANO MEMBRANES The major s tructure of the inner ear is
midcle canal 01e of three ftuld- the cochlea (&om the Greekkoclrlos, usnail ''), a tiny coiled s tntcture embedd ed
ftlled passages In the occhlea_The
middle canal Is sandwiched bet"""n in the temporal bone of the s kull (see Figme 9.8). Rolled up, the cochlea is the
the tympanlc and vootlbular canals and size of a babypea,about4mm in diameter in humans. Uncoiled , it would bea
contains the cochlear partition. Also hibe almost ten times as long-ubout 35 nun or l.4 ind1es. TI1ecochlea is filled
called sea/a mecia. with watery fluids in th ree parallel cana ls (Figure 9 .9): the tympanlc canal
hellcotrema The opening that con- (or scala tym pani), the vestibular canal (or sea.la vesti buli), and the middle
nects the tympanlc and vestibular canal {or .scab. media). The tympank and vestibular canals are connected by a
c anals at the apex of the cochlea
s mall open ing, the heUcotrema, and these two rnnals are effectively v.Tapped
Relssner'S membrane A thin ru"Ou nd the m..iddle canal. Think of the tympanic and ves tibular canals :lS one
sheath of tissue separating the vestib- long skinny balloon (the kind d owns use to m ake ha ts and animals), blmvn
ular and mld<lle canals In 1110 cochlea.
up and folded back on itself. TI1e nUddlecan.al is ano ther long balloon that is
basilar membrane A plate of fibers sandwiched, lengthwise, between the hvo ha lves of the first balloon.
that forms the base of the cochlear
panltlon and separates the middle and 11-ie three c.."lnals of the cochlea are separated by two membranes (see Figu re
tyrnpanlc c anals In the ccchlea 9.9): Retssner•s membrane, beh-¥een the vesti bula r c..m al and the middle canal;
and the basilar membrane, ben...·een the middle canal and th e tymp;,mic canal.
cochlear partlUon The combined
basilar membrane, tectala1mem- Strictly speaking, the basilar membmne is not really a m ernbrane, because it
brane, and organ of conl. which are is not a thin., pliable sheet like the tympank membrane, the oval window, or
together resp::nslble for the trara- Reissner' s membra ne. Instead, it is a pla te m ade up of s tiff fibers. The basilar
ductlon of sound wavoo Into neural me mbrane fo rms th e base of the cochlear partition, a co mplex s tructure
sO;Jnels. through w hich so und waves are trans duced into neural signals.
rol..lld window A soft area of tissue Vibra tions transmitted throu gh the tympa nic membrane a nd middle-ear
at the base of the tympanlc canal that bones cause the s tapes to pus h a nd pull the flexible oval wind ow in and
releases excess pressure remaining
frcm extremely Intense sourds. out of the vestibular canal a t the base of the cochlea. TI1is m ovement of the
oval window causes waves o f pressu re called " trave ling waves,"
to flow through the fluid in the vestibular canal, in much the sa me way that
the tnembrane of a loud speaker m oves air to create solmd waves. Because
the cochlea is a dosed system, ii reSsure nnot spread ou t in all
directions. Ins tead , a displacernent, or " bu lge," for ms in the vestibular canal
and ex tends fr om the base of the cochlea d own to the apex (look ah ead to
Figure 9.12). lf sounds are extreme ly intense, any pressure th at remi:!ins a t the
apex passes through the heli..cotrema and back to the cochlear base through
the tympaniccana l, w here it is relieved by s tretching yet ano ther membrane,
call ed the round window (see Figure 9.9).
Beca use the vestibular and tyrnpanic canals are wrapped tightly arow1d
the middle canat wh en the vestibular canal bu lges out it puts p ressure on the
middle canal. 111is pressure has the effect of dis placing the basilar membrane

HEARING: R-IYSIOLOGY AND PSYCHOACOUSTICS 269
Tectoria l
m embrane
Audit01y
Spll<>l -
gang lion
H eliootrema
Tectorial
StereOCJh a o f StereOcillii of
inner h air cells ou ter h,1ircells
8230336 Amma APP""
I
Auditory nen-e
F IGURE 9.9 Theo The upper left illustration Is from the viewpoint of facing a
perscn. The rnm&mg Il lustrations show cross S9Ctions of the cochlea at suoct;1ssivety
greater levels of detail. Not e the thrM canals of the ood'llea: vestibular, middle, and
tymp anic (u pPQr right}. Wh91'1 vibrations Q""rter the cochlea, th9 t9ctorial tnt;1mbrane
shears across the organ of Corti Oowe- right). The photomicrograph (lower left) s hows
real hair cells: the single row of inner hair cells and three rO\IVS o f outef h air cells
(rightl. (Micrograph from Keese and Kardon, 1Q79.)
organ of Corti A strLIOture rn the
basilar membrane of the cochlea that
is composed of hair cells and den-
(which, recall, lles a t the bottom of the mid d le cana l), pushing d ovvn wh en dntoo of aud rtory ne<Ve fibern.
the vestibular-canal bulge is created . hair eel I Any oell that has stereocilla
for transduclng mechanical move-
ment In the Inner ear Into neural actlvlty
THE ORGAN OF CORTI Movemen ts of the cochlear partition a re translated
sent to the brain: some hai r eels also
into neural signals by s tructu res in the organ of Corti, which exten ds alon g recave Inputs frC111 t he brain.
the top of the basila r membrane (see Figu re 9.9) . The organ of Corti is rnade
audltoty nerve A collectlon of neu-
up of a scaffold of cells that s np p or t spedali zed neurons ca lled hair cells_, and rons that c.c;wey u11crmat1on from h91r
d end ri tes of auditory nerve fibers that term in ate a t the base of hai r cells. Hai r cells In the cochlea to (afferent) and
cells in each human ear are arranged in fou l' rows that rnn d o\-vn the length of fran (efferent:i t he brain stem.

270 CHAPTER 9
l=IG URE 9.10 \Ml9r1 'Jibratk:n causgs a dlsplacement along the Sound-induced vibration
cochlear partition (see Figure 9.9 , low&r right}, the tectorial mern-
bran9 and hair ceHs m O\IQ In opposlt9 dirscti0ns (sheaQ, ard the
deflgctlon o f s tereocllla during this action riesults in the release o f
neurotransmitters.
Resting position
".:': \!{1 1..z..1______________ -
Downward phase.
the basilar membrane: o ne row of about 3500 inner hair cells and three rows
with a total of about 10,,500 outer hair cells.
Inner and outer hair cells provide the fou ndations for minuscule h airlike
bristles called stereocllla (singular stereocflium). On an inner hair cell, stereo-
cilia are arranged as if posing for a group photo, in several nearly straight
rows with the shorter stereocilia in front and the taller ones peering over their
s houlders in the back. On an outer hair cell, stereocilia stand in rows that fonu
the shap e of a V or W (see Figure 9.9).
TI1e tector1al membrane extends atop the organ of Corti. Like the basilar
membrane, it isn' t really a membrane. Rather, it is a gelatinous flap that is at-
tached on on e end and floats above the outer hair cells o n the other end. Taller
s tereodlia of o uter hair cells are embedded in the tectorial membrane, and the
stereocilia. of inner hair cells are nestled against it BecaUSle the tectorial membrane
is attached on only one end, it shears across the width of the cochlear partition
w henever the partition moves up and down. This shearing m otion causes the
srereocilia of both inner and outer hair cells to bend back and forth (Figure 9 .1 0).
INNER AND OUTER HAIR CELLS like photoreceptors in the retina, hair cells
are specialized neurons that tran.s duce one kind of energy (in this case, sound
pressure) into another fonn of energy (neural firing). Hair cells in the vestibular
organs a lso report head movements to the b rain, as you will learn in Chap ter
12. Deflection of a hair cell's stereocilia ca.uses a change in voltage p otential
that initiates the release of newotransmitters, which in turn encourages firing
by auditory nerve fibers that have dendritic "Y""P"" on hair cells (Hudspe th,
2013). H owever, it is the differences between photoreceptors and hair cells
that are the m ost interesting.
stereoclllum Arry of the hairlike 'While the retina has a lmost 100 million photoreceptors, the cochlea has
extensions on the tips of hair cells In only about 14,000 hair cells. Although outnumbered, stereocilia of hair cells
the cochlea that. v.t-ren fleXed , Initiate blow away the competition when it comes to speed and sensitivity. A:5 we11
the release of neurotransmitters.
see in Chapter 10, hair c.ells must be capable of responding so fas t that we can
tectorlal membrane A gelat1nous detect time differences of as little as 10 millionths of a. second-lOmicroseconds
structure, attached on one end, that
extends Into the middle canal of the (f.15)-in order to know the direction from whid1 a sound arrives. In contrast,
ear. ftoatlng above Inner hair cells ard wh en we watch a. m ovie, pictures shown at 24 frames pe.r second appear
touching outer hair ceUs. continuous to the visual system. H air celJs are n ot only extren1ely fast, but

also extreme ly sens itive. It m ay take 30 minu tes for our eyes to folly a d just to tip link A tiny I lament that stretches
a d ark thea ter, but o ur ears are always ready for the slightes t sound. from the tip of • stereod llum to the
Recall that the shor test stereocilia are in front o f slightly talle.r stereo::ilia side of Its neighbor.
tha t a re in front of still-taUe r stereocilla {Figure 9 .11 ). Each stereocilium is
connected to its neighbor by a tiny filament called a tip link, so the stereocilia
connected by tip links bend togethe r as a set when deflecred by the shearing
mo tion of the tectorial membrane. When a s tereocilium deflects, the tip link
pulls on the taller stereociliu m in a way that opens an ion pore somewhat like
opening a gate for just a tiny fraction of a second . This actionpennitspotassiurn
ions (K... ) to flow rapidly into the hair cell, causing rapid depolarization (see
Figure 9.ll b and c). In tum, depolarizatio n leads to a rapid influ x of calcium
ions (Cai..) and initiation of the release of n eurotran smitters from the base
of the hair cell to s timulate d endrites o f the aud itory nerve (Fettiplace a nd
H ackney, 2006; Hudspeth, 1997).
The opening o f ion pores that resul ts from the d irect connection behveen
stereocilfa via tip links is known as mech an oelectrical tran sdu ction (MET), FIG URE9. 11 StArooci lia regulatB
thg flO'Vtl of Ions Into and out of hair
w hich is responsible fo r both the extreme speed a nd the sensitivity of hair celli..
cells. (a) This photom icrograph shows
Unlike the case in vision, dep olariza tion in h earing d oes not await a cascade the tip Hnks that conn9ci
of b iochemical processes s uch as those in photoactivation. Mechan oelectrical the tip of each shorter ster90Cilium to
transduction is also extremely sensi tive: ion p ores op en deflection is as its t aller neighbor. f.P, c) Bending the
little as 1 nanometer (nm), rou gh ly the diameter of a s ingle :atom. This deflec- stereocilia atop a hair cell opens th9
tion is equiva lent to only h alf a n inch of movemen t of a blade of grass close Ion pores, p ermitting a rapid influx o f
potassi um ions (K'1 into the hair 001,
to three football fields (350 yards) away. and this depolarization opens chan-
The fir ing o f the au di tory nerve fi bers finally com pletes the p rocess of nels that allO'Vt/ calcium b n s (Ca2+-) to
translating sotmd waves into patterns of neural activ ity. ooter the base of th9 hair cell, causing
the release of neurotransmitter Into thB
S)'T'lapse bstWQQO the hair C911 and an
afferent auditory nerve fiber.
(•) (b) (c)
.. <>o
82
.
c.." •
Vesicles
C®@
Ca 2'1"

272 CHAPTER
Here's a brief summary of the wh ole process: An air pressure wave is fun-
neled by the pinna through the auditory to the ty1npanic membrane,
which \-;brates back and for th in tirne wi th the sound wave. The tympanic
mem.brane vibrates the mal leus, which vibrates the incus, which vibrates the
st<tpes, w hich pushes and pulls on the oval wind ow. l11e movement of the oval
window causes press ure bulges to move down the length of the vestibul ar
canal, and these bulges in the ves tibular canal displace the middle can al up
an d do,,...-n . This up-an d -do\o\T\ m o tion forces the tectorial membrane to sh ear
across the organ o f Corti, m ovin g the s te reocilla atop hair cells back and
forth. The pivoting of the s tereocilia initiates rapid d epolarization (followed
by equally fust hyperpolariz.ation) tha t results in spurts of n eurotransmitter
released into sy11apses ben.veen the hair cells and dendrites of auditory nerve
fibe rs. These neurotransmitters initiate actio n potentials in the auditory nerve
fi bers tha t a re corried to the brain. Ano is tcim
CODING OF AMPLITUDE AND FREQUENCY IN THE COCHLEA Now let's retum

to see h ow the two fundamental characteristics of sound waves--amplitude
and frequency- are en coded by the cochlea.
lf the amplitude of a sound wave is increased, the tympa nic membrane
and oval window move farther in a nd out with pressure fluchtation. The
resul t is that the bulge U1 the vestibular cana l becomes bigger, which m uses
the cochlear partition to move farther up and dovvn, \-vhich causes the tecto rial
membrane to shear across the organ of Corti more forcefully, w hich causes the
h air cells to pivot farther back and forth, which 1.."""auses m ore neurotransmitters
to be released, which causes the aud itory nerve fibers to fire action potentials
more quickly. Tims, sound wave amplitude is conveyed in much the sarn e
way as light wave arnplitude: the larger the ampli tude, the higher the firing
rate of the neurons that commu nica te with the brain. (We w ill discuss some
compli ca tions of this simple explanation later in the chapter.)
Codin g fo r fre quency isa bi t trickier. Earlier we said that the cochlear pa rti-
tion is dispbred up and d o\o\11 in different p laces along the bas ilar membrane
that corresp ond to d ifferent freq ue ncies in the .sound wave. This .sta te ment
is true as for as it goes, but it does n ot tell the '"·hole because different
parts of the cochlear partition are displaced to different degrees by different
sotmd wave frequencies. High frequencies cause the largest displacements
do.ser to the oval window, near the base o f the cochlea. LO\ver frequencies
cause d isplacem ents farther away and nearer the apex. ln other words, dif-
feren t places on the cochlea are ''tuned" to different frequencies. TI1is tuning
is knO\v:n as the place code for sound freq uency.
Cochlear tuning to frequency is ca used, in large part, by diffe ren ces in
the structu re of the basilar membrane along the length o f the cochlea (Figure
9.12). The cochlea narrows from base to apex, but the basilar membrane insid e
actually widens toward the apex. In addition, the basilar membrane is th ick at
the base and becomes th inner as it widens. \ Vhile the basilar membrane gets
thinner a nd wider alon g its length, the cochlea separates frequencies like an
place code Tunin;i of different parts acoustic prism. Higher freq uencies affect the narrower, stiffer regions of the
of the cochlea to different frequencies, basilar m embrane near the base m ore1 and lower frequ encies cause greater
In which lnfarnatlon about the particu- d isplacements in the wider, m ore flexible regions near the apex.
lar trE>:1001cy of an inccmlng sound
W<Ne Is coded by the place along the
In addi tion to th is passive, structura l way of being tuned to freque n cy,
cochlear partition that has the greatest other processes actively s ha rpen hming. Remember that there are two dif-
rnechanlcal displacement. fere nt types of hair cells: inner an d oute r. Over 900/o of the afferent fibers
afferent fiber A neuron that carnes in the auditory nerve-fibers that take information to the brai n-synapse
sensory lnfcrmatlon to the central ner - on the 3500 inn er hair cells (10-30 auditory nerve fibers listen to each iimer
VOUS system. hair cell ). If the inner hair ceJJ s are con veyi n g a lm ost a ll the information

FIG URE 9.12 Th9 c ochl&a.is lika

an ac oustic prism in that its sensitivity
spread s ac ross different sound fre-
qu0nc ilils along its IQ'"lgth. Th9 narrow9r
end of the basilar membrane to ward
th9 b ase is stiffer and most sensHive
sensitive to l()!Nef frequencies. Here

High frequencies Low frequencies the cochlea is Illustrated as if it were
d.is.ploce basilar displace basilar
membrane in men-.brane in aJ."E'X unoolled (8), and the shapQs of the
b,15eofcochl.e,1 . of cochlea. traveling waves for different frequencies
L
o f vibration are shown (b) .
Dimtion ol \ " Unrolled"
sound movement ---+-- c\ lea
Cochle,u base Coch lear a}'EX
:
Relative ampUtude
of m ovemen t (J.lm)
0 --- ------ - - - -
3
0 10
I
20
----
I
30
Distance from st<1pes (mm)
about smmd waves to the brain, then '\.vha t d o the 10,500 outer hair ce lls d o?
It turns o ut th at m ost of the nerve fibers tha t syn a p se w ith the outer hair
cells a re efferent fibers, con veying in fo nna tion /rom the brain. \Vhen these efferent fiber A neuron that carries
efferent fibe rs become active, outer hair cells on w hich they s ynapse become Information from the central nervous
phys ica Uy longer, maki ng the nearby cochlear p arti tion stiffe r (Fettiplace system to the periphery.

274 CHAPTER g.
FIGURE 9. 13 Threshold tuning C Ul"VQS for six auditory ngrve fibQrs, 9.aeh tunOO to
a d iffEfent frequency. curves define the lowest intensify necessary fcr the neurcn to
fire above its spont.ano90us rate at each frequency. The characteris tic frequency for
each of these six nerve fibers is at the lo\1>19St point o f the tuning curve.
BOt
and Hac kney, 2006). By making some parts of the cochlear partition s tiffer
fi f'°\J
th an othe r par ts, o uter hair ce lls m ake the cochlea m ore sen sitive and more
s ha rply tuned to particular frequencies .
r·\J
The Auditory Nerve
0 Now that we've covered the mechanics of how the auditory system trans lates
air pres.sure cha nges into a udito ry nerve firing, let'.s discu ss w h at we know
abo ut a uditory n erve (AN) fibers. Mo re s pecifically, we'll consider the type
of information con veyed by afferent AN fibers from the cochlea to the b rain.
1 Re me 1nber that sounds wi th different frequencies disp lace d ifferent

region s o f the coch lear p arti tion . lnner hair ce lls, vvhich pro\.;d e most of
fh e informa tion to the brain via AN fibers, are lined up s ingle file along the
length o f the basilM m embrane. Put these h<Vo p ieces of information toge ther,
!
o and we can infer that the responses of individua l AN fibe rs to differen t fre-
quencies sho uld be related to their place a long the cochlea r partition . Su.re
e nough, w hen scientists record from individ u(1l AN fibers in a nim..ils, they
find that d ifferent fibers selecti vely respond to different sound frequencies.
This freq uency se lecti vity is d earest w hen so tmds arevety faint; a t very
'
low inten sity levels (even less tha n 0 dB), a n AN fib e r will increme firing
to onl y a very restricted ra nge o f frequ encies. Figure 9 .1 3 shO\vs threshold
tuning c urves fb'rsevera l AN fibers. To graµh these curves, research e rs insert
an electrod.e very close to a single AN fi ber, a nd they measure hm.1,• intense
.s the s ine waves of different frequencies must be fo r the ne uron to fire faster
3l than its normal, spontuneous firing rate. The frequency tha t increases the
1 neuron's firin g ra te at the lowest in te ns ity (the lowest point on the thresh old
• 0
;': hming cu rve) is called the nemon' s characteristic frequ e ncy (C F).
From Rgure 9.1 4,. you can see how outer h air cells are required for
thresh old tuning curves to be focused on a narrow range of frequen cies.
Outer hair cells make parts of the cochlear partition s tiffer in ways that make
the responses of inner hair cells m ore sen si tive (Dong and Olson, 2013) and
more sharply tuned to particular frequencies (Hudspeth, 2013).
Up to th is p oint, the way the ea r transd ucesacoustic en ergy a t different
OJ 1.0 10.0
Frequency (kHz)
frequen cies into a pa tte rn of ne u ral resp onses seems fair ly s trai ghtforward.
A low-inte nsity sine wave tone with a certain freque n cy will cause ce rtain
AN fibers to increase the ir firing rntes, w hile othe r AN fibers continue to
fire at their s pontaneous rates. As long as the brnin knows w hich AN fibe rs
FIGURE 9.14 O utef hair OOls

Improve b oth sensltMty and fne-
w Fil..,..r4 -Fiber 6 I
quency SflolgctMty. Three threshold
tuning c urves show responses
threshold tuning cll'Ve A graph
v•.t um hair cells are ac tive, and the
pbttlng the throohol:ls or a neuron cr
dashed li ne shows what the right-
fiber In response to sine waves with
rrost tuning c urve would look Ii ke
varying frequencies at the IO\vest Inten-
if outer hair cells were not acttvtt.
sity that win give nse to a rooponse.
Hightlr-intenslty ton"3S would 00
characteristic frequency (CF) The required to excit.e th9 AN fiber, and
0.1 1.0 10.0
frequency to which a particular audi- the fib.:ir 'WOUid be less seloctive in Frequ ency (kHz)
tory nave fiber Is most sensitive. frequ ench;is to which it would fire.

have w hich characteristic frequencies, the brain can interpre t the patte m of two.tone sl.4)presslon A decrease
firin g rntes a cross all the A N fi bers to detem1ine the frequency of :m y to ne (as In the firing rate of ooo alJ::lltory nerve
lon g as it is within the range of frequ en cies picked up by the human fiber due to one tcne. when a second
tone is presented at the same time.
Unforh.matel}'i it's n o t qui te this s imple. Almost all sound s in the environ-
ni.e nt are rn ore co mpl ex than s ing le s ine wa ves, and m ost sounds \Ve hear are lsolntenstty curve A map plotting
also much louder th..1.n the very quiet sound w aves used to measu re threshold the fl ring rate o1 an auditcry nerve fiber
against varyln;i frequencies at varying
tuning curves. So a lthough the p reviou s paragraph cap tures the gist of h ow Intensities.
AN fibe rs code for so und frequencies, we have to do a bit more work to lUl-
derstand h ow high er-intensity, complex so unds a re en coded in the auditory
nerve. We will consider h vo of the .specific complications we h ave to deal ,,...; th.
Then, we'll look at one additional m echan ism, related to timin g r ather than
p lace, tha t thea ud.i tory system uses to convey low-fre quen cy components o f
sound waves.
TWO-TONE SUPPRESSION The rate at which an AN fiber responds chan ges

w hen energy is introdu ced at nearby frel1uencies. In particular, when a sec-
ond tone of a slightly different frequency is added , the rate of neura l fi rin g fo r
the first ton e actu a lly decreases--a phenomeno n called suppres-
sion (R g ure 9 .15) . S uppression effects are particula rly p ron ounced wh en the
second (suppresso r) tone has a lower frequen cy than th e firs t tone. fn other
words, iJ we're 1-eco-rding fro m an AN fiber wh ose CF is 80CX) Hz and we use
an 8000-Hz test tone, a 1000-Hz s uppre.ssor tone a gr&'l te r effect on the
neu ron 's firing rate than a 15,CXXl- Hz suppressor ton e h.:'l.s. You can see how
understanding the response of the w hole auditory nerve to co1nplex sounds
(that is, frequency combinations) is m ore compl icated tha n s imply addin g up
the responses of indlv iduaJ AN fibers to individu..'ll pure tones.
RATE SATURATION Sounds that matter m ost to listeners-most con versational

sp eech, for example---are usuall y heard at intensiti es, s ud1 as 60 or 70 dB,
that greatly exceed the thresh old for just de tecting sounds. Are AN fibers as
selecti ve for thei r ch Ltracteristi c frequencies a t levels well above th res hold as
they a re fo r the ba rely a udible sounds used to chart tfilesh old h.ui.ing cm ves?
To answer this question, we ca n look at lsolntensfty curves, whid 1 we
chart by measuring an AN fiber 's firing rnte to a ,\Ii.d e range of frequen-
80
1 40
"
li: 20
1.0 10.0
Frequency (kHz)
FIGURE 9.1 5 Two-tone suppression. The tuniflg'CUrw plots

the responses o f orie auditory n4ilrve fibQf with a characteristic frequency o f 8000
Hz. Whengver a second tone is playtid at thQ frequsnci91S and within the Ng ht
red a reas to each side, the response of this AN fiber to an 8000-Hz tone is reduced
(supprgssed),

276 CHAPTER ()
FIG URE 9.16 lsointo9onsity 'func tions for om:i AN fib9r 300
w ith a c haraci€fistic frequency of 2000 Hz. Tones o f
varying frequencies ore presentQd at 20, 40 t 60, and
80 dB. The neuron fires vigorously t o a wid.s< range of
fraquranc ioo (mo stly lower) w hen JntEflsity is Increased.
Note that th9 20-dB curve rQSQmbles an upside-do wn
threshcid tuning c urve (compare with Figure 9. 13)
bGcau8'1! 20 dB is almost as low as th9 intensiti9s at
which thresholds are mMSured.
0 500 l 000 1500 2000 2500 3000 3500 4000

Frequency (Hz)
cies, a ll presented a t the sam e inte ns ity level. Figure 9.16 s hmvs a famil y of
isointen.sity curves for one AN fiber wi th a C P of2000 Hz. The bo tto m curve
shows the average firing rate (numbe r of action p o te ntials per second) of the
n euron in response to 20-d B to nes with freq·ue ncies between 50 a nd 3300 H z.
Tl'le other cm \res track firing ra tes for 40-dB, 60-dB, and 80-dB tones over the
same freq uency range .
\Ve learn from these c un1es that for relatively quiet, 20-dB sounds (this
is a bout the so tmd leve l of ntstling in the wind ), the ne u ron is s till
quite n arrowly tun ed , firing much fas ter in response to its CF (2000 H z)
than to neighboring frequencies. At 80 dB, however, the ne u ron appears to
fire a t about the s(1me rate for any frequency in the range of 8-00-2500 Hz. In
othe r words, frequencies sudi. as 1000 Hz, to which the AN fibel' had almost
no response at lo\.V intensity levels, evoke qui te substantial responses w hen
intens ity is
1i-1e phen om en on behind this broadening of frequency selectivity is called
rate saturation. Re mem ber that AN fibe rs fire in response to the displace-
ment of s tereocilia on hair cells. The farther s tereocilia pi vo t, the faste r AN
fib ers linked to that hair cell fire. For a 20-dB tone at 1000 H z, the s tereocilia
on th e hair cell feeding the AN fiber feature d in Figure 9.16 will not be nd at
alt so the fibe r 's firing ra te remains a t its res ting level. 111e firing ra te rises
a bove this res ting leve l w he n the frequen cy of the 20-dB tone is increased
82 to 1QOO H z, a nd it teaches its h ighest level a t the AN fi ber' s characteristic
frequency, 2000 Hz.
\ Vhen the intensity is increased to40d B, however, the bulge in the ves tibu.l ar
cn n3J is so large that stereocilia start dis placing even to a 1000-Hz tone (see
Figure 9.16). If we increase the freque ncy to 1250 Hz, the firing rate increases,
and it in creases even m ore 3t 1500 H z. The problem is tha t a t about 1500 Hz,
the fibe r 1s firing rate m axes out (snh1rates). Stereocilia are pi voting as much as
they can a t th is point, so increasing the fre quency of the tone h as no additional
rate saturation The point at which
a nerve l bar Is lnng as as pos-
effect on the AN fiber 's firing rate tmtil we increase the frequency above the
sible and further stimulation Is Inca- fiber' s CF and the firing rate s tarts droppin g again. For SO-dB tones, the fibe r
pable of lnoreaslng the firln;i rate. maxes out at even lower and higher freq ue ncies relative to the CF of 2CXJO H z.

450 FIG URE 9.17 Firing rate plotted against sound inten-
sity for six auditory nierw fibers.: thre.e low-spontanoous
400 (red) and three high-spontaneous (blue). Aring rates
for all six neurons incro9ase with inc r"10.Sing sound level.
350 Low-spontaneous neurons require higher-Intensity
sounds before they beigin to fire, and they continue to
inc rease firing rate to sound lodvflls..
50
w m m m w m ro
Sound intensity (dB)
For mod erate ly intense sounds, s uch as speech, the bmin can no t rely o n a
si ngle AN fiber to determine the frequency of the tone. Forex..'lmple, we can't
use the rule "if an AN fiber with frequency of 2000 Hz is firing
very fas t1 the sound must be 2(0) H z" because, as Figure 9.16 illus tra tes, this
ne uron vdll al.so fi re at its m aximum r.:lte to a lCOO-Hz ton e if the sound wave
has a large-enough amplitude.
One way the auditory syste m gets <'\round this problen'I. is to use AN fibers
with different s pontaneous firing ra tes. Figure 9.17 shows rateplntenslty
tunctlons for six fibers, all of which listen to the same hail' cell (remember
that dendrites from 10-30 a uditory neurons synapse with each inner hair
cell). To plot these Cltrves, the intensity leve l of a tone at the AN fiber 's CF is
slowly raised from 0 dB up to 90 dB. As you can see, the resting rates of some
fibers (whose functions are plotted in red) are Jess than 10 spikes per second. rate-intensity function
plotting the lrtng rate of an au:lltory
These neurons are fibers. The blue lines plo t firing rates
nerve fiber In response to a sound
for fibers, which fire 30 or more times per second, even of constant frequency at Increasing
in silence. fibers h.a.ve resting rntes behveen these levels. Intensities.
High-spontaneous AN fibers are somewhat analogous to rods in the retina: tow-spontaneous fiber An audi-
they are especially sensitive to low levels of so und 1 responding at rates above tory nerve fiber that has a low rate oass
resting level even w hen decibel levels are quite low. The trade-off is that the than 1O spikes per secord) of spon-
firing rntesof these fibers quickly reach saturation, so their frequency selectivity taneous flrlrrJ: low-spontaneous flb€<s
is relative ly poor w hen intensity is relatively high . Low-spontaneous fibe rs require relattvely Intense sound before
they v.;11 fire at higher rates.
are more like cones, requiring more energy (higher-intensity sound waves)
to s tart responding, but retaining thei r frequency selectivity over a bro.:' lder high-spontaneous nber An al.dl-
tory nerve fiber that has a high rate
range of inte ns ity. (more than 30 spikes per second) of
Jn addition to having different AN fibers with different sp ontaneous rates, spontanoous firing ; hlgh-spmtane::us
the auditory system can accura te ly determine th e frequency of incoming fibers Increase their firlrg rate In
solmd waves by integrating i.nfom1ation across many AN fibers, and using roop:::nsa to relat1vely 10\'lf levels of
the pnttem of firing rates across all these fibers . Reme mber that in the visual sound.
syste m, we use the pattern of firing across o ur tluee types of cones to calculate mid-spontaneous fiber An audi-
the w avelength of light. The auditory system uses the same principle, but it tory nerve fiber that has a medium rate
has some 14,((1() AN fibers in each ear to discern acoustic frequency. Conse- (10-BO spikes per secrndJ of spon -
tanooLS firlrg. The characteristics of
quently, the freque ncy sensitivity of the hlm1a.n audi tory system as a whole is mid-spontaneous fibers are Intermedi-
exquisite across a \vide range of intens ity levels, despite the coarse selectivity ate between low- and hlgh-spontane-
of individual AN fibers. cus fibers.
0336 Amma Appa

278 CHAPTER ()
if ll illllt l\, lllll l Tnne

FIG URE 9. 18 Pha:sQ locking . The histogram (bottom) shows neural spik9s for an
AN fibe' in reisponse to the same low-frequtncy sine wave (top) being p ayed many
times. Note that the nQJron is rnost likety to fire at one particular phase of each cycle
of the sine wate. This phas9 locking pro\lid9S a tem poral oode to sound foequ sncy.
THE TEMPORAL CODE FOR SOUND ffiEQUENCY ln addition to the coch lear
p lace code, the au ditory syste m has anoth e'r W'ay to encode frequency. As
Figure 9 .18 illustra tes, many AN 6bers tend to fire action potentials a t one
pl!rticular point in the phase of a sound wave--a phenomenon Ctl.Ued phase
locking. Phase locking may occur because AN fibers fire w hen thes tereocilia
of h air cells move in o ne direction (e.g., as th e basilar membrane moves up
toward the tectorial membrane) but d o not fire when the s tereociHa move in
the other direction. Recall from o ur disc ussion of mechanoelectrical transduc-
ti on in s tereocilia that the encod.i ng of time is extremely accurate.
The existence of phase locking means that the firing pattern of an AN fiber
carries a temporal code for the sound wave frequency. For exarnple, if the AN
fiber fires an action potential 100 times per then dO\-vn.stream ne urons
lis tening to the AN fiber ca n infer tha t the sound wave includes a frequency
cornponen t of 100 H z.
\ Vhile reliable for lower freq uencies, temporal coding becomes inconsis ten t
for frequencies higher than 1000 Hz and is virtually absent above 4000 or SCOJ
Hz. In large part, this inconsistency is due to the refractory period of the AN
fiber. For hig h freq uen cies, fibers simply cannot produce action potentials
quickly enough to fire on every cycle of the sotmd. However1 multiple neurons
phase locking Flrlr"I) of a s ingle could, in principle, encode highe r frequ en cies as a group . For example, fo ur
neuron at me distinct point In the neurons could each fire only once every four th cycle o f a 2000-Hz sound . 1f
period (cycle) of a swnd w""" at a the four neu rons "took huns," each would have to fire only 500 times per
given frequaicy. (The neurm nee:::t not second to hilly encode the 2000-Hz sound in their comb in ed temporal pat-
fire on every cycle, but each firing will
occur a t the same p::jnt In the cyde.) ten1. 1his id ea has a long his tory 1949), and it is referred to as the
volley prfnciple {Figure 9.19 }. According to this hypothesis, neurons sustain
terrporal code Tuning of different
parts of the cochlea to different Ire-
a temporal pattern of firing much like the pattern of Revolutionary War-era
quencles, In which Information about soldiers firing g uns from the front line of a forma tion w hile the second and
the particular frequency o f an lr>::ornlng third lines took time to reload.
scund W<Ne Is coded by the timing of Inte restingly, even AN fibers with relatively high-frequency CFs encode
neural firing as It relates to the period 10\·ver-frequency energy in the temporal pattern of thei r responses. For example,
of the sound.
if you are listenin g to a fairly loud sol.md that is a combination of 200- and
volley principle The Idea that mul- 8000-Hz sin e wave tones, a nemon near the base of the cochlea that becomes
tiple neurons can provide a temporal w ildly excited by the 8000-Hz component of the sound will also ten d to be
code fa frequen cy If each neuron fires
at a distinct point In the period o f a
phase-locked to the 200-H z component. Th is ne uron thus carries infoml.ation
scund Wa.te but does not flre on every about both the high-frequen cy component (via place coding, beca use the brain
period. la1ows the neuron's CF) and the lm-v-frequency component (via te1npornl coding).

Sin.w.w• l! \MM/\/V\/VWV
Neuron A
l l l
Neu ronB
ll l1 l
l l
Neuron e
Neu ron D
l l
Neuron E
l l
Ll l l l l l1 l Ll l L
Total response
(A- E <ombm•d)
Time
FIGURE 9.19 The volley principe, Even If on.e n.eurm cannot fire in response to
every cycle o f a higher-frequency tone. muttlpe AN fibers together c an provide a
t9mporal cod9 for freqUQlcy if diffQt"ent neuron s (A, B. C. D. E) each fir€! at differ9nt
J)eflod s of the sine wa-1e.
Auditory Brain Structures

The nerve fibers that make up the auditory nerve share cranial nerve V[J]
nerve fibers for the vestibular system (w hich is discussed in Chapter 12); this
is why cranial nerve VIll is kn own not only as the auditory nerve, but also as
the ''vestibulocochlear nerve. " All AN fibers initially synapse in the cochlear
nucieus1 which consists of three subnuclei (Figure 9.20). 1lle coc hlear nucleus
con tains m any different types of specialized neurons. Some of these are espe-
cially sens itive to onsets o f sound a t particula r frequencies. Some are sens itive
to the coin ciden ce of onsets across many frequencies (they fire when multiple
frequencies initially begin, but s top firing if thesotmd continues p laying) . Some
cochlear nudeus nemons use latera l inhibition to sharpen th e tuning to one
frequency by suppressing nearby frequencies-a mechanism reminiscent of
th at used by retinal ganglion cells to respond to s pots of light instead o f broad
fields of light. Others resp ond in exact ly th e sam e \vay as the AN fibers th.at
feed th.em . Son1e neurons appear to serve as little m or e than quick relays from
the cochlea to the superior olive, an other brain ste m nucleus . These ne urons
are especia lly good a t preservin g accurate tinting information.
cochlear nucleus The first brain
As Figme 9.20 s h ows, some of the ne urons that project from the cochl ear stem nudeus at which afferent audi-
nuclei to the superior olives cross over to the opposite s ide of the brain. Thus, tory nerve fibers synapse.
't'here inputs from ead1 visua l field remain separate superior olive An brain stem
until they have extended a fair distance in the visual cortex, signals fro m both reg bl In the audltcry pathway where
coc.hle..'\s reac h both sides of the brain after only a s ingle synapse. As we will Inputs frcm both ears converge.

280 CHAPTER
FIGURE 9.20 Pathways in the

auditory system. Not all pathways
ar.e shoVYfl in this scMfnatic.
AJthough there are two parallel
pathways, information from both
ears oom{IS together wry early in
the auditory systtm . at the superior
olives.
see in C hapte r 10, this direct rela y of infonnation across both ears is essential
to usi ng tiny timing differences between the h vo ears to d etect the direction
of a sound .
Neurons from the cochlear nucle us and superior olive travel up the brain
s te m to the inferior colllculus. Most (but not all) of the input to each inferior
colliculus comes from the opposi te (C'ontralateral) ear; tha t is, the left inferior
coll ic u.lus lis tens mostly to the right ear, and vice versa.
The medial genlculate nucleus of the thalamu s is th e las t s top in the
auditory pa thway before the cerebral cortex. Like the lateral geniculate of the
visual syste m, there are many m ore ne urons that project fro m the cortex to the
media l genicu.la te {efferent neu rons} than project from the med ial gen.iculate
to the cortex (afferen t neurons). These efferent connections, some of which
presumably convey informal;iqn back to lower stages of the a uditory system,
p rovid e forther anatomical evidence that se:nsmysys.fem two-way streets,
in w hich feedback from the brain is tightly integrated w ith sensory infomla-
tion flowing up to the brain.
All stnKtu res o f the auditory system, beginning with the basilar me m-
Inferior colliculus A midbraln brane a nd continuing th rough the cochlear nucleus, s uperior o live, inferior
nucleus In the auditory pathway. colliculus, a nd medial geniculate nude us, s how a consistent organizati onal
medal genlculate nucleus TI1e pattern in which n e u rons are a ligned base d on the frequ e ncies to w hich
part of the thalamus that relays audi- they are most sensi tive. 11Klt is, ne urons n1ost responsive to low-frequ ency
tory slgMls to the temporal catex and
reoetves Input from the auditory cortex . energy lie o n one e d ge of ead1 s tructure, n e urons respondin g to high fre-
quencies lie o n the other edge, and ne urons responding to o ther frequencies
tonotopk:: organization An are s pread out in an orderly fashion in between. The per vasiven ess of this
arrangement In which neurons that
respond to different frequencies are tonotopic organization pattem re flects both the early mechanical properties
organized anatcmk::alty In order of of transdu cti on and the importance of the frequency composi tion of sounds
frequency. for a uditory percepti on.

Primary auditory cortex is often referred to as A1, just as primary visual

cortex is called VJ. Tonotopic is less consistently mai ntained
in A1 (Bandyopadhyay, Shamma, and Kanold, 2010; Rothschild, Nelken,
and t>.tizrahi, 2010). Neurons fro m A 1 project to the surrow,ding belt area
of cortex, and neurons from th is be lt sy napse w ith neurons in the iJdjacent
parabelt area (Figure 9.21). Jus t about any smmd will cause activation in
sorne part of Al. In the belt and parabelt areas, referred to as 1'seconda ry '' or
"associational" auditory areas, simple sound s s uch as sin e \ ..·aves elicit less
activiry; particularly if the stimuli d on' t change much over time. Tlrns we see
that, as in other sensory systems, prO\.""essing proceeds from si mpler to more
complex s timuli as we m ove farther along the auditory pathway. \Ve also find
greater evidence of cross-modal processing (e.g., combining acoustic and optic
information), particu larly in parabelt areas.
Compa ring the overall s tructure of the audi tory and visual systems shows
that a relatively large proportion of the processin g in the a uditory sys tem is
d one before A1. By contrast, as you learned in Chapter 3, the majority of the
mos t important visual ocClrrs in cortica l areas Vl and beyond.
Auditory capa bilities that may be most important to humans, lis tening to
speech and music, are s ub.served a l.most entirely by cortical areas. We w W
return to the role of th e cortex in auditory p rocessing when we discuss s peech
and music perception in ChapteT n.
Basic Operating Characteristics of the FIGURE 9.21 The first s tages o f

auditoiy processing begin In the t.gm-
Auditory System poral bbe in areas 'Within the SyMan
Up to this point we h ave d iscussed the anatomy of the auditory system and the fissure. The top picture ls from the side
o f the brain, and the lo\Nel' two pictures
physiology of how the system encodes the h.vo basic physical attributes of solmd
are looking down at the brair1 with the
waves: amplitude (intensity) and frequency. We've learned this through direct parietal cort8X c ut awrty. Primary audi-
observati on of anatom ical structures. Scientists exa mine basilar membranes, tory cortex (A. 1) is in the center. It ls sur-
record from a uditory nerve fibers, d issect various brain stTuctures, .md so o n. rounded by belt regions, and parabett
We turn n o\v to the findin gs of researchers who have approached the regions extend past the b91t to thg. front
and side. {From Brugge o.nd Howard,
auditory system from a different perspective. In.s tead of play ing a so und and
2002.)
trying to determin e h ow one p articular ne uron responds, we can instead
play the sound and ask actu al human Listeners--each of w hom is the s um
total o f a great many n eurons-what they hear. When human listeners are
asked to report their auditory sensa ti ons, their answers are due partly to the
acoustic properties of the sound signal and partly to th ei r °'"''1 psychological
cha racteristics. TI1is 1nethod of inves tigation is thu s called psychoacoustics.
primary auditory cortex (A1) The
Scientfats who s tudy psych oacous tics (psych oacous tician s) are a lways nrst area wlt111n the temporal lobes of
careful to dis ting uish between the physical characteristics of sounds and the the brain responsible for processing
impressions of these sotmds for listeners. As v•:e noted earlier, whereas frequency, acoustk; information.
measured in hertz, is a physica l description of the spectral composition of a belt area A region of =tex, directly
sound, the subjective attribute o f frequency for listeners is pitch. Sounds are adjacent to the primary auditory cortex
m easured with respect to frequency, but lis teners hea r pitch. Similarly, the (A 1). wltl1 Inputs from A1. where neu-
intensity of .sound is m easured as sound pressure in d ecibels, but listen ers rons respord to more complex dlar-
hear loudw?ss. If the auditory system operated exactly th e sam e as electron ic acterlstlcs of sounds.
me.asuring d ev ices, we cou ld treat the terms frequericy and pitch and the terms parabelt area A region of cortex,
intensity and loudness interchangeably. As we will see, h m.1,·ever, biological lateral and adjacent to the belt area,
where neurcns respond to mcre com-
auditory systems d o n ot work ex..'\ctly as elec tronic m easuring d evices '"'ork.. plex. ch3.racterlstlcs of sounds, as well
For example, one solllld wave may be hea rd as quite a bit loud er than another, as to Input from other senses.
even though the two waves have ex..'\ctly the sam e amplitude. Carefol s h1dy
psychoacoustics The stldy of the
of the differen ces beh·veen the responses of electron ic d ev; ces (so und-level psychdoglra = raatoo of the physical
meters and spectrum analyzers) and bi ological lis tening d evices (human dimensions of acoustics; a branch of
beings) p rovi des great insight into how the human auditory system works. psychcphyslcs.

282 CHAPTER
FIG URE 9.22 The IOW'9st CUtvQ (r,gd)

illustrates the threshold for hearing uo
sounds at varying frQG!.J9nciGs. The
c urves 18.00!ed Rl O,R "20." "30.-" and
so on are equaH oudniass curves. For 90 _
a single oontour, tongs of diffeiri:int 1re-
quenc ies have different physical intensi-
ties, but they sound 9C1ually loud. (After
Suzuki and Takeshlma, 2004.) -:=: •o
j 50 /"
-g 40
;, 30 /
20
10
0 Audibilit)· threshold
OD2 ODS 0.'1 0.2 0.5 10

Frequency (kHz )
Intensity and Loudness

The botto m curve in Figure 922 shovvs the hmna n audlbility threshold, wh ich
graph s the lowest sound pressure level that 1."'an be reliably d etected across the
frequency range of hum an hearing (20- 20, 000 Hz}. Nore tl>0t the bes t (lowest)
absolute thresholds fo r hmnan hearin g are beh\.•een 2000 and 6CXJO H z (2- 6
kilohertz [kHz]; rem embe1· tl'lat tl10se frequencies are enhai1ced by the physica l
properties of the aud itory cana l). Tiuesholds rise on bo th sides of this range,
meaning tha t h igher- and lower-frequen cy sound waves mus t h ave larger
amplitudes in order to be heard .
The other lines in Fig ure 9.22 are equal-loudness curves {Suzuki and
Takeshima, 2004). We o btain these curves by asking liste ne rs to equa te the
lo udness of sounds 'vi th differen t frequen cies. lh: s tarting point for each curve
is a lways 1000 H z, so the curve marked 1140" tracks the an"lplitude necessary
to m ake tones at o the r freq uencies soun d exactly as loud as a 1000- H z, 40-dB
tone; the curve marked 1160" represen ts the decibel levels n ecessary to ma tch
a 1000-Hz, 60-dB tone; and so on. As the figure sh ows, the sam e pattern o f
frequency-dependent sensi tivity tha t we see in the audibility thres hold curve
extends to sotmds above threshold. (See Web Activity 9.3: Equal-Loudness
Curves.)
The two ora nge tick mark s in Figure 9.22 indicate th at a 200-Hz ton e
presented at 70 dB sounds Bho ut as loud as a 900-Hz ton e p resen ted at 60 dB
(that is, both p oints full on the equal-lo udness curve marked 1160"), whereas
a 2000-H z tone presented at 40 dB sounds much lo uder than a 100-Hz tone
p resented a t the same level (ptuple tick marks). Th ese observations demo n-
s trate the inequality of sound p ressure level and lou dness: equal-amplitude
sound s can be perceived as softer or louder than each other, depending on
the frequencies: of the sound waves.
Another way to track the relationship between amplih1deand loudness is to
pick one frequency, s teadily raise the intens ity of the sound, and ask a listene r
audibil tty threshold The lowest to jud ge hO\v the loudness increases. Experiments of this type s how, once
scund pressure level that c an be reli-
ably detected at a given frequency. again, that the relationship behveen intens ity and loudness is far from perfect.
Doubling the perceived loudness o f a solmd requires m ore than a. d oublin g
e<J,Jal-loudness c urve A graph
plotting sound pressure level (dB S PL) in the a m ount of acous tic en ergy present in a. sound wave, especially above
against the frequE<lcy for which a lis- 40 dB. The s.a rne kind of relationship holds in vision : the number of photons
tener pefcelves constant loudness. mus t be more than doubled to d o uble the perceived brightness of a ligh t.

The loudness of a so und also depend s o n its dura tio n. \Vithin limi ts, longer ten'1)oral integration The process
sotmds are heard as being louder. Again 1 the same thin g ha ppens in vi sion: by which a sound at a constant level
flashes of light appear brighter when they 1-ast longeL The reason for this gen eral ls pefcelved as being louder when It
Is of greater duration. The tenn also
phenom enon is that the perception of lo udness or brighh1ess depen ds on the applies to perceived bnghtness. which
summation of energy over a brief, but noticeable, period o f time-a process depends on the duratb'l of light.
called temporal integration. For hearing, tempora l integration occ urs over an
inte rval of 100--200 milliseconds (rns). So if a sotmd is presented for less than
100 ms, it will be perceived assoh er than the sa1:n e amplitude and frequency
presen ted for 300 rns. H ovv·ever, there \Vill be little difference in loudness per-
ception if the durati on o f the sound is increased from 300 to 1000 ms or longer.
ln addition to stud ying a bsolute lo udness judgrn en ts, p.sycho<lcous tidans
are interested in how proficient huma ns are a t discriminating between the
loudness levels of two sounds . There are several different ways to measure
the smallest differen ces in intensity that can be d etected, and many measures
shm"' sens itivity to changes of less th.an 1 dB. This ability is quite impressivel
given th e wide range o f sotmd intensities (from 0 to over 100 dB) that humans
can perceive and the fact that, unlike the vis ual system, the auditory systen1
is always sensitive to this entire range (rem ember that, to ach ieve maxim um
visual sensithity, we n eed time to ada pt to lower or higher ambient light lev-
els). Although the ability to discriminate beh-veen s ubtle loudness differences
might not seem all that important to s urvival, \.Ve w ill see in Chapter 10 how
the a uditory sys tem uses differences betiNeen the intensity levels of sounds
. to detem,ine where sound sources are located.
reach ing the left a nd right e. U"S
For a time, it was difficult to tmd ers tand h ow listen ers could be sensitive
to s uch s mall d ifferences in loudness over s uch a large range. Sow1d w ave
inte nsi ty is gen erall y s ig naled by the fuing rate of a uditory nerve fibe rs:
larger intens ities (loud sounds) correspond to high er firing rntes, and smaller
intens ities (quiet sounds) correspond to _l ,xei:)iring rates.,Yo u sh ould recall
th at the intensiti es required to fire (thresho lds) Voiry r m n AN fiber to
th e n ext. For exa mple, on e fibe r m ight selectively respond to the ra n ge of
amplitudes between 0 and 25 dB, another might span tl>e range of 15-40 dB,
a third might dB, and soon (seefigure9.17J. Afull population of
neurOJ\S ,,..,rjth different tlU'esh.o lds can then encode a much b roader range of
intensities th.an is possible with any single neu ron. In addition, remember tJw t
neurons becom e respons ive to a bror1der range of frequencies when intens ity
is higher (see Figure 9.16). O n e result is that, as sotmds become more intense,
many m ore AN fibers become excited .
Frequency and Pitch

The ton otopic organization o f the auditory system , from bas ilar membrane
to primary auditory cortex, is a ve1y big hint that frequency compositi on is a
fund am ental de te nn.inant of how we hear sounds. More than anything else,
psychoacoustidans have s tudied how listeners perceive pitch, the psych o-
logical counterpar t to frequency. As is the case wi th in tensity a nd loudness,
the freq uen cy of a sotmd is related to, but not pe rfectly correlated \<\-i. th, the
perceived pitch of the sound. For any given frequency in cre.:1se (for example,
an increase of 50 Hz), listeners will perceive a grea ter rise in pitch for lower
frequencies than they d o for h igher frequencies. Consequently, listeners per-
ceive a greater pitch diffe_rence w hen a tone s hifts fro m 500 to 1000 Hz than
w hen a to ne shifts from 5000 to .5500 Hz.
Researd1 d one us ing pure tones (sounds composed of a single sine wave)
indicates that h\.Unans are rernar kably good at detecting very sma ll diffe r-
ences in frequency. For example, listeners can discriminate behveen tones of
999 and 1000 Hz- a difference o f only one-tenth of 1%! Pitch discrimination

284 CHAPTER
masking Usln;i a second sound. at the lower a nd higher ends of the a uditory system 's frequency range is no t
frequently noise, to make the d etection quite as good, but it is still impressive. Part of th e reason that discrinUnabili ty
of arother sourd more difficult. decreases for high pitd1es appears to be that the temporal code for frequen cies
white noise Noise consisting of all s tarts breaking down above 1000 Hz an d is rela tively n onfun ctional above
audible frequencies In equal amounts. 5000 Hz. Performance thu s s uffers because the auditory system has to rely
White noise In hearing is analogous to
excl usively on place coding (the pattern of AN fiber responses from diffe rent
white light In >Aslon. for which all wave-
len;iths are present. parts of the cochlea).
critical bandwidth The range of Psychoocoustidans also use masking experimen ts to investigate frequency
frequencies conveyed within a channel selecthity. In the research described in the previous paragraph1 listeners never
In the auditory system. hear more than one.sound frequency at a time. In a maskingexperiJnent, multiple
frequencies are combined, and we see how well listeners c.a n pick out certain
componen ts. \ Ve look at how effective one sound-the masker-is at hiding
an other sound . Ymlve already learned abou t the phenomenon of two-tone
suppression, in wh ich a n auditory nerve's response to one frequency can be
disrupted by a second frequency. 1f place coding underlies pitch perception, we
s ho uld expect to see the sa m e kind of phenomenon in masking experiments.
[n the classic approach to measuring frequency selec ti vity using m.asking,
a s ing le sine wave tone is placed in the middle o f a band of acous tic n oise
(Fletcher, 1940). White noise is a signal tha t includes equa l energy o f every
frequency in the human a uditory range (20-20,000 Hzt jus t as white light
includes light rays of a ll freq uencies in the visible spectrn m. A m ore limited
band of noise mi ght include all frequencies in the range of 500-1500 Hz; an
even small er band could span 500-510 Hz.
ln a typical experiment, '"''e might start with a 2000-Hz sin e wave test tone
presented along lvith a very narrow band of 1975-2025 Hz. 'A'e would
then adjLLst the intensity of the tes t tone until listeners could just hear it over
the n oise. Next we would increase the bandwid th of the noise, perhaps from 50
to 100 Hz1 so th.at now the noise would include frequencies between 1950 and
2050 Hz. As we might e"-pect, the listener wo uld need to increase the intensity
FIG URE 9.23 C ritical bandwidth and of the test tone to be able to hear it over this broad er range of noise
rnasklng. (a} To moosure the width of a Lf we keep \.videning the bandwidth, however, we wi ll eventually re,:,d1 a
c ritical b3nd, subjects listen for a tone point at w hich a dding m ore frequencies to the noise s tops affecting the detect-
in thg canter o f a band of noise of con-
stant intensity. It Is harder to detect the ability of the test tone. 111e size of the noise band at this point is ca lled the critical
tone as the b and of noise widens up bandwidth (Figure 9.23a). For tl1e experimental d ata plotted in Figure 9.23b,
t o a point. when 1utthe.r widening has the critical bandwidth is400 Hz. [n this case, to p ick out a 2000-Hz tone from
no effect on detecting the tone. This the background noise, lis teners must increase the intensity of the tone w h en
1JQint d iefiniBS th9 critk;;al bandwidth. the bandwidth is widened from SO to 100 Hz and from 200 to 400 Hz, but going
(b) For th Is plot of bandwidth data, a
2000 -Hz to ng was US9d (Schc:on w€11t
and Moore, 1900). The bandWdth o f
the noise had no effect on det9ci:ion of (b)
the tone whfW'"I it exceeded 400 Hz. so 76
400 Hz is the w idth of the critical band
a! 2000 Hz.
3 75
'\.. ---.
(") .,ei 74
Critical
"'
:E
b.mdwidth
i-5 73
-; 72
& 71
50 I00 200 400 8CO 1600 3200
Maslcing-noi.se b.1ndwidth (Hz)

from a 400-Hz n oise band to nn 800-Hz band does not require the listener to
m.ake the test tone an y lo ude r. In fact, the 400-H z noise b...,,nd is just as effec tive
a m asker as w hite n o ise covering the entire spectnun of hum.an h earing.
Resul ts from the paradig m have helped cement the role of place
coding in pitch perception by revealing si milarities between perceptual effects
and phys iological fi ndings. First of all, the \-vid th of the critical band v.id th
chan ges dep ending on the frequency o f the test tone, and these widths cor-
respond physi,cnh;µa cing..of frequencies a lo ng the basilar membrane.
Fo r exam ple, we know tha t u greater proportion of the basilar metnbrane
vibrates in response to low frequencies, and tha t hi_;he r-frequency ranges
vibrate sma lle r p o rtions of th e membrane. Correspondingly, masking stud ies
sh ow that the band\1\-;dths for low frequencies are s maller than the
critical band'"';d ths fo r h igh frequencies, because the spacin g between low
frequen cies is larger on the basilar me mbrane.
Anothe r importan t finding is that masking effects are asymmetrical Mo.re
specifically, for a m ask whose band wid th is below the critica l bandwidth of a
test tone, the m ask is more effective if it is cente red on a frequ ency below the
test tone's frequency-.1 phenomenon called the " upward spread of masking."
This phenmnenon m ay seem counterintuitive, but a look back a t Figure 9.12
shm ..·.s how disp lacement of the basilar membrane (the trave ling \¥ave) extends
from the high-fre.:iuency base to th e low -frequency apex. Dis placement fo r
low -frequency energy sounds toward the apex leaves a trail of displacement
across h igh- freq ue ncy regions toward the base. From two-tone s uppression
experi ments, we also learned that suppression is greater if the suppressor tone
is below an AN fiber's characteristic frequency th an if the s uppressor tone is
above the fiber 's dlaracter istic freq ueni..")' (see Figure 9.15).
Hearing Loss
Ro ughly30 million Americans suffer some for m of hei'\ri..ng impairment. \Vhen
we talk nbout hem·ing loss, we typica lly d o not mea n the to tal loss of all hearing
(deafness), but rathe r the e levation of sound th_resholds. For exa mple,
des that u sed to be audible at 20 dB may become inaudible unless they a re
presen te d. a t 40 or 60 dB. Of c01u se, in the end we do not jus t need to detect
-s ounds; the term !rearing really refers to using spectral a nd temporal differences
between sou nd s in orde r to something about e ve nts going on in the en-
vironm ent. As we vvill see,cornmon forms of hearing loss can affect the abili ty
to interpret sounds, even w he n the soi.mds are loud enough to be detectable.
H earing can be impaired by damage to any of the structures al ong the
chain of auditory processin g from the outer ear a ll the way up to the audi-
tory cortex. The simp lest way to introduce some hearing loss is to obstruct
the ear canal, thus inhibiting the ability of sound waves to exert press ure on
the tympanic membra ne. Many peop le d o this on purpose by wearing ea r-
plugs. A less intention.11 hea ring loss can be created by the excessive buildup
of earwax (cerumen) in the ear canal. Thls proble m is easy for clinicians to
remedy, as long as the effort to clear out the ear canal d oes no t dilm age the
tympanic me mbrane.
A nothe r type of hearing impairme nt, calle d conducti ve hearing loss,
occi.u s w he n the bones lose (or are impaired in) their ability to
cond..Jctive hearing loss Hoo.r-
freely con vey (conduct) vibrations frmn the ty mpa nic me.mbra.ne to the oval ing loos caused by p roblems w ith the
wind ow. Such impairment occurs most often w hen the middle ea.r fills ""·ith brnes o f the mldlle ear.
mucus during ear infections -a condition known as otltis media. The oval otltls media lnftamrnatlon o f the
wind ow u s ua lly still vibrates under these conditions; but with out the a roplify- middle ear. commonly In chlldren as a
ing power of the ossides, h eari ng thresholds can be raised by as much as 50 result of infectlcn

286 CHAPTER
otosclerosls Abmmlal ol dB ?that is, sounds need to be SO dB louderin order to be heard). Thankfully,
1he middle-ear bones that c auses for th e millions of youn g children who s uffer ear infections, n ormal hearin g
hearing loss. returns after mucus is absorbed back into s tu·rounding tissues; however, this
sensortneural hearing loss Hear- reabsorption ca n take up to several m o nths.A moreserious type of conductive
ing bss due to defects n the cochlea loss, otosclerosls, is ca used by abnom1al growth of the midd le-ear bon es,
« audltO!Y ne!Ve.
most typicall y around the ova l window ne xt to the s tapes. Surgery can free
ototoxic R oduclng adverse effects the stapes from these bone gro\.'\ltl:1s and improve hearing.
m o::x:::htear or vestibular organs or By far the most commo n, and m ost serious, form of auditory impairment is
ne1V0s.
sensor1neural hearing loss, which m os t conunonly occu rs inside the cochlea
and sometimes as a result of darnage to the auditory nerve. Most o ften, senso-
rineuraJ hearing loss occurs when hair cells are injured. For example, certain
antibi otics and cancer drugs are ototoxic, meaning thflt they kill hair cells
directly. ln part, the reaso n for this effect is that ion pores on stereocilifl that
have been opened through mechanoelectrical trans duction are no t especially
selective abo ut \-1.:h at they perrnit to flow into the h air cell. Physicians are well
aware of these dangers and typically avoi d us ing s uch drugs, but sometimes
i.1. pa tient faces the decision of life with d ea fn ess versus no life at all. Vira l
infectio ns can also cause s udden onse t of hearin g loss in bo th children and
adults. ln other patients, hearing loss can be present birth, or it can appear
during a do lescence o r ear ly adulthood a nd progressively worsen over one or
m ore decades. Mutations in ove.r 150diffe.rent genes have already been lin ked
to hereditary h earin g loss in humans.
A more common cause of hearing loss is damage to the hair cells by exces-
sive exposure to noise. Those exquisitely fast and sensitive hair C"el ls are also
very vulnerable to da mage from excessive sound levels. It is fairly well known
that sh ooting a gun wi tho u t ear protection can ca use hearing loss. At leas t
since 1886, v. .·hen Scottish s urgeon TI1omas Barr tested the heMing o f Glasgow
boilermakers, we h.aYe known that extended exposure to lou d sound s s uch as
the noise o f factory el1uipment also causes hea ring loss (Keats, 2014). lt is no
coincid ence th at so ma ny aging rock s ta rs and race car dri vers wear hearing
aids. (It is ironic, but \·vise, that many heavy-metal mu sic fans now wear ear
protection at concerts.) And evi dence s u ggests that cumulati ve exposure to
even everyday n oises present in the environments of industrialized countries
ca n cause hearing loss. In one study (Goycoolea et al., 1986), middle-aged and
elderly residents o f Easter Island who s tayed almost exclusively on their quiet
island their whole lives were compared \o\:ith other Easter Islanders, who made
more frequent trips to th e noisier outside world . As Figure 9.24a sh ows, the
more time people spent off the island, the more hearing loss they experienced.
Hearin g loss is a natural consequence of aging fo r many peop le, and it
is difficult to separate a person's age from the amount of exposure to n oise.
Typicall y, age-related hearing loss first affects the perception of high frequen-
cies {Figure 924b). The 20- to 20,000-Hz frequency range fo r human hea ring
really applies on ly to ymmg people; by the time m ost of us reach college age,
we m ay have already lost the ability to hear frequencies above 15,(XX) H z. The
dec rease in the ability to hear hig her-frequency sou nds continues throughout
life, v..rith the highest audible frequency becoming lovi..-er and lower as we grow
older. Fortunately, many of the a uditory s timuli that people ca re most about,
including speech and rnusic, are composed p re dominantly of IO\ver frequencies.
TI1e ability to detect sou nds is n ot the sam e as the ability to listen to and
use sounds . Recently1 Hickox and Llberrnan (2014) showed how expos m e to
mo derntely high levels o f noise cha nges the ability to use sound even when
the ability to d etect smmds remains n o m1al. 1ltis "hidden hearing loss" has
been s hown to occur in mice following only one \veek o f exposure to 84-dB
noise (Maison, Usubuchi, and Liberman, 2013). This level is typica l of inany

(a) Effects of exposure to noise (b) Effects of aging FIG URE 9.24 Environrnental
affec ts hearing. (a) The more people are
/ Never left expo59d to noise in erwironrnent,
the more th9ir hearing deteriorat9S..
20 Easter Island who spent more
s years o ff of the relatively quiet island
had relatively worse hearing . f.P) Hear-
40
1 Aw')'3-5 -....-.
ing becomBS less sensitive with age,
partlcularty for hig h frequ9nci9s. (After
"'•"
·q 60
yea.rs
1
Away: more
than Syears
"'-.._
Goycoolea et aJ., 1006.)
;i:
SJ
'100
0.5
Frequency (kHz)
indus trial en virmun ents, and it is lower than sound levels that many young
peop le pursue at con certs and throu g h e ar buds. Unforh.mately, improving
the abil ity fo r listeners to de tect sounds is unlikely to improve the ability o f
people wi th hidden hearin g loss to unde rstand sounds .
Treating Hearing Loss

The earliest d evices fm helping people wi th hearin g loss were s imple ho ms.
The s maU end of the horn would be held a t the entry to the ear c.·mal, and the
w ide end \\.·ould be used to hmnel more acoustic energy toward the listener 's
ear. Although effective, these horns were obviously sornewhat cumbersome.
Elec tronic heming aids are much tnore con venient, but they must be desiqi1ed
to d o more than simply a mplify all sounds, because extremely louei sctunds nma Appa
(above 100 dB) Me just as annoying (or painful) for impaired listeners as they
are for listeners '"rith nom1al hearing. For a person w ho cannot hear sounds
until their intensity is <.'It least 70 dB, compared wi th about 0 dB for healthy
hea rin g, sounds th at co uld normally va ry be h veen 0 and 100 dB must be
squeezed between 70 <.'Ind 100 dB (Rgure 9 .25). Nearly a U modern hearing
aids use some means to amplify the s ignal vvhile also compressin g intensity
d ifferences to keep the highest intensities wi th in a com fo rtable listening level.
Very loud
Lo ud
- NonTu'1
- Impaired
----------------- -----------
LI-----------
FIG URE 9.25 When hearing thresh-
olds ar& inc r&a.Sed by impairrmmt. a
sound mus t have m ore energy to be
heard, but lncrnases faster
than it does with healthy ears. 'Whli!n
a hearing ak:t Is used to Inc rease the
Sp<E<Oh intensity o f soun ds, all variation in
components
sou nd levels must be com pr.;esed into
a smaller rangEi of intensity because
WfY loud sounds c an be just as i:alnful
Very soft for listeners. 1o11;1th heating impairment as
0 20 40 60 100 120 they are for with healthy hear-
Sound pr12ss ure (dB SPL) ing. {After C. D. Gels/.er, 1008.)

288 CHAPTER 9
Hea ring aids can a lso be tuned to provide the greates t a mplifi cation only
fo r frequencies in the region of grea test loss (fo r mos t people, higher frequen-
cies w ill need to be mnpli fied tnore) . An additional method is to move energy
from frequency regions in w hich hear ing is poor (usually high frequencies)
into regions w here hearing is nom 1al (A le xander, 2013}. Beca use the lower-
fre quency region is alrea dy being used for lmver-frequency sounds, this strategy
requires squeezin g toge the r .som e lo we r-freque ncy so unds to m a ke room for
the high -frequency sound tha t is being moved down
One ad vantage of the old horns over electronic hearing aids was that they
permitted listeners to direct their hearin g toward the sound source they were
most interes ted in . \ Ve m.ay thir1k about hearin g aids as a mpli fying the voice to
whid-1one is listening, but they also am plify all the other sotmds in the environ-
8230336 Am men The backg rotmd noise in a car, or even the sound of a refrigera tor, can
become lo ud enough to compete \vi th the sound of a person's voi ce. When the
entire ra nge of hearing is comp ressed fro m a range o f 100 d B to only 30 d B, a
10-dB difference between the nunbling of the car 's engine and the voice o f the
perso n in the passenger seat becomes compressed into only a 3-dB difference.
H e..u in g aid s are grad ually improving, and they have provided relief to
rnill ions of Americans, including for mer presiden ts Ronald Reagan and Bill
Clinton. However1 des pite researchers' rnany clever innova tions for improving
the signal th..'l t arrives at the ty mpanic membrane, da mage to the mechanisms
tha t trans.duce sound waves into ne ural signnls is p roving diffinilt or im pos-
sible to overcome completely. By analogy to vision, the best eyeglasses, contact
le nses, o r even laser s urgery ca nno t dum ge an im age enough to overcome
retinal degeneration. The best ad vice is to never need a hearin g aid. Protect
y o ur e..u s by avo iding exposure to loud soun ds an d by using heari ng protec-
tion such as ea rplugs Ol" mu ffs when necessary. U someone else ca n sing along
to the song you' re lis tening to o n your personal medi a player, turn it d m vn!
Electronic Ears
Mo dern medical .science and engineering are pro\'i ding some degree of
t-earing to many people i.vho are deaf. Cochlear prosthetics. more com-
mo nly known as ooc hlear implants (Fig ure 9.26a), are tiny fie>d ble ooils
v11tl1 aba.Jt two dozen miniature electrocle contac ts along their length.
Surgeons deli cately thread these electrode arrays through th9 round
wirdow as far toVi.ra:rd the apex of the rochlea as The elec-
trcde af'ra'.>' is connected to a ti n:I radio recetver und er the scalp, and
sig nals are transmitted from a small micro phone d evice on the outside
of the head b ehind t11e ear (Figure 9.26b). Signals com ing in from the
m icrophone act tvate the miniature elect rocles at apprcpriate positio ns
al ong the oochlear imp lant, which In turn stimulates associated auditory
nerve fi bers.
AJtho ugh cochlear implants are a mo dern medical miracle. they
cannot provide hearing that apprceches what nature provides. We are
rot surprised, how ever. that a handful of electrod es canro t replace the
funct ion of 14.000 hair cells. Some people benefit m ore than otllers
from Implants. and many adults 'Nith electric al hearing ca werse flaw-
lessly aver the pho ne. Young children, receiving implants as young as
age 1 or 2, do test of all b ecause young brains are particularly plasti c.
Children's brain circuitry develo ps to get the m ost infcrmatiai possible
from their electroni c ears.

(n) (b) / lvlicrophone .md tran.sin.itter FIG URE 9 .26 Cochlear implants give
som9 p 90ple who an;i dQaf th9 ability to
hear. (a) The cochlear implant ejeciroOO
array. The flexible array o f 91ectrodes
is insc;irted through the round window
as far as possible t oward the apex of
the cochlea (/::>) The 9lectrode array is
oonnected to a smaJI be·n eath
the scalp. and a small micropt-one and
transmitte< a re placOO over the rec9iv4"1t"
on th9 outsKJe of the head.
8230336 A
Summary
1. Sounds are fluctuati ons of pressure. Sound w aves are d efined by the frequen-
cy, intensity (amplitude), and phase of fluctu ations . Sound frequency and Refer to the
intensity correspond to o ur perceptions of pitch and lo udness, respecth.·· ely. Sensation and Perception
2. Sound is funn eled into the ear by the ou ter ear, made more intense by the Ccrnµink:Jn Website
middle ear, and transformed into n eural signals by the inner ear. sltes.slnauer.com/wolfe4e
3. 1n the inner ear, cilia on the tops of inner hair cells pivot in response to pres- for actMtles. essays, stu:Jy
sure fluctuations in ways tha t provide infomiation about frequen cy and inten- questions. ard other sttL!y aids.
sity to the auditory nerve and the brain. Auditory nerve fibers convey infor-
mation through OOth the rate and the timing patterns with which they fire.
4. Different characteris ti cs of sounds are processed a t multiple pl.aces in the
brain stem before information reaches the cortex. In.formation fro m both
ears is brought together very early in the chain of processing. At each stage
of auditory processing, including primary auditory cortex, neurons are
o rganized in rela tion to the frequencies of sounds (tonotopkally).
5. Hwnans and o ther mammals can hear sounds across an enomlous range of
intensities. Not all sound frequencies are heard as being eq ually lo ud, howev-
er. Hearing across s uch a wide range of intensities is accomplished by the use
of many auditory neurons. Different neurons respond to different levels of
intensity. In addition,. m ore ne urons overall resp::md when .sounds are m ore
6. Series of channels (or filters) process sound s within bands of frequency.

Depending on frequency, these cha1mels vary in hm ..• wide (m an y frequen-
cies) o r narrow they are. Consequently, it is easier to d e tect differences
beh·veen some freciuencies than beh\reen o the rs . \·Vhen energy from multi-
ple frequencies is p resen t, lower-frequency energy makes it relatively more
diificult to hear hig her frequencies.
7. Hearing loss is caused by damage to the bones of the middle ear, to the
hair cells in the cochlea, or to the neurons in the auditory nerve. Although
hearing aids are helpfol to listeners \\•ith hearing impairment, they cannot
restore hearing as \·vell as glasses can improve vision.

CHAPTER IQ

Hearing in the Environment
8230336 Amfrl a
Tl-E AUDITORY SYSTEM'S ABIUTY TO rnANSFORM TINY AR PRESSURE CHANGES

into a rich percep h u:tl world is a n a mazing wonder of bioeng ineering. From
the ftu-meling o f sound w.:tves by the p innae, to the mechan ics of middle-ear
bones, to the tiny p e rh1rbations of the basilar pa rtition a nd hair cells1 to the
sophisticated n e ural en coding in the brain s te m and cerebral cortex--so n1 e
remarkable m echanis ms have e volved to interpret a cous tic info rmatio n a bout
the world around u s.
Researd1 that reveals these Umer workin gs of the audito.ry sys tein typically
uses very s in1ple s timuli unde r constra ined s itua tions -often isolated pure
tones heard through headphones by listen ers sitting in an otherwise perfectly
quiet laboratory. Although these methods are invaluable for understa nding
how the a uditory sys te m functions, th.is is obviously not the wa y we experi-
ence sounds in our d aily li ves. In thls chapter we get "ou tside the head " to
investigi.1.te how hearing he lps us learn about the real world .
We start by looking at how it is p ossible to determine the location of a sound
source. In many respects, sound loca liz..."l. tion parallels visua l depth perception,
which you lei.1.rned about in Chaprer 6. \'Ve then him from lt41ere to what., how
percephml aspects of complex sounds are composed o f simpler sounds in 1nuch
the same '";ay that vi sual representations of objects are built up from siinple
feahues (see C ha pter 4). The third part of the ch i.1.pte r deals with auditory scene
(m alysis, whe re we will see why some so unds gro up togethe r, w hile separating
from othe rs, in ways that resemble the Gestalt principles introduced in Chapter
4. We see how the auditory .syste1n .seamless ly fills in gaps to form a cotnple te
and cohe rent "picture" of our ,wditory env ironrnent in an auditory analog to
visual object oa:lus ion (see Chapter 6). Pinally, we expl ore how auditory a ttention
has much in comm on w ith visual attention thi.1.t you learne d aho ut in C hapter 7,
w hile also serving a special role in keeping us ever vigilant for s urprises in \.Vorld .
Sound Loca lization

Suppose you were out ca mping in your local state park one mild s ummer
night, enjoying the las t e mbers of your campfire, when s uddenly an O\VI be-
ga n hooting somewhere in front o f you . You w ould instantly know w he ther
the owl was perched to the le ft, to the right, or directly behind the fire pit.
Moreover, if you were w illing to le.!l ve the com.fort of the fire, a nd if the owl
were cooperative enou gh to s it still a nd keep hooting., you could ea sily \.Valk
to the exact source of the sound e ven though you \vould.n 1t be able to see the
owl until you w ere very d ose to it.
When you think about it a bit 1 you w ill realize that this feat of auditory
localiza tion is quite different from d e te rmining the location of a v isual object.
If you could see the owl in front of you, you wo uld know that it was to the le ft

292 CHAPTER 10
::::::!:Vision
==:::::::. Sound
FIGURE 10.1 The position of the owl is easUy encoded by th9 visual system
because the owl's lmage falls on different parts of the retina (and thus activates dif-
ferent ree&ptors} depending on whethGr it Is to thQ left (A) or to the right {8) o f the
observer. In the auditory system, however, the same receptors are activated rggard-
k1ss of the owl's position.
or the right of your fovea because its image would appear on the right or left
side of your retina (Ffgure 10.1 ). But the owl 1s hoots enter you rears in exactly
the same place (funneled through the pinnae into the middle and inner ear)
regardless of where the mvl is.
You may recall that we face a similar dilemma when trying to determine
how far away a visual object is. As we saw in Chapter 6, visual depth percep-
tion involves processing and integrating a set of "cues"--stimulus aspects
that provide indirect evidence about how far away an object is. The auditory
system uses a similar approach to determine the location in space from which
a sound is coming. (See Web Activity 10.1: Auditory Localization Cues.)
8230336 Just as having two eyes turned out to be one of the keys to determining
visual depth relations, having two ears is crucial for determining auditory
locations. For most position.s in space, the sound source will be closer to one
ear than to the other. Thus, there are two potential types of information for
determining the source of a sound (Fjgure 10.2 ). First, even though sound
trave.ls fast, the pressure waves do not arrive at both ears at the same time.
Sounds arrive sooner, albeit very slightly, at the ear closer to the source. Second,
the intensity of a sound is greater at the ear closer to the sou rce. These are our
first two aud itory localization cues.
lnteraural time difference OTO) lnteraural Time Difference
The dlffa-ence In time between a
sound arrMng at one ear versus the let's first consider what we can learn from the lnteraural time difference, or
other. ITD. If the source is to the left, the sound will reach the Jeftearflrsl lf it's to the

HEARING IN THE ENVIRONMENT 293
Sound
FIGURE 10.2 Thg two 9al'S receive slightly different inputs w hm the sound sourcg
is locat9d to one side or the other. For frequancies greater than 1000 hertz. the head
blocks some of the mergy from reaching th9 oi:::poslte ear.
rig ht, it will reach the right ear firs t Thus, we can tell whether a sound is com- azimuth 1he angle of a sound
ing from o ur right o r le ft by determining which ear receiva the soWld first. The source en tl1e horizontal plane relative
tenn that is used to describe locatio ns on an imaginary circle extending around to a point In the center of the head
between the ears. Azimuth Is mea-
us in a horizon ml plnm--front, back, left, and right-is azimuth (Figure 10.3). ,eured In degrees, with o degrees being
A m ore detailed analysis o f the ITO can tell us even more. See if you can Straight 81-1. 111e angle lncre"""6
answer the following questions: \·Vhere would a sound source need to be clockV.1se to.Yard the right, v.1111 1oo
located to produce the maximum possible ITD? 'What location would lead to degrees bel'lg d rectly behnd.
the minimum possible ITD, and what wottld the ITD be in this case? Finally,
600
Measured values
FIGURE 10.3 lnterauraJ time differences for sound

sources varying in azimuth from directly In front of
lis teners (0 degrees) to directly behind (1SO dggr9QS).
t-\ow much variation there is in the time it tak.9S for a
sound to reach 90Ch ear dePQnds on where a sound
20' 40" 140" llD' 180'
comes from In spaoo. {Data from et al.,
Front Dlledion of sound source Back 1967.)

294 CHAPTER 10
FIGURE 10.4 lnteraural time diffg- 0°: ITD = Oµs

ences for different positio ns around - 2D'o ITD o -200µ• . -- 2Cl'o ITD = 200µ •
the head. Blue indicates locations from
w hich sound reaches the left 981' first:
red Indicates locations from which
sound reaches the right ear first. (Data
from FQdd gr son ot al., 1 Q57 . - W'o ITD=-480 µ, . • 60°: nD = 480 µs
I
-120"0 ITD = -480 µ; •

0 •
\
I
OO'o rro . 640µ .
- 16()'oITD = -200 µ• • _
1801;1: IID = 0 µs
what would happen a t intermediate loca tion s? Figure 10.4 illustrates the
answers to these ques tions. The ITDs fo r sounds coming from various angles
are represented by colored circles. Red circles indicate positions fr om which
a sound w ill reach the right ear before the left ear; blue circles sh ow positions
from w hich a sound w ill reach the left ear first. 111e size and brightness of
ead1 circle rep resent the m agnitude of the ITD.
As Figure 10.4 shows, ITDs are largest, about640m icroseconds (m illionths
o f a :second, abbreviated µs) w hen sounds come directly &om the left or directly
from the ri ght, althou gh this value varies somel'Vhat depending on the size
o f your head. A sotmd coming from directly in front of or directly behind the
listener p roduces an ITO of O; th e sound reaches both ear.s sim ultaneously. For
intermediate locati ons, the ITO \<Vill be somew here beh veen these two values.
Thus, a sound source located at an angle of 60 degrees will alw ays produce
an ITO of 480 µs , and a sound coming from - 20 degrees w ill always prod uce
an IID of -200 µs . That might n ot seem like m uch of a time difference, but
listeners can actua lly detect intera ural delays o f as little as 10 µs (Klumpp
and Eady, 1956), w hich is good enough to detect the an gle of a sound source
to within l degree.
TH E PHYSIOLOGY OF ITDS The portion of the auditory system responsible

for calculating ITDs obviously needs to receive input &om both ears. As we
saw in Chapter 9, binaural input enters almost every stage of the auditory n er-
vous system after the auditory nerve (see Figure 9 .20). As information m oves
upward throu gh the system, however, with e very additional synapse the tim-
ing between the tvlo ears is likely to become less precise. TI1e med Ial superior
olives (MSOS) are the first places in the a udi tor y system w here inputs from
b oth ears con verge (Figure 10.5), and s u re enough, firin g rates of neurons in
the tvLSOs increase in resp onse to very brief time differen ces between inputs
from the two ears of cats (T. C. Yin and Chan, 1990).
ITO detector s form their connections from inputs coming in from the two
medial olive (MSO) A ears during the first few months of life (Brand et al., 2002; Kapfer e t a l., 2002),
relay statrn In the brain stem where
Inputs from both ears contribute and d eveloping the ability to use ITDs to localize sounds depends critically
to detectlm of the lnteraural time o n hav ing experien ce w ith separate sounds coming from diffe rent places
difference. in space (Seidl and Grothe, 2005). The d evelo pmen tal sequence is simila r

To higher bl',Un centers
Cochlear Cochlear
Sound
\
AuditOI)' Br,1in stem
______. /
oochl"
Section
'
from pons
FIGURE 10.5 Aftier o nly a single synapsie in the cochlear nucleus, Information from
each ear travels to both the mt:ildlal superior olive (MSO) and the lateral supgrior oltve
tLSO) on each sKje of the brain stern. MNTB =medial nucleus of the trapiezoid bocfy.
to formation of binocular neurons in the visual cortex, a nd it probably has

a si mil ar cause. Jnterpretation of ITDs (e.g., assoc ia ting a given ITD w ith a
give n angle, as sho\o\TI in Figure 10.4) is critically dependent on the size of the
head. lf babies came ou t of the womb w ith the ir ITD LT1echanisms prewired
to the size of their infa nt heads, thei r sound localization abilities wo u ld
s teadily worsen as their head s grew during chi ldhood and ad olescence (H .
i:;,.J<jues et ai. 2011).
1
lnteraura/ Level Difference
The second cue to sou nd locali:z.ation is the lnteraural le vel differe nce, o r ILD, lnteraural level difference pLD)
in sound intensity. Sounds are more intense at the ear closer to the sound source The difference In leVel (Intensity)
bec1 use the head partially blocks the sound pressme wave from reaching the between a sa.md arrMng at one ear
versus the other.
opposite ear. The properties of the ILD relevant fo r auditory are
si milar to those of th e ITD:
• Sounds are m ore intense at the ear that is closer to the sound source,
a nd they are Jess intense at the ear farther away from the so urce.
• The ILD is largest at 90 a nd - 90 degrees 1 and it is non existent at 0
degrees (d irectly in front) and 180 degrees (directly behind).
• Between these two extremes, the ILD correlates wi th the angle of
the sound som ce, but because of the irregular sh.ape of the head, the
correlation is less p recise than it is wi th ITDs.
Although the general relationship betv.'een ILD and sound source angle is
alm os t identical to the relationship betvveen ITD and an gle, there is an impor-
tant difference between the hvo cues: the head b locks high-frequen cy sounds
mud1 m ore effectively than it d oes low-frequency sotmds. 1111.s is because the
lon g wavelengths of lm1,.·-frequency sounds 1'ben d around# the hea d in much
the same way that a large ocean wave crashes over a piling near the sh ore.
Thus, as shown in Figure 10.6, fLDs are g reatest for high- frequency tones,
and fLD cues work really well to d etermine loca ti on as long as the sounds
have highe r-freq uency energy. IL Ds are greatly reduced for low freq uencies,
becoming almos t nonexis tent below 1000 h ertz (Hz). Om inability to localize
low frequencies is the reason it d oes n ot matter where in a room you place
the low-frequency s ub woofer of your stereo system .

296 CHAPTER 10
FIGURE 10.6 lnteraural (intensi-

I fill)Hz ___.. -----
ty) differences LDs) for tones of differ-
9flt fn:iquencio9S prQS6nt8d at diff8rsnt
[ .:;::;:-::: I I I "7---">i
positions around the tiead. Note that
the bigg9St differences are for fr9<:1uen-
c i9S greater than 1000 Hz, at which
point the head crootes a shadow.
c urves are not syrnrn9trical toward
the fror1t and back, because of filter-
ing charact9rlstics o f the pinnru11. (Data
from Fedderoon et al., 1 Q57.}
82
200
{iJ'
Front Direction of .soL1nd
THE PHYSIOLOGY OF ILDS Neurons that are sensitive to intensity differen ces
behveen the two ears can be found in the lateral superior olives (LSOs) (see
Figure 105), which receive both excita tory and inhibitory inputs. Excitatory
connections to the LSOs come fro m the ipsilateral ear (that is, excitatory con-
nections to the left LSO originate in th.e left cochlea, and excitatory conn ections
to the right LS() come from the right cochlea). Inhibitory input.s come from
the contralaternl ear (the ear on the opposite s ide of the head) \oia the rnedial
nucleu s of the trapezoid body (IvlNTB).
Neurons in the LSOs are very sensitive to differences in intensity across
the hvo ears because excitatory inputs from one ear (ipsilateral) and inhibitory
inputs from the o ther ear (oontralateral) are wired to compete. When the sotmd
is m ore intense at one ear, connections &om that ear are be tter both at exciting
LSO ne urons on that side and at inhibiting LSO neurons on th e o ther side.
Cones of Confusion
If you examine Figures 10.4 and 10.6 for a bit, you should see a potential
proble1n with using ITDsand ILDs for sound localization: An ITD of-480 µs
arises from a smmd source that is located a t eithe r an angle of-60 degrees from
the line of s ight (ten o'clock in Pigure 10.4) or an angle of -1 20 degrees (eight
o'clock). Adding information from intensity differences d oes n ot h elp us here,
lateral s'-"er1or olive (LSO) A relay
station In the brain stem where Inputs because the lLDs for th ese two angles are also identical. If we also cons ider
frcm both ears contribute to detection the elevation of a .solmd source (how far above or belm"'· our h ead the soun d
of the k1teraural level difference. source is-a factor ·we've been ignoring up to now), we find th.at a given ITD

(a) FIGU RE 10.7 Elevation a::fds another

dimension to sound bcalization. "1)
Con.as o f confusion. lntero.ural time
differQn09S arc;. plott9d h9t"e
aqoss azimuth and QIQvation. A sound
wtth zero azimuth and ..*vatkrl ls
d irectty in front of a listener's head . All
? o f tl'lQ locations in spaca that shar9 th9
same cok>r prCNld9 exactly the same
ffDs. Rod contours p ottod beoooth
the color&d surface llk.Jstrat9 how al l thQ
locations on the red line give rise to the
same ITD. (Part b after Wghtman and
Kistl.;.r, 1QOO.)
(b)
or ILD could arise from any point on the s urface of a cone of confusion tha t
extends perpendicu larly fro n1 the left or rig ht ear (Rgure 10.7).
Although many books speak of a single cone of confusion, actually an infinite
nurnber of cones are nested inside one another. ln fact, the widest "cone" is
reall y a dis k extendin g frorn directly in front of you, up to directly over your
head , back to directly behind your head, and contimting to directly below you.
As strange as it rnay seem, this disc is the n'lost confusing of the cones. Cones
of confusion are rea l perceptual phenomena, not jus t theore tica l problems
for the a uditory systenL In the real world , thankfully, we need not hold our
heads in a fixed position . As soon as you move your head, the ITD and ILD of
a sound source shift, and only one spatial location M ii be consistent wi th the
ITDs and fLDs perceived before and after yo u move your head lFigure 10.8).
Pinna and Head Cues

Another reason why cones of confusion are n ot major practical fo r
the auditory syste m is that time and intensity differences are not the only
cone of confusion A region of
cues fo r pinpointing the location of sound sources. Take a look at one of your positions In space where all sounds
pinnae {or, more take a look a t a friend's pinna). You11 see that produ:Je the same tlrne and level
the s h ape of the pinnil is quite complex, \vi th lots of idiosyncratic nooks and (lnter<ilty) differences and ILDsJ.

298 CHAPTER 10
(a) (b)
,.,.--\-
---
pp
FIG URE 10.8 Evaluating ITDsand

ILDs frcrn t'WO different head positicris
helps In sound locollzatb n . (a) At first.
a sound souroo coming from the blue
frog will lead to an ITD and an ILD that
vvould also bg oon sistent with a sound
corning from tt-ie gret<l frog. qJ) lf the
listeni;;ir' s head Is rotated slightly coun- cramties (Figure 10.9). Remem ber that the pinnae hmne l sound energy into
ttirclockwise, the rTDJlLD will no longer the ear ca na l. Because of their complex sh apes, the pinnae humel ce rtain
be consistent with the location of thti sound freq u encies more efficiently tha n o the rs. In addition to pi nnae, th e
gr99n frog. Th9t"e Vvill still b9 an a mbi-
guity, hO"WWer. Now the rTDllLD w ill be s iz.e and s hape of the res t of the body,. especial ly the upper torso, affect w hid1
consistent w ith th9 red frog as wall as frequencies reac h the ear m os t easil y. Because of these effects, the intensity of
the blue frog, But only the blue location each frequency varies slightly according to the direction of the sound . This
Is consistent w ith both th€1 first and tM variation provid es us w ith auditory localization cue.
second s9ts o f lTDs and ILDs.
FIG URE 10.9 Pinna shapes vary

quite a lot betw'*'n people. Listeners
learn how their pgsonal pinnae affec t
how they hear sounds from diffeJent
places In tile en\'ironm€1nt.

(•) FIGURE 10.10 Directional transfer functions (DT Fs).

15 (a} DifferienCGS in intensity at eardrum for sounds
of varying frequency are p totted In this DTF for a single
p oint in space. (b) Thi9 s9fies of DTFs plottad hgr-e is
10
for the same azimuth, b.Jt at different elevations. (Part a
from Kistler and Wightman, 1992; b from Wightman and
5
-ll Jenison, 1005.)
:;
-5
m'la Appa
200 500 1000 200J 5000 10,000
Frequency (Hz)
(b)
20
s:
i
:; -20
Here's a simple example to illustrate h ow the effects of pinna and upper

body shapes are measured. Suppose you' re in an anechoic room, a room in
w hich the walls are padded so that very little sound enters from the outside
and very little sound bounces (reverberates) off the walls. The room is full of
speakers at many locations-up, doV1.m, and all around you. Tiny microphones
are inserted. inside your auditory canals, right next to your eardrums. Now
you can measure just how much energy from different frequencies actually
reaches your eardrums from different loca tions . Figure 10.10a s h ows the
measurements a t the eardnun in a similar experimental setup-in this case
for sounds played over a speaker 30 degrees to the le ft of a listener and 12
degrees up from the listener 's head. A lthough the am ounts of energy at all
frequencies were equally intense coming from the speake r, you can see that the
amounts of energy were not equally intense at the eardrum . Some frequencies
(e.g., 5000 Hz) had higher intensity when they arrived a t the eardrum; others
(e.g., 800 Hz) had less in tensity.
These changing spectral shapes across changes in elevation provide cues to
auditory loca lization . Figure 10.6 showed how rela tive intensities of different
frequencies change depending on azimuth, and illustrated that those curves
are not synunetrical toward the front and back, because of the filtering charac·

300 CHAPTER 10
directional transfer function (DTF) teristics of the pinnae. For the final piece of the puzzle1 look at Figure 10.10b1
A measure that describes how the which s hows the sol.md recorded by in-ear mkrophones when the speaker
pinna, ear canal, head, and tcrso
is rnoved up and dow n in elevati on. As you can see, the relati ve intensities
c hange the lnteretty of sounds Wth
diff€fent frequenc ies that arrive at eat:h
of diffe l'ent freq uencies con tinu ously change wi th changes in e leva tio n as
ear from different locations in space well as azirnuth. The sum totaJ of these intensity shifts can be measured and
(azi muth am elevation). combined to determ ine the directional transfer function (usually referred to
by its acronym , DTF) for an individual.
TI1e importance of the DTF in sound localiza tion is easily und ers tood if we
conside r the difference behveen hearing a concert live and listening to music
through a set of hea dphones. ]n person1 we perceive the so und of the French
horns as comin g from one side of the orch estia a nd the sound of the flutes as
comin g from the other sid e. But w he n we wear h ea.dphon es (especially the
type inserted directly inside the auditory canals),sounds are delivered direct ly
to the ea rdrums, bypassing the p innae. Auditory engin eers can use multiple
microphones to simula te the JTDs and JL Ds tha t result from the musicians'
different locations (the Bea tles were emly u sers of this type o f technology),
but DTFs are not sim ulated . As a resu lt, you m ay be able to ge t so me sense
of direction w hen listening to a concert tluou gh head phones, but the sounds
will seem to come from in sid e your skull, rnther than from out in the world .
The situati on is a kin to tha t of visu al depth perception. Pictorial cu es ca n give
a linUted sense of de pth, but to get a true pen:eption of three-dirnensionality,
'\.Ve really need binocular-disparity information th at we n onnally get only
w h.e n '"·e're seeing real objects.

Actua lly, just as stereoscopes can be designed to simulate binocular dis-
parity, it is possible to si mulate DTFs. Instead o f using h.vo camera lenses in
place of two eyes, tw'o microphones are placed near the e<udrums as described
earlier_Th:•n the sound so urce, s uch as a concer t, is recorded from these two
8230336 AmmJ Ap microphones. When this special stereo recording, called a 11>i.n a uraJ record -
ing," is played over headp hones, the lis tener experiences the sounds as if they
were back out in the '\.vorld w here they belong. Unfommately, however, every
set of pinnae is differen t (see Figure 10.9), so fo r this simul a tion to work, all
listeners need their own individual recordings.
Just as heads (and thei r corresponding lTDs and ILDs) grow to be larger,
ears grow and chan ge during development and <1.re ofte n s ubject to mutila-
tions of varying degrees (e.g., piercings). Research su ggests that listeners learn
about the way DTFs rela te to places in the en \oironment through their extensive
experience listeni ng to sOlmds, w hile othe r sotrrces of information, s uch as
vision, provide feedback about location (\Vighh:nan a nd Kis tl er1 1998). This
leantl ng through experience su ggests tha t children may update the way they
use DTF infonnation during d evelopment, and it appe..us tha t s u ch learning
ca n continue into adulthood. Hofman, Van Riswick, and Van Opsal (1998)
inserted plastic m old s into folds o f ad ults' p innae. As expected , listeners im-
mediately becarne much poorer at localizingsounds. But after 6 weeks of living
with these m olds in their ears, the subjects' localiza tion abilities had grea tly
improved. Somewh at these listeners a lso remained quite good
at loca lizing wi th thei r "old ears" w he n the molds were rernoved. It would
be interesting to know h ow well Leonard Nimoy (who played Spcx:k in the
origi nal Stnr Trek series) could localize sotmd s wi th his Vulcan ear molds. The
experience of listeners in this s tud y su ggests that switching between human
and Vu.lean pinnae every day may have b ecom e jus t a normal part of Nimoy's
auditory life.
the re are some limi ts to the a bility to a djus t to growing or
re1nolded pi nnae. Babies grow larger heads, and pin.nae really d o get larger

in old age. TI1ese larger ears do help o lder adults use lower-frequency cU mrta Appa
however, this improved ability to use lo\I\.' &equencies is insufficient to offset
th e effects of age-related hea ring loss, and o lder individuals are poorer at
localizin g e leva tion (Otte e t al., 2013).
Auditory Distance Perception

How d o listene rs know hov.- far away a sound is? Althou gh it is important
to know what direction a sound is coming fr om, n one of the cues we've
d iscussed so fur (ITO, fLD, DTP) p rovide mud1 infom1ation concerning the
dis tance behveen a listener and n smmd source that is much m ore thnn an
arm's length away.
The simplest cue for judging the distance of a sotmd source is the relati.l.>e
hlteusityof thesotmd. Because sounds becon-1e less intense with greater distance,
listeners have little difficulty percei vin g the relative d istances of two identi-
cal sound sources. For example, if you he..1 r a pair of croa king bullfrogs, the
louder croaks should be comin g from the closer frog. Unfortunately, this cue
suffers from th e sam e problem as reL1 tive size in depth perception. Interpret-
ing the cue requires o ne to make assumptions about the sound .sources that
may tum out to be false (e.g., the softe r-sounding frog m ight be very dose,
wi th its croaks muffled b y s urrolmding vegetation). In Chapte r 6, you saw
lots of rabbits w hen learn in g about the use of relative size in visua l depth
perception, and the problem in hearin g is much the same . At the end of this
section, we will return to the challenge of fig uring out just how loud {big) a
sound is perceived to be w hen distance changes. But first, we w ill continue
to consider dist:ince percepti on.
The effectiveness of relative intens ity d ecreases llttickly as dis tance in-
creases, beca use sound intensity d ecreases according to the Inverse-square
law. \ Vhen sound sources are close to the listener, a small difference in distance
can produre a relatively large intensity difference. For example, a sotmd thnt is
l meter away is more intense by 6 decibels (dB) th an a sound that is 2 meters
away. But the sam e 1-meter dilferen ce between smmd SOUJ'('es 39 and 40 meters
away produces an intensity change of o nly a frnction of 1 dB. As one might
expect from these facts, listeners are fairly good (i t using intensity diffe rences
to detem1ine d istance w hen sounds are presented within l meter of th e head
(Brrn'tga rt, Dmlach, and Rabinowitz, 1999), but listeners tend to consistently
unde restimate the distance to sound sources farther a\..,.·ay, and the amount of
underestimation is larger for greater dis tan ces {2.ahorik, 2002).
Intens ity works best as a di sta n ce cue when the sound sou rce or the
listene r is mo..,;n g. If a croaking frog s tarts h opp ing toward you, you will
know it because its croaks tvill become louder and louder. Listeners also get
some informati on about ho w far away a source is when they move through
the en vironment. This is because, in a manne r akin to m otion parallax in the
perception of visual depth, sounds that are farther mvay do not seem to cl1ange
direction in relation to the li stener as much as n earer sounds d o.
Another p ossibl e cue for auditory di stance is the sp1..">Ctml 1.xmtposition of
sorni.ds. T11e sound-absorbing qualities of air dampen high frequencie.!! m ore
than low frequencies, so when sound sources are far away, higher frequencies
d ecrease in en e rgy m ore than lower frequencies as the sound waves travel inverse-s(J.Jare law A prin:lple
from the source to the ea r. Thus, the further away a sormd source is, the ''mud- stath)J that as distance from a scurce
dier" it sounds. This change in spedral composition is noticeable only for Increases. Intensity decreases faster
such that decrease In Intensity is equal
fairly large dis tances, greater than lOCXJ meters. You experience the change in to the distance squared. This general
spectral composition when you hear thunder from near your v..-indow . or from law also applies to optics ard other
far away. You hear th\mder as a loud ''crack# n earby, but thw1der from farther forms of erargy,

302 CHAPTER 10
......
/ ........, ______ _
./
.·/
l=IGUR E 10. 11 The relatll/Q amounts of direct and rev9rb9rant energy coming from
the listener's neighbor and the singer wHI Inform the llsten&r about the relative dis-
tanC9S of the two sound sour09s.
away sounds more like a ''boom." Note that this auditory cue is analogous to
the visual depth cue of aerial perspective (see Chapter 6i aerial perspective
involves that fact that more distant objects look more blurry).
A final distance cue stems &om the fact that, in most environments, the
sound that arrives at the ear is some combination of direct energy (which
arrives directly from the source} and reverberant energy (which has bounced
off surfaces in the environment). The relative amounts of direct versus reverberant
energy inform the listener about distance because when a sound source is dose
to a listener, most of the energy reaching the ear is direct, whereas reverberant
energy provides a greater proportion of the total when the sound source is
farther away. Suppose you're attending a concert. The intensities of the musi-
cian's song and your neighbor 's whispered. comments might be identical, but
the singer's voice will take time to bounce off the concert hall's walls before
reaching your ear, whereas you will hear only the direct energy from your
neighbor's wlllipers (Rgure 10.11 ). (See Web Essay 10.1: ReverberaUons
and the Precedence Effect.)
As it happens, reverberations appear to be important for judging the
loudness of sounds. You do not judge that a coyote is howling softly just
because it is far so how do you estima te how loud the howls really are?
Recall that the sound to which you are listening rapidly decreases in energy,
following the inverse-square law. However, reverberations do not fall off
so quickly, because the surfaces that sounds bounc:e off do not move when
the sound source becomes closer or farther away. Following a clever set of
experiments that allowed them to dise.ntangle listeners' use of direct sound
energy and reverberant energy, Zaho:rik and Wighhmm (2001) suggest that
listeners maintain constant perceptions of loudness across d1anging distances
by scaling direct energy relative to reverberant energy.

Complex Sounds
Simple sotmds like sine waves and b ands of noise are ve ry useful for explo r-
ing the fund a mental op erating characterfsticsof a udito ry systems, ju st as sin e
wave g ratings a nd sin g le-waveleng th lig ht sources are essentia l tools for vi-
sion researchers. But pure s ine wave to nes, like pure sin g le-wavelength lig ht
sources, are rare in the rea l '"'·orld, w here objects and events tha t matter to
listeners are m ore cotnplex, m ore interesting.. a nd therefore more chal lengi ng
for research ers to s tudy.
Harmonics
Many environmen tal sounds1 includin g the human voice and the sounds of fundamental frequency The ICM'-
m.uskal ins trumen ts, have harm onic structure (Figure 10.12 ). h1 fact, harmonic est -frequency component of a com-
sounds are a mong the m ost con.u11on typ es of SOlmds in the en vironment. TI1e plex pencxJlc sound.
lowest frequen cy o f a harmonic spectrum is the fundamental frequency. With
na tural \·ibra tory sources (as opposed to pure tones prod uced in the laboratory),
th ere is also energy at freq uen cies that are integer multiples of the hmdamental
freq uen cy. For example, a fe.m::ile speaker may p rod uce a vowel so und \vi th a
hmdam en ml fre que ncy o f 250 Hz. Th e vocal cords wi ll prod uce the greatest
energy at 250 Hz, less ene.rgy at 500 Hz, less s till a t 750 Hz, less at 1000
Hz, and so on. ln this case, 500 Hz is the second ha nnonk , 750 is the third, and
1000 is the fo urth. For har moni c complexes, the perceived p itch o f the com-
plex is determin ed by the fundamental frequen cy, and the ha rmonics (often
called "overtones" by musician s) add to the perceived rk hness of the sound.
The a uditory system is acu tely sensi tive to thefiltur.:ll relationships beh'''een
harmon ics. [n fact, if the fi rst harmonic frequenfy) is-rem oved
from a ser ies of har monics, as s how n in Figure 10.13, and only the others
(second, third, fo urth, and so on) are p resented, the pitd1 that lis teners hear
corresponds to the fund amental frequen cy---even tho ugh it is no t part of the
sound l Listeners hear the missing ftmdamen tal. (See Web Activity 10.2: The
Missing-Fundamental Effect.) It is not enm necessary to have all the other
ha rmonics present in order to hear the missing fundam ental; just a few w ill do.
The most s traightforward explan ations of the m.is.sin g-hmdarnental effect
involve the tem poral cod e for pitch d iscussed in Chapter 9. One thing th.a t
Fundamental
frequ ency
FIGURE 10.12 ManyenvifonmentaJ sounds.

including vob9s, arG harmonic. The lo¥Vest fre·
H.u monics quency o1a hannonic sound is the fundamental
frequency, and there ar9 p eaks of e ne rgy at Integer
Frequ""')' (Hz) multiples of tM

304 CHAPTER 10
..
=- - - -Missing fwtd.amental
al l harmonics of a funda men.tal h ave in comrnon is
fluctu,'\tions in sound pres.sure at regular intervals
cor responding to the fond am en tal frequency. For
exa rnp le, the waveform for a 500-Hz tone has a
peak every 20 mil liseconds (m s) (Figure 10.14b).
The wavefom1s for 75()..and 1000-H z tones have
peaks every 1.3 and 1.0 ms, respectively (Figures
10 .14c and d ). As sh o\o\Tt in Figure 10.14e, these
three waveform s com e into aligmnent eveiy 4 ms,
w h.ich,conveniently, h appen.s to be the period of the
Ji.mdamen ta l for these three hnrm onlcs:
250 Hz. lndeed ,eouy Ho rn1orudoi2S0H z Wi!L have
an energy peak every 4 ms. Some neurons in the
auditorv nerve and cochlear nucleus '"'ill fire action
potentia'.isevery 4 ms to therollec:tion of \Vaves sh mvn
in Figure 10.14e, providing an elegant mechanism
2 3 4 5 6 7 8 9 10111213 1415 16 1718 19 20 21 22 23 24
Hammnics to explain why listeners perceive the pitch of tlus
- - - - - - - - - - - - - - - - -.. 600{)
complex to ne to be 250 Hz, even though the to ne
Frequency (H z) has no 250-H z component.
FIGURE 10.13 If the fundamental (lowest frequency) o f a har-
rronic sound ls remov'9d , listQflefs still t\Qar the pitch of this "'m issirrg Timbre
Loudness and pitch are reJ ati vely easy to d escribe,
bee.:,use they correspond fairly well to simple acous-
tic d imensions (arnplitu d e and frequent..)', respec-
ti vely). Bu t the richness of co mp lex sow1ds Hke
those fo und in our world depends on more than
(a) I (b)
I
V M.issing fundament,1-l
:i!
Ii> (r)
""] :i!
"
; l AOAAOAAAAAAOAAO
Harmonics
- - - - - - - - F,. en-o-.v -(H_z)_ _ _ _ _ _._ 6c:m
-q-u- (<)
FIGURE 10.14 When only thrM harm onics of the same fun-
damental freq uency are presented (/:>-dl, listeners still h ear the
pitch of the mlssing-fundrun.:rrtaJ frequency (a) because the
harmonics share a common B"lergy ftuctuation 4 rns, the
perbd o f a 260-Hz signal (e). 4ms

simple sensations of loudness and pitch. For e..xarnple, a trombone and a ten or timbre The psychol::glcal sensation
saxoph on e might play the sam.e note (that is, th ei r hvo notes will have the by which a listener can Judge that two
exact sa m.e fundamental frequency) at exactly the same lm.idness (the sound sounds with the same loudi1ess and
pitch are dissimilar. Tlrnbre quality Is
waves will have identical intensities), but we would have no troubl e d iscern- con'leyed by harmonics and other high
ing that h vo different instnunents \Vere being played.11-le percephtal quality frequencies.
that differs between these ti-1;·0 rnusical instruments, as well as v°'wel
sotmds s uch as those in the words lrot, heat, and hoot, is k:no\oVTl as tinibre. (9ee
Web Activity 10.3 : Timbre.)
What exactly is titnbre? You won 't find a good answer to this questi on
in the dictionary, beca use the official definition of timbre is /(the quality that
makes listeners hear n.vo different sotmds even thou gh both sounds have the
same pitch and loudness" (Am erican Standards Association, 1960). However,
differences in timbre between musical instnunents or vowel sotmds can be
estima ted closely by of the exten t to which the overall spectra of
two sounds overlap (Plomp, 1976). Perception of visual color depends on the
relative levels of energy at different wavelengths (see Chap ter 5), and very
si milarly, perception of timbre is related to the relative energies of different
acoustic spectral components (R g ure 10 .1 5). For example, the trombone and
tenor saxoph one notes \vhose spectra are plotted in Figure 10. lSn share the
same fundan1ental frequency (middle C, 262 Hz). However, notice that the
trombone's third (786-Hz) component is s tronger tha n its fourth (1048-Hz)
component, whereas for the saxophone, the relationship between the energies
of these h<\o·o components is reversed.
FIGURE 10.15 Timbre. (a) Three dlffererl: musi·

cal instrurnents playing the same note (middle C.
262 Hz) sound diff8f9flt b9causei they h ave different
spectral shapes.(/:>) producoo byth.:i
6000 600J
same female talker, also with a fundamental fre·
Frequency {Hz) quency of 262 Hz. sound very 10 listeners.

306 CHAPTER 1 0
Auditory "Color" Constancy

Once we take off our headphones and get on out into the ·world , listen-
ing environments can dramatically alter the sounds that anive at our
ears. ·we already learned ab::>ut the importance of revert:eratlcn energy
for perception of distance and lcudness of sounds. Even the way you
da:::crate can charge :sounds on their way to your ears. Higher frequen-
cies are reinforced by hard surtaces such as tile floors and concrete
walls, but they are dampened by soft surtaces such as thick carpet
and curtains. Thls is much like the problem of oolcr constarcy that you
learned about in Chapter 5. In vision, the goal Is to perceive the same
colors even thOLtJh the spectrum of illurnlnation can be quite different
depending on the type of lighting (sunlight, incandescent. and so on).
In hearing, sutiaces in the en\'ironment refle:::t and absorb energy at
different frequencies In W"fS that change the spectral shape that finally
anives at you r ears.
Klette and Kluender (2008) used the different spectral shapes
of the vowel sounds ·ee· and ·oo' to learn how heating calibrates for
changes In the listening environment. They took advantage of the fact
that the spectra for 'ee ' and 'oo' differ in t\.vo lmp:>rtant '>N'iflS (Fjgure
1 0.16a). First. the vowel sound ·ee' has a relatr.telyftat spectrum with
almost as much energy In high frequencies as In low frequencies (upper
lefi in the figure). In cmtrast, energy in 'oo' is dcrnlnated by low fre-
quencies w ith less and less energy as frequency increases (lower right).
We can S"f that the "tilt" of the ·ee' spectn.rm is much flatter than the tilt
of 'oo. · Second, the sounds 'ee' and 'oo' also differ In where peaks are
present ln their spectra. The second peak In the 'ee' spectrum Is at a
hi gher frequency than the second peak in the ·oo• spectru m . Listeners
use both tilt and the frequen:;y of this second peak to tell ·ee' from 'oo'
(Figure 10.1 Gb) (Klefte and Kluender, 2005).
To learn how the auditory system adjusts for listening context,
Klefte and Kluender (2008) created stimuli that enabled them to sepa-
rately measure how much tilt and second -peak frequency cmtr1bute to
the perception of these vowels. Then they had listeners Identify vowels
after healing a sentence like Myou w ill now hear the vows." but an
interesting tv.;st. They created some sentences so that the overall tilt
of the sentence was the same as tl1e tilt of the foUowirQ vowel, either
'ee'-like or 'oo'- like. To other sentences they added a peak in the spec-
trum, all the way through the sentence, that matched the frequency of
the second peak in the vowel ('ee' or 'oo') that listeners would Identify.
Usteners heard the very same vowels in dramatically different ways de-
pending on vl'hldl type of manlpulated sentence the vowels.
When tilt ·was the same for both the p-eceding sentence and the vowel,
listeners used only the frequency of the second peak t o identify the
vowel (Figure 10.16c). \M"ren the second peak was present all the way
through th3 p-eceding sentence, listeners relied mostly 01 tilt to identify
the vowel (Figure 10.16d).
Listeners calibrate for reliable spectral charactertstics of a listen-
8230336 Amma Appa ing caitext much as observers calibrate for the spe::;trai oomposltion of
illuminating light when percei.,;ng color (Alexander and Kluender, 2010 :
smp et al., 2010). IM"ren the brain int erprets spectral tilt as a conse-
quence of the em'ironrnent in vJhlch a sound is heard (concrete versus
curtains), listeners igncre tilt and use only t he seoond peak. Conversely,
when there seems to ah.'v'3fs be energy around the second peak in a
listening emironment , listeners use only tilt to decide w hich vo·wel they
heard.

FIG URE 10.16 Listeners use both

spectral tilt and the frequencies o f
spectral peaks to identify vowels. {a)
SPQCtra o f natural ' 9€1' and ·oo' are
shown In th<> t.l>ll'l' lo-
a oq
right, respectively. The vowel 'ee' has
(l .... ·oo•
.....
a relatively flat spectrum. a"ld 'oo' has
a tilted spQCtrum, SLCh that there is
muc h less energy at higher freq.JQn·
cies. The second spectral pgak. is
lower in fr9QU9"C)' for 'ex>' than for 'ee' j
"' "' "'
'lile.' ExpGTimGntally manfpulatOO 20 20
vowel sounds are also stown with 10 10
the tilt of ·ee' and second peak of 'oo' Hybrid:
in the Llppeir light and the tilt of 'ex>' Natural 'err! 'el!! tUt, 'oct pe<1k
-10 -10
and second peak of ·ee' in the lower
left. (b) Listgners use both tilt and fre· Tilt 0 1000 a:ro 300:> 4000 .scoo 1000 axo 30CO 4000 .scoo
.
quency of the second peak to identify
vow9's.. Lighter ar9aS correspond to
sounds that subj.:cis Mar as 'ee';
"°50 "°
"1
darker areas represent sounds that :s 40 40

listeners h'11ar as 'oo. · (c) Upon hearing
a vowel atte< a sentence that has the
·oo' ] "' "' ....
20 20
s ame tilt as the vowel, listena-s use
only the frequency o f the second peak
to identify the vow9' . lp') Ho\11/ever,
j 10
Hybrid:
•OO• tilt, 'et!! peak
10
NaturaJ 'od
when the pr9C9ding sentGonca Includes -10 -10
n:o
a peak at the samg frequency as tt1e
second peak in the VO'N91, IBteners
1000 200)
"""
Frequency (Hz)
4000 5000 1000
Frequency (Hz)
?(.(() 4000 5COJ
use mostly tilt to identify thQ \'OW'QI.
(b) Tes! vowel (c) Test vowel after sentet\C2 (d) Test vowelafta- senteru:ewithsame
in isolation with same tilt frequency
-2.4 -2.4
-3.9 -3.9 -3.9
-5.5
- 7.0
-<1.5
0.1 0.3 0.5
"----- - l0.1
1400 1600 1800 :a:oo n:o

Frequency of second peak('Hz\
Attack and Decay

Another very important quality of a complex sotU"\d is the way it begins (the
attack of the sound ) and ends (the sound's decay) (Rgure 10.17). 0m a uditory
systems are very sensi tive to a ttack and decay characteristics. For example,
important contrasts be tween sp eech sow1ds in the words bill a nd tcill or the
words chip and :hip rela te to differe nces in hmv quickly sotU"\d energy increoJses
at the onset (the rate of attack). attack The part of a sound duri ng
The sam e musica l ins tnunent can have quite different a ttacks, fro m the Which arnplltude Increases (onset).
rapid onset of a plucked v io lin string to the gradual onset of a bowed s tring decay The part of a sound durlrg
(Figure l 0.l 7a and b). How quickl y a sound decays depends on ho w long which amplitude decreases (offseQ.

Amma Appa
308 CHAPTER 10
FIGURE 10.17 We use th9 onsets of (•) Violin (pluck) (b) Vmlin(bow)
sounds (attacks) to id9ntify thsm. 1h9
attack is quit8 diff9f'gnt wMn a \'lolln
s tring ls plud<ed (a) versus bowc:1d «>).
Similarly, different speech sounds, such
as ·ba' {c) and 'wa' (d.I. have diff9rMt
attacks.
(c) Spmh ('ba') (d) Spe•oh ('wa}
it takes fox the vibrating object creating the sound (e.g ., the violin string) to
dissipate energy and stop moving. One of the more challenging aspects of
designing music synthesizers that could mimic real musical instruments was
learning how to mimic the attacks and decays of the instruments.
To appreciate the importance of attack and decay in sound quality, explore
Web AetMty 10.3: Timbre, where you can hear a normal piano note and then
hear the piano note played backward . The sound of a piano note is caused
by a small hammer hitting a string. The amplitude of the resulting sound
increases very quickly before more gradually dissipating. If a recording of a
piano note is played backward, the sound no longer even remotely resembles
that of a piano. Instead, the backward piano note sounds more like the same
note played on an accordion.
Auditory Scene Analysis

The acoustic environment can be a busy place. In most natural situations, the
so\md source that one is listening to is not the only source present. Consider,
for example, conversing with a friend at a party where many other people are
talking, music is playing, chips are being munched, the door is being opened
and closed, and so on. Now consider simpler environments, such as those
you choose for studying. To read this chapter, you probably chose the spot
where you are right now because it was relatively quiet, but stop and listen
carefully. Is a heater, air conditioner, computer, refrige.rator, or some combi-
nation of these devices hrnnming in the background? Can you hear people
talldng somewhere in the vicinity? Are chair legs sliding across floors? Places
with only a single sound source are very uncommon. Indeed, there are few

truly quiet places outside the laboratories of hearing scientists and the testing source segregatloo or auditory
chambers of audiologists. scene analysis Processng an audi-
Environments with multiple sound sources are the rule, not the exception . tory scene consisting of multiple sourd
sources Into separate eound Images.
How does the auditory system sort out these sources? Note that the visual
system also has to contend with a busy w01ld, but eyes can be directed to any
part of the scene that is of interest to the visual system. Moreover, the rods and
cones on the right side of the retina always see the objects on the left, politely
leaving the receptors on the other side of the retina to collect data about objects
on the right. For an auditory scene, the situation is greatly complicated by the
fact that all the sound \vaves from all the sound sowces in the e n vironment
are summed together in a single complex sound wave (Figure 10.18). You can
m.ove your ears around all you want, but everyone's voice at the party still has
to be picked up by the same two sets of cochlear hair cells. Separating different
sounds from one another is like living in a world in which everything is made
of glass: it is difficult to distinguish separate objects, because they all merge
into a single combination of shapes, as Figure 10.lSe illustrates.
Someh ow, however, the auditory system contends quite well with the
situation: our perception is typically of a world wi th easily separable sounds.
We can understand the conversation of a dance partner a t a party, and we can
pick out a favori te instrument in the band. This distinction of auditory events
or objects in the broader auditory environment is commonly referred to as
source segregation or auditory scene analysis.
Spatial, Spectral, and Temporal Segregation

The auditory system uses a number of strategies to segregate sound sources.
One of the most obvious strategies is spatial separation between sounds.
Sounds that emanate from the same location in space can typically be treated
as if they arose from the same source. Moreover, in a natural environm.ent in
whid1 sound sources move, a sound that is perceived to move in space can
more easily be separated from backgrmmd sounds that are relatively stationary.
(a) Frog (b) Bll'd

FIGURE 10.18 Separating sounds
from one anoth9r is living in a \'isual
wol1d made o f glass, because all the
waveforms from a ll the sounds around
us are summed Into a single wavefam
arrMng at the ears. Here the sounds
ar9 a frog, a bird, and a splash, sho..,.,-i
both separately (a-c) and as they occur
togethgr at the ear ¥1). In this Wlif'f,
scunds are transparEnt 111<9 the glass-
ware In (e). (Part e oourtesy o f Andrea
Kluender.)
(c) Splash (<f) Frog+ bUd + 'Pl..h (•)

10336 Amna Appa
310 CHAPTER 10
FIGURE 10.1 9 WhGn t ones that (•) (b)

are dose in frequoocy ocx:ur In rapid
suocQSSion , thgy are heard as a single
wartjing stream (a), but whQn succes-
sive. rapidty alternating ton 9s have very
d ifferent frequencies, thgy are heard as
tVY'O separate streams (b).
!
• • • •y,,,,.• • • • • • • • • •
Tm,.
auditory stream segregation The Listeners m ove too, so if a sound stays in the same place rela tive to the path of
organ ization of a complex a listener, it w ill be easier for that so und to be sorted out frorn other soun d s.
acoustic signal Into separate auditory In addition to location, sounds ca n be segrega ted on the basis of their
events for which each stream Is heard
as a separate event.
spect ra l or tem pora l quali ties. For example, sounds w ith the sa me p itch or
.si mila r pitches are m ore like ly to be treated as coming &om the sam e sotuce
nnd to be segregated from other sounds. Sounds tha t are pe rceived to emana te
from the sam e sotuce are o ften d escribed as being pa rt of the same uauditory
s tream ,'1 a nd divid in g the a ud itory world into separate au ditory objects is
kn own as auditory s tream segre gation.
A very sim p le examp le of au di tory stream segregation involves hvo tones
""-ith s imilar freq uencies tl1.:1t are alternated (see Figure 10 .1 9a and WebActtv-
ity 10A: Auditory Stream Segregation). 111is sequence sounds like a sing le
coheren t stream o f tones tha t warble u p an d down in freq uency. Bu t if the
alternating tones a re m a rkedly di fferent in frequen cy (R g ure 10.19b t two
streams of tones are heard-one higher in pi tch tl1an the othe r {G. A. Mille r
and Heise, 1950).
Auditory stream segregation is :a powerhJ percep hial phenom enon tha t
is not lim ited to si mp le tone:s in the laboratory. Before strea m segrega tion
was "discovered " by audi tory scientists, th e com poser Johann Sebastian Bach
exploi ted th ese a uditory effects in h is compositions (Figure 10.20). Th e sam e
s uch as a pipe organ, would rapidly play interleaved sequences of
low and high n otes. Even tl1ough th e musician played sequence in tl1e order
*bm1· ¥ CUfijt£r!(ff tld

*b • dP tft! t&F ,,.
FIGURE 10.20 This mu sk;: al
seq uence from J. S. Bach's ''Toc-
cata and Fug ue in D Minor• utilizes the
s tream segrtigation prlnci p es •discov-
a tf6 trd ti •trtr1
f ti
ered" by auditory resoo.rc hec-s in the
twentieth centu ry. V\lhen played in rapid
suC09SSlon , the higher no tes (redj are
heard as one mebdy separate from the
lc;iv(Qr not es IJ::ilue).

Hl-Ll-H2-L2-H3-L3, listen e rs heard two m e lodies>-one high (Hl-H2-H3)

and one low (Ll-L2-L3). (See Web Activity 10.4: Auditory Stream Segrega-
tion.) Thi s bit o f knowledge about auditory pen:eption was known not only
to Baehr but also to other Baroque composers of the seventeenth and early
eighteenth centuries.
When th.inking about auditory scene analysis, it is sometimes useful to
describe effects using Gestalt principles s uch as those elucidated for vision
in Chapter 4. The examp les of auditory stream segreg'1tion presented in this
secti on can be described as applications of the Gestalt principle of si milnrify
(see Figure 4.1811 in Ch.:1p ter 4): sounds that are s im.ilar to e,1ch other tend to
be grouped toge ther into streams.
Grouping by Timbre
When a sequence of tones that have increasing and decreas ing freque ncies is
presented (Figure 10.21a), tones that deviate from the rising/ falling pattern
are heard to 11pop out'' of the sequence (Heise and Mill er, 1951). Wha.t happens
when two patterns over lap in frequency- one increasing and then decreasing
(•)
••••• •••••
•• •••
!
••
•••
•••••
••••• •••••••••••••••
.. ...
. .
(b) Same timbre (c) Different timbres
-. -.... ••••• .
. . ..
..." "'
"
""'
.1.. l·. ...

• •• •• •• •• ••
•• • Tune
• • Time
•
FIGURE 10.21 Timbre affects how sounds are grouped, (a) tndividuaJ tones (gr9€<1)
Mp::>p of a stream if they do not fit the patterns of rising or falling fr9Quency for
the othg- tones (red). f/;i, c) Sounds that sh31'e the same timbre grcup togi::ithtf. \<Vhi<m
sounds in a succession alt share the sarne they are heard as streams
aocordirg to similar fr.equ0ncy. But if sounds have diffGfent timbres (c), thQY will S9pe.-
rate aoccrding to tirnbr&, even if their frequenc ls:is cross another pattern with a differ-
Elflt timbre.

312 CHAPTER 10
in frequency, and on e decreasin g and then increasing in frequency (Figures

10.21b and c)? If the tones a !'e s irnple sine waves, t\vo streams of smmd are
heard v.i thout over lapping pitches (Figure 10.21 b); one s trea rn includ es all
the high tones, and one includes all the low tones. However, if ha rmonics are
added to one of the sequences, thus creating a richer timbre, h,,,:o overlapping
patterns are hea rd as distinct (Figu re 10.21c) (van Noorden, 1975).
Auditory stream segregation ba sed on groups of no tes ,vith similar tim-
bres ca n be seen as a no ther ex.a mple of the principle of similarity a t work. As
Figure 10.16 illustrated , timbres are very different fo r pianos, trombones, and
saxophones. Grouping by timbre is pa rticularly robu st becau se sounds wi th
similar timbres usually arise from the same sound source. Th is principle is
the reason w hy listeners are able to p ick out the melody pl::1yed on a single
ins trument-a trombone, for ex..1mp le---even when another instrument, such
as a saxophone, plays a differen t or even op posite sec1uence o f notes(\ Vessel,
1979).
Neural processes th.'"lt give rise to s tream segregntion can be found throu gh-
out the a uditory sys tem, from the first stages of auditory p rocessin g to the
primary auditory cortex (A 1) to secondary areas of the audi tory cortex, sud1
as the belt and parabelt areas Q. S. Snyder and A lain, 2007). TI1e brain s tem
s hows neurnl evi dence of stream segregati on based on si mple cu es such as
frequencies of tones, but segregation based on m ore so phis tica ted perceptual
properties of sounds is m ore like ly to take p lace in the cortex.
Grouping by Onset
In add ition to timbre, sotmd components that begin a t the same time, or ne.nrly
the same time, s uch as the ha rmonics of a music or speech sotmd , \viii also tend
to be heard as coming from the same so und source. One way this phenomenon
helps us is by gro uping differen t harm onics into a single complex so und. Fre-
quency components w ith different onset times less likely to be grouped.
For example, if a sin&e harm onic o f a vowel so w1d begins before the res t of
the harmonics of the vowel, that lone harmonic is less likely to be perceived
as part of the vowel if the onsets are sim ultaneous (C. J. Darwi n, 1984).
R. A. Rasch (1 978) shov.-ed thnt it is much easier to distinguish two notes
from one an other w hen the onset of one precedes the onse t of the other by at
least 30 ms. He n oted that musicians playing together in an ense1nble such as a
string qu artet do not begin playing no tes at exactly the same time, even w hen
the musical score instructs them to d o so. Jn.s tead, they begin n otes s lightly
before or after one an other, and this s taggered start probably helps listeners
pick ot1t the individua l instruments in the gro up . Part of the signa ture sty le
of the Rolling Stones h as been to carry th is practice to an extreme. Members
of the grot1p some times begin the sam e bea t wi th s t1ch widely varying onsets
tha t it is unclear whether they are playing toge ther or not.
Grouping of sounds with comrnon onsets is consistent with the Gestalt law
of com mon fate. Consid er the consequences of dropping a bott le on a hard
smface and listening for whether the bo ttle b reaks. When any object botmces,
each bounce resu lts in a set of overtones tha t relate to the size, shape, and
8230336 Amm1 nMterial of the object. lf the dropped bottle d oes not break (Figure 10.22a),
this patte rn of overtones rep eats as a group, and the intens ity of the sotuids
decreases wi th every additiona l bounce. But if the bottle breaks upon landing
(Figure 10.22b}, indiv idual shards of the bottle ,.yi_l! have different spec tral
compositions, each sh ard bein g its resonator. Onsets of the overtones for
different shards will di.ffer beca use the pieces bounce independently until they
bounce no m ore. TI·ms, even w hen the initiaJ burst of noise is removed from
the sotmd of the bouncing a nd breaking bo ttles, listeners can use pa tterns of

(a) Bouncing
I
Intensity
TlITle
FIGURE 10.22 Spectrograms of a bottle bouncing (a) or breaking (p). VYhen the
bottle brooks. there are multiple patterns of o nsets for multiPe pieces of glass. Lis-
teners c.an use o f onsets to aco.Jrateo/ detiermlne \l\'hether or not the bottle
broke.
on.sets to acc urately determine whethe r the bottle b roke QN. l-1 . Warren and
Verbrugge, 191\4).
When Sounds Become Familiar

In additi on to the simple Gestalt principles that we' ve already d iscussed, listen-
ers make use of experience and familiarity to separa te different sound smu ce.s.
When you know what you're listei\ing for, it's easier to pick out sounds from
a background of other sounds. An obvious example is h ow 1.1ukkly you rec-
ognize someone sayin g your name even though there are m.r1ny other sotmds,
including other voices, in the room.
You might be s urprised to learn how quickly you can come to recognize
a complete ly new sound once you 've heard i t a few times. Sounds in the en-
dro1unent, s uch as b ird calls, often occur more than once. To test how much
expe rience lis teners need in order to benefit from familiarity, f\kDermott,
Wroblewski, and Oxenharn (2011) created complex novel smmds by f(lmQirung mma Appa
natural sotmd characteristics in wav.s that listeners had never heard before.
They repeatedly played these sou;1ds at the same time and intensity as a
backgrmmd of other n ovel so unds that did n ot repeat, as sh own in Figure
10.23. A lth ough listeners could not segregate a sound from its backgrmmd
w hen they Listened to a sing le instance, they could nonetheless segregate and
identify the sound when it repeated _Listene rs need ed only a fevv repetitions
to perform well above ch an ce, even though they had never heard the comp lex
sounds befme they came to the laboratory.

314 CHAPTER 10
(n)
Tug.et
4000
¥ 1900
.t
•"'g. 7SO
230
100 200 30-0 100 :!)() 300 100 200 300
lunillill
TII1'le(ms) TUYle(pl.S) Tune (ms)
(b)
Ls
-
8.
l
2
0. 0
1 2 3 5 10
rn =- =========
Ntunber of mixturES
FIG URE 10.23 Ustengs' task Is to identify a novel complex sound (a) when t arget
and d istractor ovgrlap in one oombin€1d sound (mixture). Wh9n a n ew oornp lex
sound {red) is repeatedly played at the sa1Tl9 time as differeint sounds (no t red) jus t a
few times, listeners quickly becom e familiar enough 'vVith the repeated sound that they
it out from other SOllnds in the background.
Continu ity and Restoration Effects

As a lrea dy d iscussed, the sound we' re trying to lis te n to a t an y given tim e
is us ually no t the onl y sound in the env ironm ent. In addition to dea ling
\.vith overlapping a u d itory streamsr we a lso oh en have to deal wi th the to ta l
masking o f on e sound source by another fo r brief per iods . Sup pose you' re
lis tening on your cell phone as a friend gives you directions to the restaurant
w here you' re to m eet for ltmch. A car m ay honk, a baby may cry, or your cell
phone m ay pro duce a short burst of s tatic, but if yo u're paying attention and
the interruption is not too long, you will p robably be able to 11h ear through "
the interruption. This effect is consis tent with the Gestalt principle of good
con tinuation (see Figu re 4.14 in Chapter 4): the co ntinuous a uditory stream
is heard to continue be hind the m asking smmd. Auditory researchers have
b beled these phen om ena "continuity e ffects" or "perceptual res toration ef-
fects" -the la tter label aris ing bee.a use the a uditory sys te m appea rs to restore
the portion of the continuous stream th at\·vas blocked out by the inte rrupting
sotmd (R. M. Wa rren, 1984). In this sense, aud itory res torati on is analogous to
the visual sys tem' s filling in the portions of a background object that is s itting
behind a n occluding object.
Continuity effects h ave been d em ons tra ted in the laboratory with a w ide
variety of target smmds and i.ntem1pting sotmds. The simplest version of su d i.
an experiment is to d elete portions of a pure tone and replace them n oise
(Figure 10.24a ). The tone ,,,,.;U so und continuo us if the n oise is intense enough
to have masked the ton e, had it been p resent (R. Warren, 1984).
H ow d o we know th a t lis ten ers really hear a so und as continu ous in
experimen ts s uch as this? O ne of the best ways to d ete rmin e w hat p eople

(a) Noise (b) FIGURE 10.24 When a sound is

,---'---..
d.:JletGd and replaced w ith [I lolld noise.
list.:.iers w ill hear tha sound as If it con-
tinu9S through the nois9. This is true
whEf'I a tone maintains the same fre-
qllency (a) and whElfl the tone steadily
f increasee or decreases In frequency (b).
0.
Time Tune
reall y perceive is to use a signal de tec tion task. Klue nder and Jen ison (1 992)
used signal detec tion m e tho d ology with a slightly m o re complex: version of
the continuity e ffect (Flgure 10.24b). ln their e xperiments, listeners hemd to ne
glides, in wh.id1 a sine ""·ave ton e varies continuo us ly in frequency over time
(the resulting smmd is si mi lar to th.:"l t of a slide w histle). \Vhen intense noise
is superim posed over pa rt of the glide, listeners rep o rt h earing the glide con-
tinu e behind the n oise. (5.e Web Acthltty 10.5: Continuity and Restoration
Effects.) Kluende r a nd Jen ison crea ted stirnuli in w hich th e middle gnrtion
of the glide ei ther was present wi th the noise or was completely remdved... In AMm.:iAppa
trials in w hich the noise was shortest and most intense, the signal de tection
measure of discriminability (d') d ropped to 0. For these tri als, perceptual
restoration was compl ete: listen ers had no idea w het he r or not the glide was
actually present ' "'1th the noise.
The compelling nature of perceptu al res tor,1tion s u ggests that at some
point the restored m issing sounds are encoded in the brain as if they were
actually present in the sign a l. lmagi..ng studies of humans , w h o ca n report
when they d o a nd d o not hear the to ne thro ug h the n oise, s how metabolic
activity in Al tha t is consis te nt with wh::\t lis teners report hearin g, w hether or
no t the tone is present {Riecke e t aL, 2007, 2W). Macaque mo nkeys also h ear
tones being restored even '"1 hen interrupted by noise (Petkm.-, O'Conn or, and
Sutter, 2003), and ne urons in A 1 of m onkeys s how thesa n1e responses to real
and restored tones O 'Connor, a nd Sutter, 2CXYJ). These data from the
auditory cortex canno t telJ us w he ther the glides were res tored in the cortex
or at a point earlier in auditory p rocessing. H owever, they make it easier to
understand why perceptually restored sounds really sound present.
Restoration of Complex Sounds

Cornplex sounds such as mu.sic and sp·eech ca n also be perceptually restored.
When De\Vitt and Samuel (1990) pla yed familiar melodies with n otes excised
and replaced by n oise, listene rs p erceived the missing notes as if they were
present. Res tora tion was so complete that listeners could n o t report which
notes had been rem oved and replaced with noise. The researchers also tes ted
whether familiarity of the melodies matte red. Just as you might expect, liste n-
ers were much less likely to uhear" a miss ing note in an unfamiliar melody.
Just in case you think only human listen e rs would care about m elodies,
listen up. Seeba and Khunp (2009) trained European starlings (Figure 10.25) to FIGURE 10.25 European starlings
peck w he n they heard a difference between h\.·o parts o f starling song, called perceptually restora bits o f starl!ng
songs, and they arg morg to
motifs. \\/hen starli n gs heard motifs i,..;ith short s nippets filled with noise or rootore song parts when they ate
w ith silenre, they were more likely to peck as if there was a diffe ren ce between familiar w ith the starling that produced
on intad and an internlpted motif w hen silence filled the gap. 1l1is observa - theim.

316 CHAPTER 10
tion suggests that the s tarlings res tored the mis.sing bits of motifs w hen n oise
\ ..·as inserted into the gap. Not a ll s tarling songs a re h owever. In the
sam e set of experiments, the researchers used bits of son g tha t '"''ere either
familia r to the s t.arling in the experiment (the bird 's own song or the song of
a cage ma te) or tmfarnilia.r (from starlin gs the s ubject had never heard ). Just
like humans listening to familia r and unfami liar m elod ies, starlings are more
likely to resto re m issin g bits of a familiar song .
Percep tua l restorati on in sp eech is even m ore compelling than restoration
of mus ic. R. M. \Va n -en and O busek (1971) played the sen ten ce " the s ta te
gove rn ors me t wi th their resp ective legi '1ht\1res con vening in the capital
city" wi th the first s in legislatures rem oved and replaced by s ilence, a cough,
a burst of n oise, or any of a few o ther sounds. Despite the missing s, listen-
ers h eard the senten ce as if it were intact and complete. Even w hen lis teners
were explicitly warned tha t a sma ll part of the sentence had bee n removed
Mtd replaced w ith silence or an othe r sound, they were un able to accurately
report w here the sentence had been changed, excep t w hen the m issing s was
replaced with silence.
Lis tening to familiar m elodies and to rea l sp eech sentences, as opp osed
to siinple soun ds s uch as s ine waves a nd tonal glides, p e nnits li steners to
use m ore than just auditory processi ng to fill in miss ing infomu'ltion . Clearly,
these "h ig her-order" sources of infom1ation are used for listening to sou rces
in th e face o f other acous tic clutter. [n a particularly compelling exa mple,
li steners res tored a missing so und on the basis of ling uis tic informa ti on
th <lt fo llowe d the de le ti on (R M. Warren and Sh e rman, 1974 ). For example,
Warren a nd Sher ma n played uttera nces s u ch as " the •eel fell off the ca r " and
"the •eel fe lJ off the table'' (the aste risk h ere represents a patch of n oise).
Listeners were mu ch m ore likely to report hearing 11 w heel" in the for m er
uttera nce and " mea l" in the latter, even though the context information
cam e well .ifter the mi ssi ng phoneme in the sentences. (See Web Activity
10.5: Continuity and Restoration Effects.) As this result hin ts, m eanin gful
sen tences actually become m ore intelligible w hen g..1psare filled with intense
8230336 Amma A1
n oise tha n w hen ga ps are left s ile nt (Figure 10.26). Think ab out this: adding
n oise improves comprehension l
(•) i
t
1
FIG URE 10.26 Adding noise

improves comprehensioo. (!I) Sp eic-
trogram o f the SQrltQnCli'I •the mailrn..>tn
brought a lette<. • fP} Listeners h ave
gri90.t diffic ulty und9rstanding the s9n-
tenoo when intervals are n:imovOO and
replaced \tllith silence. (c) WhEin the
same intQtvals are rep aced by noiss.
however. it becomes easier to under-
s tand the seintQllCQ, b9cauSl9 of per-
c ep tual restoration. (Afigr Basl1ford and
Wam;m, 1987 .) T1..me

Aud itory Attention

H ave you eve r been so engrossed by a task, s uch as reading this book, that acoustic starUe reflex The very
you d idn't even hea r som eone cal.ling rour name? :At o ther times, maybe also rapid motor resp::;nse to a sudd€f1
w hile re<lding this book, you ever n oticed tha t it took onl y the bares t sound. Very ff!!W neurons are Involved
In the basic startle reflex, which can
distraction to d raw your a ttention away? We said a lot about 'visual a tte ntion also be attocted by ernotlonal state.
in C hapter 7; we will not delve so deeply into auditory attention, but thel'e are
som e parallels to the visua l case. Auditory attention a lso diffel' fro n\ visual
attention in ways that reflect the differences beh·veen senses. You just learned
about how the au ditory system is exquis itely fast and sensitive. Ea rl ier, '"''e
pointed out that hea ring works g reat at a distance, in the dark, and around
obstacles. These facts make hea ring our primary sense for being vigilan t in
om surroundings-your first line of defen se in a sensory world .
We see the auditory system play ing the role of sentinel in th e acoustic
startle reflex. Just like its name implies, this is the very rap id bod,ily
ment that arises fo llowing a loud abn.ipt sow1d. This reflex is very fast: nui.Scle ma Appa
t\ldtches may fo ll ow the sotmd by as little as 10 m s (r\.1usiek, 2003). Beca use
the tim e between ear (so und ) and s pinal cord (movem ent) is so brief, there
C'1Tl be n o more than a few brain.stem ne uro ns behveen them . Being afraid
increases acous tic s ta rtle (Davis, 2006). Directors of horror movies seem to
know this as they gr.:ldually ramp up your anxiety before the big event, and
then the s urpri se is us ually loud .
1l1e acoustic s tartle reflex is unselective--aln'lost any loud sound \-V ill do.
In o ther cases, auditory attention is selective-i t is p icking one sound source
out of severa l. Many sounds occur at the same time in natural environments,
w ith all o f the sounds becoming merged at the ears. This makes listening to
only one sound amon g many a serious challen ge. This problem has much in
common wi th these lectivevisuul attention that you lea m edabout in Ch.apter 7.
Effects o f a ttending to a particular sow-id source can be so strong that we
completely miss o ut on he<uing other sounds in a kind of inattentiona l deaf-
ness. Skilled music ians were no be tter than tmtrained lis teners at n oticing an
electronkguit<U' improvisation that mixed in with Richard Strauss's Titus
Spoke Zarnthustra (theme musk fro m the sci-fi classic 2001: A Space Odyssey)
w hen both groups were asked to count (and, thus, to attend to) the munber
of timpani beats (Korei mann, Gula, an d Vi touch, 2014 ). This inattentional
dea fness has its limits, because lis teners had less trouble n oticing the guitar
w hen it was made sufficiently lo ud .
While inattentional deafness m ight seem to be a bad thin g, it really repre-
sents a n extreme example of auditory p rocesses that help us to listen in our
acoustically crowded world. Imagine yourself in a room full of people who
are speed dating in their sea rches for true love (Figure 10.27). You earnestly
focus on the person who you are sizing up, but man y other voices compete
wi th the one voice yo u are trying to w1derstand. Listene rs ca n use the aco us tic
characteristics of a talker to track what tha t voice is saying despite the clutter
of othe r voices. While we might think that thi s relies on being familiar with a
talker or actively concentrating on that voice, we would be wrong (Bress ler et
al ., 2014). In.stead , it appears that the bra.in does this a utomatically, foll owing
principles like those for auditory strearn segrega tion d escribed earlie r in the
"Spa tial, Spectrat and Temporal Segregation" .section.
Suppose tha t, after s haring a bJ"ief exch ange on favori te colors, you req u ire
no more time wi th the person to w hom yo u've been lis tening. You begin to
tune in to a neighboring conversation \'V"itho utmoving away jtLs t yet from your
currently assigned d atin g c,mdidate. TI'is requires shifting attention. There are
several p oi nts to be about this situation. Firs t, there is a cost to m oving
to the next .sotmd sou rce: listeners beco me less accurn te in und erstanding

318 CHAPTER 10
FIG URE 10.27 The organizers of this

speed-dating event oouldn 't seat three
1X>tM tial couples at one if WQ
didn't have the ability to attend to one
voice am ong many. Now that although
each of these individuaJs to
be att9nding to th9 pernon across
the tabh;i , any of them could actL1-
alty be attending to one o f the oth91·
convQl"Satio ns.
wha t they hear w hen they h ave to swi tch be l:\veen talkers (Lawo and Koch,
2014). Second, yo ur ability to look at one person l·v hilea ttend ing toa different
con versa tion illustra tes the flexibility of yom attentional apparatus. Finally,
if you a re a ttendin g to a nothe r con versatio n 1 reall y em barrassing things can
happen wh en you realize that your partner ha.s stopped speaking a.nd is wai tin g
fo r a response to a question to which you have not a t tended . You rn n svdtd1
ntten tion back a nd fo rth behveen stream s, but you cannot fully process two
streams of speech any rnore tha n you can read t\vo sentences a t the sam e time
(see Figure 7.1 in Chapter 7).
Summary
1. Listeners use sma ll differences, in time and intens ity, across the h-.•o ears to
Refer to lhe lean1 the direction in the horizontal plane (azimuth) fro m \\.'hich a sound
Sensation and Perception comes.
Comparion Website 2. Time and intensity differences across the two ears are not sufficient to fully
sltes.slnauer.com/woKe4e indica te the location from w hich a soWld comes. In p ar ticula r, they are no t
fCT esS1¥'. study enough to indica te whether sounds come from the fro nt or the back, or fr om
h igher or lm<Y·er (elevation).
qlEStions, and otrer study aids.
3. TI1e pinna, ear c·anal, head, and torso alter the intensities o f di fferent fre-
quencies for sounds coming from different places in space, and listeners
1.tse these changes in intens ity across frequency to ident ify the location from
w hich a soWld comes.
4. Perception of auditory d istance is s imilar to perception of visual d epth
because no single ch a racteristic of the signal can inform a listener about
how d istant a sound source is. Listen ers m ust combine inten sity, spectral
compositi on, and rela tive a mounts of d irec t and reflected energy o f s01.mds
82 to estlnlate d istance,_ to a sound source.
5. Many natural sounds, including music and human s peech, have rich
harmon ic structure with energy a t integer multiples of the fundam ental
frequency, and listeners are especially good a t perceiving the p itch of har-
monic sounds.

6. Important perceptual qualities of complex solU1d.s are timbre (conveyed by

the relative amo unts of energy at d ifferen t frequencies) and the o nset and
offse t proper ties of attack and decay, respectively.
7. Because all the sounds in the e nvi ronment are su1nmed into a single wave-
for m tha t reaches each ear1 a major challenge for hearing is to separate
sound sources in the combined signal. This general p rocess is known as
auditoty scene analysis. Sound source segrega tion s ucceeds by u.sing mul-
tip le characteristics of sounds, including spatial location, similarity in fre-
q ue ncy and timbre1 onse t p roperties, and familiarity.
8. In everyd ay environments, sounds to wh ich a person is listening often are
interrup ted by o ther, lo uder sounds. Pe rcept ual restoration is a process by
which m issing o r degraded acoustic signals are perceptually replaced.
9. Auditory attention has many aspects in conuno n w ith visual a ttention . It
is a 'b a beh veen able..tp make use of sounds o ne needs to h ear
in the midst of competing sound.S, and being on alert for new a uditory
infomlation.

CHAPTER 1 3J 336AMmaAppa

Music and Speech
Perception
SOUNDS FROM MUSICAL INSmUMENTS AND HUMAN VOCAL TRACTS obev the
same laws of physical acoustics as all other sounds. G uitar s trings and ln;man
vocal fo ld s are vibratory structures, with similarities to mbbe r bands and s us -
pension bridges. Trombones and voca l tracts act as reson.ntors following the
sam e laws as e mpty bottles and h ollow logs. In th is sense, spoken words -and
musical notes are n othing m ore than e:,y
In o ther "'.rays, h owever, music a n s peech c<i n be d isting'li ish ed from
m ost o ther environmental sounds. Much like visua l a rt s uch as paintings and
sculph.tre, music and speech are crea ted with perceivers in m ind . Both music
and speech serve to conununicate, and both can con vey em o tion and deeper
m eanin.gs. In song, music and speech conspire to move the listener. A Ithough
d ogs, birds, and whales share acoustic messages and even sing to one another,
there is n o questi on that the depth and breadth o f human communica tion by
music and language has n o riv al in the acous ti c world . (n this chapter we ex-
plore these communicati ve aspects o f heari ng: music and speech
Music
People have been using music as a way to express the mselves and influen ce
the thoughts and emotions of others for a very long time. 1he oldes t d iscov-
ered mus ical instruments-flutes carved from v ulture bones-are at least
30,000 years old (Conard, Malina, and Mi.inzel, 2009). You probably know o f
the grea t ancient Greek sch olar Pythagorn s (c. 580-<:. 500 BCE) from the Py-
thagorean theorem in high sch ool geom etry. To say that Py tha.gom s was ob-
sessed with nwnbe.rs would l>e an und ersh1tement. And the numbers that he
'1nd his fo ll owe rs cared about most were those found in mus ical scales. They
were con vinced tha t the rnu siciJ intervals they found m ost pleasing s hould
provide the g re atest ins igh ts no t only to mathema tics, but to the universe as
a w h ole. Althoug h music may n ot actually explain the known universer we
all appreciate how itnportant rnusic is to culttue and , perhaps, to personal
cultural identity.
Musical Notes
From Chapter 9 you know that on e of the most important characteristics: o f
any acoustic s ignal is frequen cy. You also know that brain s tructu res for pro-
cessin g sounds a re tono topically organized to correspond to frequen cy. The
psychological quality of perceived frequen cy is pitch. When you im agine d
pitch w hil e ren ding C hapter 9, you p robably imagined mus ical p itch. Figure pitch The r;eychologlc aJ aspect of
11.1 illus trates the ex tent of the freque1\cy range of musical sounds in relation sound related malnt>J to perceived
to human hearing. frequency.

322 CHAPTER 11
r - - - - - - - - Accordion - - - - - - - - - - - - <
l - - - - - Piocolo- - - - - 1
I - - - Soprano .saxophone - - - I
Bass s axo phone -
=-- ======
"
nma Appa
A E'> C D E" F G A E'> C O E: f C A E'> C O E" P C A B c DE: r e A 8 CD[ F COE F G A B (OE F AB C
FN!qt1etlC)' (Hzj
FIGURE 11 .1 The sounds of music
ext€f"ld across a freq uQncy range from
abc:ut 26 to 4200 hertz {Hz).
TONE HEIGHT AND TONE CHROMA Musical pitch is on e of the characte r-
istics of musica l n o tes, the soun ds that con sti h.1te m elodies. A very important
concep t in lmderstanding musical pitch is the octave. \Vhen we described
pitch (or psycho..1cou st ic pitch ) in 0 1apte r 9, it seemed dear that the nearer
any hvo sounds \'.:ere in frequency, the nearer they were in pitch. Here's where
octaves come in. \ Vhen one of hvo periodic SOlU\ds is dou ble the frequency of
the o ther, those n.vo sounds are one octave apart. For example, midd le C (Cd)
h as a hmdamen tal frequen cy of 261.6 he rtz (Hz). Notes tha t are one octave
below a nd above midd le Care 130.8 Hz (C3) and 523.2 Hz (C 5), respectively.
Not only d o these three so unds have the .same names on the mus ical scale
(C), but they also so und similar. In fact, C 3 (130.8 H z) sounds more similar to
Cd (261.6 Hz} than to a sotmd with a cl oser fre<iuency-for example, (164.8
H z)t C learly there is more to musical pitch than just frequency.
TI1e pret."'eding example illus trates the con cep t of "jus t intonation/' in which
&eq uencies of sounds are in simple ratios w ith one another (e.g ., 2:1 fo r an
oc tave). But in typical Western music, the frequencies of n otes are adjusted
octave The Interval between two slightly from simple rati os so that combinations of notes w ill so und equa lly
oound frequencies having a ratio of
2:1.
good when played in higher- or lower-freq uen cy ranges (keys). The set of
notes (scale) used conunonly in \¥es ten1 music is called "equal temperament.'1
tone height A round qualitycor-
respmdlng to the level of pitch. Tone
Because of octave relations, m usical pitch is typica11y described as havin g
height Is monotonlcaly related to two dimensions. The firs t is tone height1 wh ich relates to frequency in a fairly
frequency. s traig htforwa rd way. The second dimension, related to the octave, is tone
tone chroma A sound quality chrorna (drromn is the Greek word for "color''). \Ve can vis ua lize mu sical pitd1
s hared by tones that have the same as a helix. Frequency and tone height increase wi th increi.'\Sing heigh t on the
octave lntervaL helix, as s hown in Figure 11 .2. The circula r laps ::.u'Ound the helix correspond

MUSIC AND SPEECH PERCEPTION 323
FIGURE 11 .2 This helix Illustrates the tvvo characterlstbs of musical pitch: tone
height and tone chroma. Tone height
to changes in tone chroma. At the same point along each lap around the helix,
a spe::::ific sound lies on a vertical line, and all sounds along that line share
the same tone dl.rom.."I and are separated by octaves. (See Web Activity 11.1 :
Notes, Chords, and Octaves.)
In your early years of schooling, you probably learned to sing the notes of
the musical scale: do, re, mi, fa, sol, la, ti, do. Perhaps you even tested how many
times you could sing the scale at increasingly higher pitches. In that case you
were actuaJ ly singing your way up the musical helix. Your pitch traveled a hill
tum upward with each repetition of a particular note-for example, do or re.
In Chapter 9 you learned that both a place code and a temporal code can
be used in the perception of pitch. Neurons in the auditory nerve convey
frequency information both by their location in the cochlea (place) and by
the timing of their firing (temporal). For frequencies grearer than 5000 Hz, ""'"'
temporal coding does n ot contribute to the perception of pitdt, and pitch notes
discrimination becomes appreciably worse because only place coding can
be used . Most musical instruments generally produce notes that are below
4000 Hz. Could tone chroma somehow be related to the temporal encoding
of pitch (Moore, 2003)? We do not know. We do know that a sequence of pure
tones with frequencies greater than 50)() Hz does not convey a melody very
well (Attneave and O lson, 1971), and listeners have great difficulty perceiving
octave relationships betw'een tones when one or both tones have a frequency
greater than SOOOHz (Ward, 1954).
CHORDS Music is further defined by rid1er,complex sounds called chords, chord A comb'1atlon or three or
whid1 are created when three or more notes are played simultaneously. (The more musical notes ViJ!th different
1
simultaneous playing of two n otes is called a "dyad. ' ) The major distinction pitches played simultaneously.
between chords is whether they are consonant or dissonant. Perceived to be
most pleasing, consonant chords are combinations of notes in which the ratios
between the note frequencies are simple. llUs is why Pythagoras was so taken
by the relationship between pleasantness and mathematics. You already know
one of the consonant relation.ships, the octave, in which the frequencies of the Tone height
tw'o notes are in the simple ratio of 2:1. Other major consonant intervals are the
perfect fifth (3:2) and tlle perfect fourth (4:3). Disso11a11t intervals are defined by
less elegant ratios. For example, the minor second (16:15) and the augmented
fourtl\ (45:32) do not sound very pleasing. Indeed, during the Middle Ages
the augmented fourth was called the "devil in music" (Seay, 1975).
Because chords are defined by the ratios of the note frequendes combined
to produce them, they are named the same no matter what octa ve they're
played in. For example, the G -major chord consists of G, B, and 0 , and it can
be played as G 2 + B., + D,, G4 + B4 + D., G 6 + B6 + D7, or in any other octave,
provided the ratios remain the same. Note th.at in the musical he lix in Flgure
11 .3, the relationships between the notes of the chord on the helix stay the
same, as in Figure 112; the only thing that changes is the pattern's height.
CULTURAL DIFFERENCES Musical scales and intervals vary widely across Musical
cultures. Although potent relationships between notes such as octaves are EO.:ale
notes
relatively universal, different musical traditions use different numbers of
J=IGURE 11 .3 Chords are made up of three or more not9S and can be played with
different tone hGights whlle their chromatic relationships are maJntalned. He-e, th9
G-major chord (purple shading} Is shown being playact at diffQ1'"9nt heights.

324 CHAPTER 11
n otes a nd sp aces between notes vvithin an octave. All o f o ur disc ussion has
concerned the hep ta tonic (seven-note) scale . An other conu n on scale, penta-
ton ic, haS fi ve notes p er octave, an d you 've heard th is sca le in gosp el, jazz,
rock, a nd blues inusic, a m ong other North A me rican genres. The pentatonic
scale is trad itiona l in 1nany othe r p a rts of the world , especia lly Asia .
You m ay be th a t Asian la n g uages su ch as tv1a nd a rin (a C h in ese
lan g uage), Thai, and Vie tnam ese a re ton e la n guages, givin g a sin gsongy
impressio n to som e Eng lish listeners. Speakers of to ne la ng uages use d "\anges
in voice pitch (fund amental frequency) to d istin g u ish different wo rds in th ei r
lan guage. For exan1ple1 the Mandarin word ma means " horse" or "mothe r''
d ependin g on how voice p itch l'ises or fa lls.
lt appears tha t language m ay a ffect the types of m usical scales people
use. C hanges in p itch d irection are larger and m ore freque nt in spoken tone
langu ages su ch as Ma ndarin, Thai, and Vieh1am ese tha n in English, Frenc h,
and Germa n. Parallel acous tic d iffe re nces Ltre fou nd between the p enta tonic
8230336 Anma Appa music of Ch ina, Thailand, a nd Vieh-iam an d the heptc1ton.lc music of Westem
cultures. Pitch ch an ges in A.sian m usic are larger an d occ Lu m ore o ften (Han
et al., 2011 ).
ln scales fo r ""· h.id1 fewer notes comp rise an octm.-e, n otes n-iay be more
loosely hmed tha n a re notes in the hepta tonic Western sca le. \Nhen the re are
fewer notes to distinguish, a wide r ran ge of p itches could qu <11.ify fo r a given
note. For example, th e s!Cudro and pJlog scales h ave fewer tha n seven
n otes \'1'1. thin an ocrave, a nd there is variation in a note's a1..--ceptable
frequencies.
Be::ause people a round the world have very d ifferent listenln gexperiences,
you might expect the m to hear musical notes in different ways. lndeed, w hen
Javanese a nd\ Veste m m usician s hear in tervals be tween n otes, their estimates
of the intervals vary according to how '\'\--ell those n otes correspond to Javanese
versu s \ Ves tem scales, respectively (Perlman and Krum.h an sl, 1996). Further-
more, infa nts seem equ.l pped to learn w ha. te\·er scale is u sed in their en 'viron-
ment. Lynd 1 and Eilers (1990) tested the d egree to w hich 6-m onth-old infan ts
noticed. in approp riate notes 'vi.thin both the trad itional Westem scale and the
Jova:nese rf'log scale . TI1e in fan ts to be equally good a t detecting such
"mistakes" \.vi thin both Wes tern and Javanese scales, but adults from Floridn
were reliabl)'· be tter at de tecting deviations from the Western scale.
ABSOLUTE PITCH You may 1-iave heard that some people are "gifted " wi th
absolute pitc h (AP) A rare ability absolute pitch (AP), which is o ften referre d to as ;,perfect p itch." AP is a rare
w hereby some people are able to V€fY abi lity of some listene rs to accurately n am e or recrea te particular n o tes in
ao::::urately narne or produce notes
isola tion . In this way, "a bsolu te" is u sed to distin gu ish AP fr om the use of
without ocmpariscn to other rotes.
"rela ti ve" pitd 1, the way m ost people id en tify notes rela ti ve to one a nother.
A P relates sp eci fi ca lly to musical n otes, becau se p eople with AP s h a re the
same basic sense of hearing as o ther peop le. Their auditory syste rns are no
more sensitive than n om1al, a nd they are no bet ter than us ual a t d etecting
differences between sounds .
AP is very rare, occu rring in less than on e in 10,0CXJ people (Takeuchi an d
Hulse, 1993). A mong musiciar1s, this rare skil l is often conside red to be highly
d esirable. lv1a ny believe tha t Ludwig van Beethoven had AP, a nd it a ppears
fair ly certa in th at \Volfgang Mozart had A P by the tim e he was only seven
years old (Deutsch, 2013).
Resea rchers disagree as to how some people come to have AP. One idea is
tha t people might acquire AP follow ing a grea t dea l of practice; however, it
is very d ifficult for adults to acquire AP even thro ugh a grea t d ea l of p rac tice
(Br<ldy, 1970). ln contrast to leaming AP, others have suggested that some peopl e

are born with AP (Athos e t al., 2007), and this is consistent \vi th the fact that
AP appears to nm in fa milies (The usch, 2009) . However, children in the sam e
family share more than gene tics. In a ho me in wh id1 one child receives music
lesson s, it is like ly th.at s iblings also receive rnusic lessons . There is variati on
in AP, and this variation presents a challenge for identifying p ossible genetic
components. Most traits tha t are influenced by genetics, sud1 as height, show
va riation-they are not a ll or none-and variation in exp er tise also occurs fo r
skills that are learned . The best explan ation for AP m.ightbe that it is acq uired
through exp erience, but tha t e xpe rience must occ ur at a yo\.mg age. In studies
m easuring abilities a cross man y people, the age at which m ictll App a
begins is \<\--e ll correlate d wi th h1ture p ossession of AP (Deu tsch, Le, Shen, a nd
Li, 2011; Lee and Lee, 2010).
AP might n ot be so absolute, and adults with AP m ight be m o re fl exible
in their lis tening than typically tho ught. Afte r first testing the identification
of notes by l isteners wit h AP, H edger, H ea ld, and Nusbaum (2013) had their
participants listen to Joha nnes Bra hms 's Symphony No. 1 in C minor whi.le
"sking the m to pay d ose attention to indi vidu"l rne lod.ies. \ Vhat the resea rch-
ers d id no t tell the ir listeners was that_, during the fi rst part of the piece, they
very grad uall y ud ettmed" the music by making all the n otes a bit lower (flat)
before playing the m usic at these lower frequ encies for a while longer. Afte r
the m usical passage was complete,. AP lis teners were m ore likely to jud ge fln t
n otes as bein g in-hme a nd to judge in-hme no tes as being mistune d . While
it may be very diffi cult to train adul ts to become AP listener s, perception of
musical n otes by adults \vith AP can be s h ifted by musical exp erien ce.
Music and Emotion

AA ear can break a human heart
I<!. qulcKty as a spear
We wish the ear had not a heart
So dargerously near
- Dlcklnscn, 'The Saddest Noise, the SWeetest Noise"
Listening to music affects people 's moods (Eich, 1995; Pignatiello.
Camp. ard Rasar. 1ra3) and emotions (Slobcda, 19ffi). When listen-
ers hear pleasant-sounding chords precOOing a word, they are faster
to respond that a word such as charm is positive. and they are slower
to resp:1nd that a word such as ev# is negative (Sollberge-, Reber, and
Eckstein. 2003). Given the power1ul effects of music on mocd and emo-
tion. some clinical psychologists practioo music therapy, through which
people sing. listen. play, and m ove to music in efforts to improve mental
and physical health.
Music has deep physiological effects. Music can prom ote i:::ositive
emotions, reduce pain, and alleviate stress. and it may even improve re-
sistance to disease (Gangrade , 2011; Roy, Peretz, and Rainville. 2007).
Evi de nee acrcss many studies suggests that music can have a positive
impact en f:0,in , anxiety, mooct. and overall quality of Hfe for cancer pa-
tients (Archie, Bruera, and Cohen, 2013). Vvllen peope listen to highly
pleasuraJ::je music. they experience manges in heart rate. muscle elec-
trical acttvity, and respiration , as well as blood now increases in brajn
regions that are thought to be involved in reward and motivation (Blood
and Zatorre , 2001). Music Is a powerlul human inventirn indeed.

326 CHAPTER 11
cJ
' I ( F
Q
r
'
ii!
•·
j
I ==
FIGURE 11.4 The path;1m of increasing and d9Crea.sing pitc h9G can r01Ttaln the
same, but the mebdy will changg If notes have different duratbns. Here. serise.
of no tes with the same meody contour are shown with notes that differ only in
duration.
Making Music
melody A sequence of notes cr Notes o r ch o rds can form a melody, a seq uence of sounds perceive d as a
c hcrds perceived as a single coherent single coh erent structure. The n otes of a familiar melody, su ch as 11 Twinkle,
structure. Twinkle, Little Star" (a.k.a . "No'"1I Know My ABCs/' ''Baa Baa Black Sheep,''
terrpo The speed of the and Variation K.265 [300e] by Wolfgang Amadeus Mozart), "belong togecl1er"
presentation of sounds. percephlally because they fonn this melody. Note that a me lody is defined
by its contour--the patte rn o f rises and declines in pitch-rather than by an
exact sequence of sound freq uencies (Handel, 1989).
You've a lready learned o ne simple w.Jy in which me lody is not a sequence
of specific sounds: shift every note of a melody by one octave and the resulting
melody is the same. \'\Then you s ing \.•.:ith other people possessing higher or
IO\•Ver voi<:es, they sing the same melody at very different pitches. Even with.in
a single octave, the same melody can be heard from different notes if the steps
bet>veen n otes s tay the same.
In addition to varying in pitch, notes and chords vary in duration. The
ave.rage duration of a set of notes in a melody defines the mu:Mc' s tempo. Any
melod y can be played a t e ither a fast or a slow tempo. But the relative dura-
tions within a 5eGuence o f notes are a critical part of the melodies themselves.
If the n otes of a given sequence are played with different relative durations,
we will hear completely different melo dies (Figure 11 .4 ).
RHYTHM In adaition to varyln!i' n speed[ nnl•ic varies in rhytlun. The fact

that music has rhythm should go without saying. After .ill, how else would
we dance to it? Less obvious, perhaps, is th'-lt many-or even most-acti\.ities
have rhythm. I Valkingand galloping hove rhyclun. So do finger tapping, wav-
ing, and swim. ming. Perhaps it is the very commotmess of rhythm th.:lt ca uses
us to hear nearly ..,11 sounds as rhythmic, even w hen they're not!
Over a century ago, Thadde us Bolton (1894) conducted experin1ents in
wh ich he played a sequence of identical so unds perfectly s paced in tim.e; they
had n o rhytlun. Nevertheless, his listeners readily reported that the sotmds
01..-"'C'UJ"I'e<l in groups of h-vo, three, 01 four. Moreover, they reported hearing the

,& a a A a a Aa aAa a !2_a aAa aAa a Aa a &, a a
-BbbbBbbbBbbbBbbbBbbbBbbbBbbb
- -
FIGURE 11 .5 Whein tiNo rhythms are played togeth0r and one rhythm fn this case
A aa) is dominant, list€1no91"S to pet""CQive the timing o f beats in the no ndo minant
rhythm {Bbbb) adjusted to c onfotm w ith the dominant rhythm.
first sotmd of a group as "ai...-cented/' or 1'stressed ," while the remaini ng sow1ds
were tmaccented, or \IDStressed . You've probably ha d a similar experience '"'+tile
riding in a train or car. Even though a train travels over jtmctions in the rails
at nearly equal intervals, you h ear the sound as "CLICK dick C LICK click."
When yo u ride in a Gu· at a steady speed, you hear "11-IUMP thump THlJl'v[p
thump " as yo ur tires roll over cracks in a con crete road.
A s Bolto n' s s tudies sh ow, listeners me predisposed to grouping so und s
into rhy tlunic patterns. Several qualities contribute to w he ther so unds will
be heard as accen ted (stressed} or unaccented (un stressed ). Sounds that are
lon ge.r, louder, a nd highe r in pitch all are more likely to be hea rd as leading
thei.r gro ups (Wood row, 1Q09). The timi ng rebtion.ship between on e sound
and the others in a sequ en ce al.so helps determine accen t. Fo r example, we are
m ore likely to hear a se ries of three sounds as" Aaa Aaa Ana" than as "aAa
aAaaAa."
Listeners prefer1 or <'l t least expect1 sequences of notes to be fai rly regular1
and this tendency p rovides opp ortunities for cornposers to get crea tive wi th
their beats . On e way to be crea tive in d eviating from a bland s uccession o f
regula r beats is to introduce syncopation. Syn copation is any a . at o n a App a
from a regular rhythm, for example, by accenting a no te that is expected to
be unaccented or n ot play ing a n ote (replacing it wi th a rest} w hen a note
is expected. Syncopation has been used for ce nturies and ca n be found in
compos itions by all the grea t composers, including Bach, Bee th oven, and
Mozart. Most collegesh1de nts may be m ore familiar with syncopation from
jazz, reggae, and ska..
O ne particula rly inte resting exa mple of syncopation is syncopa ted audi-
tory poly rhy thrns. When h vo different rhythms are overlapped , th ey can
collide in inte resting ways. For example, if one rhy thn1 is based on 3 beats
(AaaAan.Aan. Aaa) a nd the o ther on 4 (BbbbBbbbBbbb), the firs t accented
sound for both rhy thms \vill coi nci de o nly on ce every 12 beats. Across th e
11 intervening beats, the two rhythrns will be out of sync. \'\i'hen we liste n to
syncopated polyrhythms, one of the ti.vo rhytluns becom es the d ominant or
controlling rhythm, and the other rhythm tends to be pe rcep tu ally adjus ted
to accomm odate the firs t (Figure 11.5). In particular, the accented beat of the
subordinate rhythm shifts in time (Handel and Oshinsky, 1981). Thus, syn-
copation is the perceptio n that beats in the s ubordinate rhythm have achtally
traveled backward or forward in time !
These findings reveal that rhythm is, in large part, psychological. We can
produce seq uences o f so unds that are rhy thrnic a nd are pe rceived as s uch.
But we also hea r rhythm when it d oes not exist1 and no tes e ffectively travel
in time to m aintain the of consistent rhy tlun.
MELODY DEVELOPMENT Like rhy thm, melody is essentially a p syd1o logical

entity (Handel, 1989). TI1ere is nothing about the partk ular sequence of notes syncopation Any deviation from a
in 1'Twinkle, Twinkle, Little Star" that rnake.s them a melody. Rather, it is our rat)ular rhythm.

1ma Appa
328 CHAPTER 11
experience with a particular sequence of no tes or with s imilar sequences that

helps us perceive coherence.
Studies of 8-1nonth-old listene rs reveal that learning of m elodies begi ns
quite early in Hfe . Sa.ffra n et al. (1 999} crea ted six s impl e an d d eli berately
n ovel "me lod ies" com posed of sequences of three ton es. Jnfants sat on their
paren ts' laps w hile hearin g only 3 minutes o f continuous ra ndom repetitions
of the melodies. Nex t, infants hea rd both the original melodies an d series of
new three-tone sequences. These new sequen ces contain ed the same no tes as
the originals, but o ne part o f the sequence was taken from one melody and
another part &o1n anoth er melody. Because the infants responded differe ntly
to the new m elodies, we can deduce that they had learned something about
the original melodies.
This ability to learn new melod.ies is not limited to simple sequences of
tones. In a s tudy with 7-month-old infants, parents played a recording of two
Mozart son a ta m ovements to their infants every day for 2 weeks (Saffrn n,
Lornan, and Robertson, 2000). After an o ther 2 weeks had passed, the infan ts
were tested in a laboratory to see whether they reine mbered the movements.
Infan t listeners responded differently to the original movements than to similar
M ozart m ovemen ts introd uced to them fo r the first time in the laboratory.
Speec h
Most people wh o lis ten to speech als o p roduce speech. Talkers speak so tha t
they c..'m be unde rstood, a nd the relationsh.ip between the production and
perception of speech is an especially intimate one. Therefore, it is import.a nt
to know some things about speech productio n before tryin g to un de rstand
speech perception.
Humans are capable of producing an incredibl e range of d istinct speech
sounds (the 5000 or so languages acr0'5s the world use over 850 different speech
sounds) (Maddieson, 1984). lf you' veever heard Kenny Muh..-m un..1d, the "Hu-
man Orch estra," you' ve experienced the unrivaled versa tility of hunmn sound
production. Th.is flexibility arises frorn the tmique s tructme of the human vocal
tract (Figure 11 .6 ) (Liebennan , 1984). Unlike tha t of other animals, the human
lary'Tlx is posi ti on ed q uite Joi,...,• in the throat. O ne no torious disa dvantage of
su ch a low larynx is that humans choke on food mo re easily than a ny othe r
an imal d oes. The fact that these life-threa tening anatomica l liahiliHes were
evolutionarily tnunped by the survival advantage of ora l communica tion is
a testamen t to the imp ortance of lang uage to human llfe.
Speech Production
The production of sp eech has th ree b<1sic compon ents : respiration (lungs),
p h onatio n (voca l fo ld s, and articulation- voca l tract-see Figure 11 .6).
Speaking fluently requires an imp ressive d egree o f coo rdinati on am ong
these comp on e nts.
RESPIRATION AND PHONATION To initia te a speech sound, a ir mus t be

p ushed out of the lungs, through the h ad1ea, a nd up to the larynx. The dia-
vocal tract The airway above the phragm flexes to d raw air into the lw1&S 1 and elas tic recoil forces air back out_
larynx used for the procluctlon of
s peech. The vocal tract lrcludes the At the larynx, air must pass through the hvo vocal folds, which are made up
cral trac t and nasal tract. of mu scle tissue that ca n be rid jus ted to vary how freel y air passes through the
phonation The process through opening between the m. These adjustments are described as types of phonation.
which vocal folds are made to vibrate TI1e rate a t w hid1 voca l folds v ibra te d epends on their stiffness and m ass.
when alr pushes out of the lungs. Consider gu itar stri ngs as an a nal ogy. Just like guitar strings, vocal fo ld s

(b)
,/
AhreoJ.ar
ridge
Oralb<ut
.,,..,,.,,, ...
-----;------- r..th
Pharynx
Epiglottis
FIGU RE 11.6 The anatcrny unda-lying speech prOOuctim. {.:l) Speech production
has thr9"! basic components: respiratio n j:honatlon {voe.al folds), an::::! articula-
tion (Vocal tract). {b) The voe.al tract Is made up of the oral and nasal tracts.
becom e s tiffer a nd vibrate fuster as their tension increases, creatin g so unds

with 1-Ugher pitch. The pitd1 o f a guita r s tring a lso depends on its thickness or
mass. Thinner g uitar strings vibrate more quickly and crea te higher-pitched
sounds. Similarly, children, w ho have relative.lysm..Ll.l vocal folds, have
pitched voices tha n adul ts do. Adult men generally have lower-pitched \l'o.iais 1Ma Appa
than wom en have, because o ne of the effects of testosterone during puberty is
to increase the mass of the vocal fo ld s. By va rying the tension of voc..1-1 fo lds
(stiffness) and the pressure of airflow fro m the lungs, indi vidual talkers can
vary the hmdamental frequ ency of voiced sounds.
If we were to measure the sound right after the larynx, we would see that
v;bra tio n of the vocal folds creates a harrnonic spectnim, described in Chapter
10 and illustrated in Figure 11 .7a. If wecoLtld lis ten to just tlU.s part of speech,
it would sound like a buzz. TI'e fi rs t harmonic cor responds to the actua l rate of
physical vibra tion of the folds-the fundamental frequency. Talkers can
m ake interesting m odifications i.n the way thei r voca l folds vibrate--creating
breathy o r creaky voices, for example--and singers can vary voca l-fold ten-
sion a nd air press are to sing notes with w idely varying frequ encies. However,
the rea lly extraordi.nruy pa rt of producing speech sound s occurs above the
laryn x and vocal folds.

330 CHAPTER 11
FIGURE 11.7 The spectrum of (a) Haunon.ic spectrum

sound coming from the. vocal folds is
a harmonic spgctrum {a). Aft,er pass-
ing through the vocaJ tract. w hich has
1
r9SOnances based on tllQ vocal tract' s
shape at the time Ip), there. are pgaks
(fonnMts) and troughs o f energy at dif-
farient frequ.encies in tM scunds that
come out of the mouth (c}.
111111111111111111111111111111
Frequency
(b) Fiher functi on
Frequency
(c) Vowel output
1
l11IIIIll11111IfII11 •• 111I1 II.
Fre..1uency
ARTICULATION TI1e area above the larynx-the oral tract and nasal tract
combined-is referred to as the 11vocal tract" (see Figure 11.6b). Humans have
an unrivaled ability to change the..6ib..1pe of the. voca l tract b)! manipu lating the
jaw, lips, tongue body, tongue tip, velum (.soft palate), and o ther vocal-lTact
s tructures. These manipulations are referred to as artlculatlon. As you will
recall from Chapter 9, changing the size and shape o f the space through which
sound passes increases and decreases energy at different frequencies. We cal l
these effects "resonance characteristics," and the speclTa of s peech sounds are
shaped by the way people configure their tracts as resonators. Figure 11 .7b
illustrates the filtering effects of the vocal tract for the V0\1\-'el sound 'eh,' as in
wet. Figure 11 .7c por h·ays the n et result of passing the periodic energy from
articulation The act or manner of the larynx through the voca l tract.
producing a epooch sound using the
Peaks in the speed, spectrum are referred to as formants1 and fomrnn ts
vocal tract.
are labe led by munber, from lowest frequ ency to highes t (F 1, F2, F:y and so
formant A resonarioo of tl1e vo:::.al on). TI1ese concentrations in energy occm a t diffe rent freque ncies, depend-
tract. Formants are epeclfied by ther
center frecL.OOCI' and are denoted ing on the Length of the voca l tract. For shorter vocal tracts (in child ren and
by Integers that Increase with relative s malle r ad ults), forma nts are at higher frequencies th.:m for longer vocal tracts.
frecuency. Because absolute frequencies chan ge d epending on wh o's talking, listeners

(•) (b)
-!I
2
l
Frequency Amplitud• (dB)
FIGURE 11 .8 For sounds that do not vary OV8r time, frequency spectra can b9
repr9S9nted by graphs that plot amplitude on the y-axis and fr9QU&ncy on the x-axis
(a). To stow sounds whose spectra change over time, we rotate the
graph so that frequQf"CY Is plotted on thQ y-axis (b): then we pklt time on th9 x-axis
(c}, with the amplitude of each frequency during each time slice represented by color
(rsdd9r shO"Nlng greater intensity). The spectrogram in part shO\NS the acoustic
signal produced by a male uttering the sentenoe "we were a.Net; a ygar ago...
must use the relationships between formant peaks to perceive speech sounds
(Kluender, Stilp,and Kiefte, 2013). Only the fost three formants are depicted
in Figure 11.7c. For the most part, we can distinguish almost aU speech sounds
on the basis of energy in the region of these loVY-est three formants. However,
additional formants do exist, at higher frequencies with lower amplitudes.
In Chapters 9 and 10, many of the sounds that we discussed had constant
frequency spectra. That is, ii a sound started with a SO.decibel (dB), JOO.Hz
component and a 60-dB, 2!X}-Hz component, these frequencies continued at
these amplitudes for the duration of the sound. One of the most distinctive
characteristics of speech sounds is that their spectra change over time. To rep-
resent this third dimension (time) in addition to the dimensions of frequency
and amplitude represented in frequency spectra of static sounds, auditory
researchers use a type of display called a spectrogram. In a sound spectro-
gram, frequency is represented on the y-axis, time is tracked on the x-axis, and
amplitude is indicated by the color of any point on the graph (Figure 11 .8).
Formants show up clearly in spectrograms as bands of acoustic energy that
undulate up and down, depending on the speech sounds being produced.
CLASSIFYING SPEECH SOUNDS Speech sounds are most often described

in terms of articulation. This is because in the early days of studying speech,
people did n ot have electronic recording or the ability to analyze sounds. In-
stead, they paid close attention to their own vocal tracts and described speech
sounds in terms of the articulations necessary to produce them. You will get spectro!J"am In oourd analysis, a
three-dimensional that plots
the most out of the following discussion if you "sing along" just as these early time on the horlzootal rods. frequency
speech researchers did, producing the speech sounds yourself and feeling the en the vertical axis, and arnplltWe
various articulatory maneuvers necessary to speak them. (lnternty) oo a cdcr cr gray scale.

332 CHAPTER 11
FIGURE11 .9 VoW'el soundsofEng1ish ,

shoviii ng ho w the frequMcioo o f the first for-
rnant (F 1) and s9oond fo rmant (F2) r9atetp
how high or IOV\I' the tongue is (f1) and h ow
far forward or b ack the tongue Is (Fil. The
d ashed line illus trateis limits o n how far the
tongue m oves when vowels are produced . beet r- - - - - -- --
For example, the VOWQls '99' and 'oo' are
produced with the tongue verf high in the
I
I
' ...
mouth. In the Ca.SQ of '98,' the tongue is fur 'I \
forward; but for 'oo, ' the tongue is far back.
.g
)j
,;:- t
"'
6 bit I
I
I
"' "'.,:'' bird

I
I
I
; bomb
b:d. . ,,"'
b aug ht
Lo w - - - - - - - - F1 frequency - - - - - - - Hi gh
High - - - - - - - Tongue position L ow
(u) Voiced
'bah ' 'd.1h ' Vowel sounds a re all made w ith a relatively open vocal tract, a nd
they vary m ostly in how high or low and h ow fur fo rw ard o r back th e
tongue is p laced in the oral tract, a long w ith w hether or n ot the lips are
rounded . We produce the 1 ee1 sound in the word beet by placing th e
ton gue up and forwa rd, the 'aw' 111!Wu.ght'Qy rnovin g the tongue d olvn
and back, and the 'oO in boot by moving the tongue up a nd back w hile
rounding the lips (Figure 11.9).
We produce consona nts by obs tructing the vocal tract in some way;
(b) Voice less and each consonant sound can be classified according to three articula-
'poh' 'tah' ' kah' tory d im ensions:
1. Pince ofarticulation (see Figure 11.6b). Airflow can be obstructed
• At the lips (bilabial speech sounds: 'b,' 'p,' 'm' )
• At the ju.st behind the teeth (alveolar speec h
sounds: d, t, n )
• At the soft p alate (vela r speedl sounds : 'g/ ' k,' 'n t{)
2. Mminer of articrdation. Airflow ca n be
FIGURE 11.1 0 English stop consonants • Totally obs tnicted (stop s: 'b,' ' d,' 'g,' 'p,' 't,' ' k') (Figure 11 .10)
rnay b9 {a) voCed , as In 'b<ID, ' 'dah,' ;;md
• Partially obstructed (fricati ves: 's,' 'z,' 1f,' 'v,' ' th,' 'sh ')
·gah ': or VJ ) voiceless., as In ·pan,· 'tah.'
and ' kah. ' lhie main artic ulatory diffo9'r9/1ce • Only s Hghtly obs tn1cted (laterals: ' I,' ' r'; and g lides: 'w,' 'y)
between vobed and voiceless stop conso- • Fi.rst blocked , a nd then al10V1>-ed to sneak through (affricates:
nants Is that. for the latter. talkers delay vibra- 'ch,' 'j')
tion o f the voc.:..'11 folds by ctbcut o m1-twentieth
o f a second after opeir1ing the vocal tract to • Bloc ked at flrs t from going throu gh the m outh, but allowed to
bQgin the sou nd. go thl'o ugh the nasi.11 passi.1ge (nasals: 'n / 'm,' ' ng')

100 FIGURE 11. 11 BecaUSQ the Spe9Ch

00 signal has oo r"'fk.-my redundant acoustic
c haracteristics. listeners c.:.-m under·
,., SJ stand sp99ch when Qll 9nergy 9ithq
below Cf abov9 1 BOO Hz ls rerroV9d .
7tl
'" 60
lh9 gretiln line shows p9rlcrmance
whEf'I energy above different frequen ·
"Ji Oil c ies (Mgh pass) Is available to 1St9ners.
.5 and the red line shows PQrforrnanc.:i
40 wh.:n en9'gy below differ€<'1t frequeo·
c ies (low pass) Is present. The rGod
& 30 c utve shovvs intelligibility when th9
;)J
c utoff is above the given x value, and
10 thGo grQQfl c urv9 shOVtls intelligibility
0 wheo the cuto ff is below the given x
100 200 300 400 600 SJO 1000 ;)JOO 4000 w:JJ 10,000 value. Notice that inteilligibillty is QXC.91·
Cutoff frequency (Hz) lent Vvhen all energy is rtiomoved M ow
about 1500 Hz (gr13en). and also 'ltVtlsn
all .enggy is removed above about
2000 t-tz (rocf). (After Fletcher and GoJt ,
l QW.)
3. Voicing(""' Figure rl.10). The vocal fo lds may be
• VibratD;g which can be felt by a finger on the
th roa t: b, m, z, I, r ')
• Not vibrating (voiceless consonants: 'p/ 's,' 'ch')
1l1ese speech sounds from English are o nly a small sample of the sounds
used by languages around the world, Most la.nguages use fewer consonants
vowels than are u sed in English _Some so unds are ciuite comm o n across
langua ges, and o thers, s uch as English ' th' and ' r/ are fairl y uncvm.m pn
around the world. When ma ny or most lan guages include pa rtk u.krr SpJ.ech MITTolAppa
sounds, often the reason is that the differences between them are particululy
easy to perreive. Humans mus t comm1ulicate in difficult environmmts where
the listen er is far away or there are man y competing sounds. To be effective,
speech sound repe rtoires of languages have d eveloped over generations of
individuals to indude mainly sounds that are relatively easy to telJ apart.
In addition to usi ng relatively easily d istinguishablesounds,.a nothe:i· rea-
son speech provides effective comn"lunication is that all distinctions behveen
vmo1,·els and consonants are signaled \vi th multiple differences bet\veen sounds.
Because more than one acous tic p roperty can be used to tell h '\"osounds apart,
di stinctions are s ignaled redundantl y, and t11is redundancy he lps listeners.
The speech signal is so redundant that if we remove all ene rgy below 1800
Hz, lis teners will still perceive speed1 nearly perfectl y, and. the same is true if
we remove all energy above 1800 Hz (Figure 11 .11 ).
Speech Perception
Speech production is very· fast. In casual conversa tion ""'"e produce about 10--15
consonants and vowels per second,. and if we're in a hurry we can d ouble this
rate. To ad1ieve thi s acoustic feat, our articula tors (tongue, lips, jaw, and so
on) must d o many differen t things very quickly. However, articulators can
move only so fas t, and m.ass and inertia keep articulators from getting all the
way to the position for the next consonant o r vowel. Experieni...-ed talkers also
adjust thei r production in anticipation of where articulators need to be nex t.
ln these ways, producti on of one speech sound overlaps production of the
coarticulation The phancmenon In
next. This ove rlap of articubti on in space and time is ca lled coarticulation. speech ""1ereby attnbutes of succes-
As we htrn from s peech production to speech perception, we will find that sive speech units overlap In artlculalcry
although coarticulation does not cause much trouble for listeners tmders tand- cr acoustic pg.ttems.

334 CHAPTER 11
'bah ' '<Lili' ing speech, it has made it harder for s peed1 perception researd1ers to
explain how ""·e d o it.
r. ,.___ 1'111--•W••""I
J r;;;; F,
F
COARTICULATION AND LACK OF INVARIANCE To get a bette1· sense
of w hat coarticulation is, say the worcl moody a few times, and note the
Iii p2
1
acti vity of yo ur tongue as you form th e ' d ' sound. You will find that
it starts in the back of yom mo uth, w here it mus t be to fom1 the 100'
vowel so und, then it tou ches the alveolar ridge just be hind the teeth
to fom1 the 'd / and finally it ends up toward the front of the mouth to
lo---·
form the "ee.' Now say the n on sense word eedoom. Revers ing the vowel
r·---· 111
..... F
sounds m eans sending the tongue on the op posite frorn front
to alveolar ridge to back. As a res1tlt of the very different pa th taken by
the tongue in these two utterances, the acoustic qualities of the 'd' in
I
I •••• i•••• "•••11: •
2
1
the two utterances are quite d ifferen t.
This contex t sensitivity is a signa ture property of speech . One might
expect context sensi tiv ity to ca u se a s ignifica nt problem fo r speech
perceivers, bec.1.useitmeans. there a re no "invariants'' that '"1ecan count
on to uniquely id entify d.iffere.nt speech sounds. For 'b/ ' d ,' 3fld 'g' in
Figure 11.12, notice tha t the of the first formant (F 1) is. pretty mud1
fltll••• ...--..
t ,....---.flll••• 1'1111••1111
Fl the for ail three consonants when they precede the sam e \'O\vel. F 1
1
F
2
is helpful in telling b,' 'd/ and 'g' apart from other speech sounds, but
J: it d oes no t help much in tellin g these sounds apa rt from one another.
An F1 like tha t s hown in Figure 11.12 is necessary fora s.ound to be 'b,'
. .. . . . .. . . . .. . . F1 but it is n ot s ufficient to infonn the listener tha t the sound is 'b,' a nd
Tun• not ' d ' or 'g.' F2 is very imp ortant for telling 'b' from 'd ' fr om 'g,' but
wh at F2 tells the listener depends on th e quality of F3 and the nature of
FIG URE 11 .12 Spectrog rams o f 'd'
sounds (c9ntgr column), along with the following voweL
stop-consonant ccusins 'b' (!aft) and Exp laining h ow lis te ners understand speech despite all this variati on has
·g • (right). O'!anges in formants across been o ne o f the m ost sign ificant challenges for speech researchers. Contex t
time (formant transitions) for these sensiti vity due to coa rticulati on also presents one of the grea test difficulties
sounds differ dramatically depending in d eveloping computer recogniti o n of speech . \ Ve cannot program or train
o n the fo HowinQ vovvets: 'ah' (top rOYV),
'oo' (middle), and 'ee' (bottotT\). a computer to recogn ize a s p eech s ound----consona_n t or vmvel -w itho ut
al.so ta king into cons id eration w hich speech so und s p recede and follow
tha t sou nd . And we canno t id e ntify those preceding and fo ll owi ng sounds
wi thout a lso takin g into con side ration w h ich sounds precede a nd follow
the m , and so on.
CATEGORICAL PERCEPTION [fit is cha llen ging to get computers to recognize

speech, why d o children ac'-1uire th is ability so early and easily? Something in
the a.caustic signal must lead them to pe rceive the ' d ' in deem, doom, and dam
as the same consonant. Shortlv a fte r World War U, researchers invented ma-
chines tha t could produce spe;dilike sounds, and they s tarted testing listeners
to try to determine exactly wlmt the acoustic cues \Vere th.at enabled them to
dis tingui sh different speech sounds. They folUld, for example, tha t by varying
the transitions of F2 and F3 , they could produce sounds tha t listeners reliably
reported hearing as 'ba h,' ' dah,' or 'gah' (Figure 11.1 3). (See Web Activity
11.2: Categorical Perception.)
TI1ese researchers knew that making small, incremental changes to simple
acoustic s timuli s uch as plU'e tones leads to gradual chru1ges in people's percep-
tion of these s timtill. For example, tones sound just a little higher in pitch \vi th
ead1 s maJI s tep in frequency. s peech sound s. were not p erceived
in this way. \\/hen researchers s tarted with a synthesized 'bah' and gradually
varied the fomK'lnt transitions m ovin g_ to a rCI, 'dah' at"'ld then 'gah .' lis tene r.s'
responses did not gradually cl-\.;1. nge fro m 'bah ' to 'bah'-ish 'dah ' to 'dah' to

8230336 Amma MUSIC AND SPEECH PERCEPTION 335
1,
Sounds ' \/ Sounds

like 'dah' \ like 'gah'
\
'-----------
Tmw
FIGURE 11 .13 The sound spootrograms at tha bottotTl o1this figure Indicate audi-
tory stirnu" that c hange smoothly from a dear 'bah' on the left through 'dah' to a
dear 'gah ' on the right. PQl"C€1ption, howe\'i'lr, do9S n ot chang9 smoothly. AH the
sounds on the left sound like a 'bah' cutvii:i) until we reach a sharp 'bah' - 'dah'
b ord9r. Listeners are much better at discriminating a 'bah' from a 'dah' thon at dis-
criminating two 'bah's or 'M'o 'gah's. The dashed black line indicates d iscrimination
performance. The Sl'tn1t\ Is true for 'dah' (red cu1Ve) and ·gah' (green c urve).
' d ah '-is h 'ga h' to 1gah.' Instead , lis tene rs identified these sounds as ch<:m ging
abruptly from one consonant to
Furthernlore, listeners appeared inc.apa.ble of hearing that mud1 of a nything
was differen t w hen two sounds were labeled as the sa me consorn:mt. In this
second part of the experiments, researchers played pairs o f synthesized speech
sounds a nd asked listeners to tell them apart. Lls te ners performed almos t
perfectly w hen detecting s mall differences between hvo sound s if o n e was
' bah' and the other was ' dah.' But if both sounds Vl'ere 'bah' or both were 'dah,'
performance dropped to n early chance levels (d ashed line in Fig ure 11.13),
even though the differences in formant trans iti ons in the first and second pai rs
of s timuli were equally large.
This pattern of results has come to be called categorical perception. As
illustra ted in Figure 11.13, three qualities deflne categorica l perception. The
first two were just described: a sharp labeling (identification) fon cti on and
discontinu ous di scrimination performance. The thir d d efinitio nal quality
of categorical perception fo llows fro m the first two: researchers can predict
di scri mina tion perfonn an ce on the basis of labeling d ata. In short, listeners
report hearing differences between sounds only when those differences would
result in different labels for the sounds, so the ability to discriminate so unds
can be predicted by how lis teners label the sounds.
HOW SPECIAL IS SPEECH? Ca tegorica l percepti on of speech sounds is n ot

limited to the 'bah'/'dah' /'gah 1 dimension; it has been sh own for many dif-
fe re nt contrasts between sounds in English, as well as other languages. 1llese
categorical perception For speech
findings, along'"; th the failure to find invariants that distinguish speech so1.md.s
as well as other complex sourds and
from each o ther, led many speech researche rs to s uspec t that lnm1ans had Images . the phenomeoon by whic h
evolved s pecial mechani sm s just for perceiving speech. One very influential the dlscrlmlna:tlcn of Items Is no better
version of the "speech is s pecia l" idea, called the " rnotor theory" of speech then the ability to lab<j Items.

336 CHAPTER 11
FIGURE 11 .14 The rnotorth90ry

was bcistered by, among o thtt" things.
a somawhat pecutlar fl nding b y McGurk
and MacDona k:J (19 7 6). Experimental 1. A videotape .shows
Sl.lbject s wtire sho wn a video of a a }"'E'l"Son rep E"ating
son th!:! syllable ' g.cth' OVQI'" and the sotmd 'Gah.'
over. The audio track. however. was
pla)oing the sound ' bah' repeat9dly.
lncredibty, s ubjects con:sistentty report- 2. An audio track plays
ed hearing a third S)'11able, 'd ah ,' even if the sound ' Bah .'
they kn9'W h ow thQY WGre being f ool9d
(because they heard ' bah' w hen th€1oy
c losed their eyes}. (fhls effect really
has to be expgi,gncQd to be
t ty n now in Web Ac tivity 11 .3: The
Mc Gurk Effect. )
perception (Liberman and Mattin g l)'; 1985; Liberma n et al ., 1967), he ld tha t

processes used to p rod uce speech sounds can som ehmv be nm in reverse to
understand the acoustic speec h s ignal . (Figure 11 .14).
Over time, however, a num ber of prob lems wi th the m otor theory cropped
up . First, it hlrns out that speech pro...-luction is a t least as complex as speech
perception, if n ot m ore so. Every a spect of the acous tic s igna l relates to a
particular aspect of the vocal tract. So if the acoustic s ignal is complex, this
us t ¥ thares ul! of complexity in p roduction . Trying to exp lain
FIGURE 11 .15 Speech perception speech percep tion by re ere nee to prod uction is a t least as d ifficult as explain-
by n onllumans. (8} Japanesie quail can ing speech perception on the basis of acoustics alone.
te3ITT t o t€111 'd ' fro m 'b' and 'g ' praced-
A second reason to d oubt that p rocesses for perceiving speech are unique
ing different vawels just as human s do.
(b) C hinchillas h avg shown c at9g0ri- to humans is tha t numerous d e m ons trations have s ho\."11 that n onhuman
c al r:ieroeption o f sounds varying from ani mals ca n learn to respond to speec h signals in much the sarne way that
'dah ' to 'tah.' (Part a oourtesy of Keith human listeners d o {Kluender, Lotto, and Holt, 2005; Kluender et al., 1998). For
R. Klu9f'ld'91': b court9Sy o f Annie Huy- exa mple, Japanese quai l {Fig ure 11 .15a) can be taught to tell ' d' from 'b' an d 'g
ler. Pioneer Valley C hin chillas.)

FIGURE 11.16 Pec:ple categcricalty

pere&tve changeis between imageis
across the sa me sort of acoustic variation depicted in Figure 11 .12 (Kluender,
o f familiar animals such as monkeys
Diehl, and Killeen, 1987). Chinchillas (Figure 11 .15b) have alsos h°'"'·n classic and oows. The labels that obs9rVQl"S
ca tegorical-perception effects (Kuhl, 1981; Kuhl and Miller, 1978). use sh ift abruptly bet\-Ve9n
Ftuthemlore, we n ow know that categorica l pe rception, one of the defi- and •oow" vvhen they ld0ntify Images
niti onal characteris ti cs o f s peech that was thoug ht to be so unusual as to likQ those in tha s,g1ies. shown
ObsBVers also ara bette.r at dlsctimi-
require a special tnll limited to speech s otmds.
nating two Images Wien they labBI one
O ther types of auditory sti muli,, such as musia.i.l interv.:ils (Sm ith et al., 1994), as ·monkg;/' and one as "caw." (After
are a lso pe rceived c."l tegorically, as are vis ua l s timuli su ch as familiar objects R. Garnpbell et at .. 1 007 .)
(Newell and BtUthoff, 2002), human faces (Levin and Beale, 2000), and facial
exp resslons (De Gelder, Teunisse, and Benson, 1997). People even perceive
differences behveen familiar animals ca tegorica!Jy (R Ca mpbell et al , 1997)
(Figure 11.16), and m onkeys perceive im ages o f cats versus d ogs categorically
(Freedman e t al., 2001).
COARTICULATION AND SPECTRAL CONTRAST Con temporary research has

turned increasingly to inves tigating how s peech perception is explained by
genera l '"'ays that hearing, and percepti on m ore broadl}'i works. For exampl e,
the perception of coartic ulated speech appears to be a t lea.st partially explained
by som e fundamental principlffi of perception that you 1ve already read abo ut.
Le t's tum to o ur exa mple o f s top conson;:m ts such as 'b ' and ' d .' T'"'O
of the acoustic feah1res tha t contribute to the perceptio n of 'b,' as contrL.lSted
w ith the perception of ' d/ are the onset freq uei11.."y and trajectory o f
the second forman t (F2 ). Beca use o f coarticuli."ltion, production o f one 'eeb.ili' 'oob.ih'
speech sm.md a lways affects production of the next sOLmd ; fo rmants
for one sound always are m ore like (assi milate to} th e sounds tha t
precet..1.e and follm"'·· The onset of F2 varies depending on whether 'baJ11
or 'dah ' fo ll m··.rs vowels such as 'ee' (hi gher) o r 'oo' (lo \\.'er). F,
Percept ion o f the syllables 'bah' a nd ' dah' works in a way that
nicely fi ts the facts of coarticulatio n . The very same F2 onset is heard
F,
as 'dah' following 'oo ·and as 'bah ' follow ing 'ee' (Figure 11.17). Wh.y
d oes perception work this way? Because roarticulation al ways causes
a speech sotmd to become more like the previous speed1sound, a ud i-
tory processes that enhance the contrast between s uccessive sounds Tune
undo this assi milation. Listene rs are more likely to perceive 'ba h' (low 'eedah' ' oodah '
F2) when preceded by the vowel sound 'ee' (high F2), and to perceive
FIGURE 11 .17 Because of coarticulation, consonant sounds such as

'bah ' and 'dah' ar.e acoustically very d ifferent, depending on the preceding
vowgl. Here, 'bah' is stiown following 'ee' (top l9ft) and ·ex:>' (top right). and
J
._:
F,
'dah' Is sho'Wfl following the same vowels Note that F2 in 'etibah' F,

(top left) is acousticalty identical to F2 in 'oodah' (bottom right). Listeners
F,
hear the s ame conscnant sound as 'b' following 'ee' and as 'd' following
'oo' because of the contrast betwe9n th€o spectrum of 'ee' and 'oo' and the
spgctrurn of th9 following oonsonant. Time

338 CHAPTER 11
'd a h ' (high F2 ) w hen preceded by 'oo' Oow F2) . Preceding sounds do not even
have to be speech sounds. [f, instead of playing 'ee' and 'oo' before syllables
varying from 'bah' to 'dah,' we present only a single small band of energy at
the frequencies where F2 would be in 'ee' 0 1· 'oo/ perception of the following
syllable changes just as it does vdth the full 'ee 1 or 'oo' vowel sound (Coady,
Kluender, and Rh ode, 2003). The onset of F2 fo r 'bah ' and 1da h' is perceived
relati ve to whether the preceding ene rgy is or h igher in frequency. We
perceive syllables s ud 1 as 'bah' and ' da h' in terms of th e rela tive cha n ge in
the spectrum-how the onsets contrast YVith the energy that precedes the m
(Alexander and Kluender1 2009).
You've encotmte red contrast effectsse\.-eral times before in this book- for
exa mple, brighb1ess contrast in Chapter 2. Remember that melodies are de fin ed
by changes behveen adjacent n otes, a nd not by the exac t n otes in particular.
While learning about vision, you saw many examples of contrast . Contras t
plays a la rge role in the pen::eptiOJ1 o f brightness color, and size, as well as line
orienkltion, position, curvature, depth, and spatial frequency. Here, spec tral
contras t helps listeners perceive speech, d e'.!o--pite the klck of acous tic invariance
due to coarticulation.
USING MULTIPLE ACOUSTIC CUE.S \Vha t do speech sound s, musical inter-

vals, and fo.ces all have in common? They are a ll stim uli that people have a
great deal of experience perceiving. We spend a large chmtlc of our waking
lives listening to speech <-tnd identifying people by their foC'E.'S. Another thi n g
tha t m akes distinguishing between individua l s peech soun ds a nd individua l
faces simi lar is tha t many s mall d ifferences must be used together in order to
d iscri min a te different speech so unds and different faces (e.g., s mall chan ges
in formant transitions a nd s m all cha nges in nose shape). At the same time,
oth er stimul us differences mus t be ignored so tha t multiple instances of the
sa me speech sound or multipl e images o f the same face ca n be classified
properly (e.g.1 acoustic varfation introdu ced w he n d iHerent spea kers utter
the sa m e speech sound, or image V'1riation introduced w hen a face is viewed
8230336 &jom different angles).
To the ex tent that perception dep ends heavily on experience, speed1 is
specia l because (1) humans have evolved tmique a na tom ical mad1inery for
producing it and (2) '\Ve spend a great deal of time practicin g the perception
of speech . The fact that there are no acoustic invariants for distinguishing
speech sounds is really n o different from many comparable si tu ati on s in
visua l perception. FoT example, \Ve saw in Chapter 6 that on e particular cue
for de pth perception m ay foil u s1 but by taking multiple cu.es into l'lccotm t, we
rarely make large mistakes vvhen calc ulatin g d istance rela tions.
The compa rison with face recognition may be even more apt. Caitlin's nose
may be quite similar to Sara's n ose, Cai tlin'seyes may be exactly as far apart
as Hannah's eyes, and Caitlin's mo uth may be shaped jus t like Emily's m outh.
But given enough experience wi th all four faces (and enough experie nce \vi th
face recogniti on in general), we can use the pattern of facial fea tures to p ick
Caitlin out fr om a line up every time---even if she's covering her mouth with
Tun• he r hand. Tum.ing to speech perception, utte ra nces of the sy llables 'aba' and
FIGURE 11 .16 The sl rn!>e 'apa' can be distingu ished from each otller by a t least 16 different chamcteristics
tion between 'aba' 6eft) and 'apa' of the acoustic signal (Figure 11 .18 ) (Lisker, 1986)_ Listeners can make use of
(right) includes at 19as1 16 acwstic their experie nce with the co-occu rren ce of these multipl e acou stic differences
differences. Som e diffEfEf'ICes that to understa nd sp eech .
e asy to include duration o f the 1irst
vowel. duration of the interval between To s tun up1 we d on' t need individual acoustic invariants to distinguish
s)oilables, and the presence of low-fre· speech sotmds; we jus t n eed to be as good a t pattern recognition fo r sotmds
quencyenergy in the middle o f 'aba.' as ""·e are for visua l irnages. A nd one of the things that the billions of neurons

MUSIC AN D SPEECH PERCEPTION 339
in the brain d o bes t is integratin g nuJtiple sources of informa tion to recognize

patte rns {Kluende r and Alexnnder, 2008). Computer simulatio ns of neural
connectivity (n eural ne t work m odels) d esigned to min1ic the use of multiple
attributes and associations be tween a ttributes are arnong the m ost s uccessful
artificial pattern recognizers yet invente d . lnterestingly, rnany neural ne h'llork
m odels (e.g ., tl1ose d eve loped by J. A. Anderson et a l. [1 977) or by Dampe r
and Hamad [20CXJ]) also s how ca tegorical-perception effects w hen the rno dels
are trained on en ough exan1ples.
Learning to Listen
In our discussion of vision earlier in the book, \•vesm"'· tha t experience is incred-
ibly imp ortant for visual perception, particularly the higher-level perception
of objects and events in the world. Experience is every bit as imp orta nt for
auditory perception, especially for the perception of speech. And tui.like vision,
experience w ith speed1 begin s very early in development. In fact1 babies gain
significant experience w ith speech even before they' re bornt
Measurements of hemt rate as a n indicato r of the ability to n otice change
between s peech sounds have revealed that late-tenn fetuses can discriminate
between different vowel .sounds (Lecanu et et a.L, 1986). Prenata l experien ce
with speech sounds appears to have consid erable influence on subsequent
perception. For one thing, a newborn prefers hearin g he r m other 's voice over
other women's voices (DeCasper and Fifer, 1980). When 4.-day-old infants in
Paris were tested , they preferred hearing French instead of Russian (Mehler
et a l., 1988). Perhaps m ost amazingly, newbo rns prefer heari ng particular
child ren's s tories that v..iere read al oud by their m othe rs during the third
FIGURE 11.19 How we hear speech
trimester of p regnancy (DeCasper and Spence, 1986). sounds d0pends on our experienc e
w ith the SP9Qeh sourds o f our first
BECOMING A NATIVE LISTENER As we h 1we seen, speech sounds can dif- language, Because o f experience w ith
fer-fu..inany> a . Aroustfc d.f.fkrences tha t 1na tter critically for one language one language, it is often diffic ult to pef·
may be irre levan t or even distrncting in an o ther language. Fo r example, the c eive and produ:::e distinctions in a new
language. For @rampe, mo st Japanoo.e
English language makes use of the distinctio n between thesmmds 'r' and 'I/
piecple learning English as a S9cond
whereas these two sounds are both very si milar to only one sound (cal led a language have trouble distinguishing
''flap ") in Japanese. As another example, Spa n ish is one of many lang uages between 'I' and 'r ,' both whai th91 lis·
that uses only the five vowel sounds 'ee' (as in tai and w hen thgy speak.
beet), 10 01 (as in boot), 'ah' {as in bomb), 'a'/ (as
in and 'oh ' (as in bont), whereas English
employs numerous other vowel sotmds.
As an yone w h o has s poken to a native
Japanese spea ke r in English knows, the ' r ' / 'l'
d istinction is very difficult for Japanese people
to pick up when they learn English as a second
lan g uage (Figure 11 .19). Beca use the differ-
ence between '1' and 'r ' is irreleva nt to native
Japanese speakers when they're learnin g their
native lan gu age, it is adaptive fo r them to leam
to ign ore it1 thus all ow ing them to focus o n
speech sound distinction s that are important
in Japanese. When people have spen t m ost o f
their lives listening to Japanese and not hearing
the diffe1-ence between 'r ' and 'I/ we are n o t
surprised that they have difficulty learning to
produce the 'r ' and 'l ' By the same token, a na-
ti ve Span ish speaker who complains that your
d og just "be.:-tt" him is probably not d,Umi ng

340 CHAPTER 11
100 tha t Rover threw a punch; rather,, the difference betv.·een 'ee" and 'ih ' is
less percep tible to the Spanish s peaker, beca use both of these English
so unds are simila l' to the Spanish 'ee.'
Interesting ly, s tudies s how tha t infants begin filtering out irrelevant
.5"' acous tic differences long before they begin to utter speech sounds (even
l before their babbling s tage). One study found that by 6 months o f age,
infants from Seattle were more likely to notice acoustic differences that
di sting uish h vo English vowe ls than to notice el1uiva lent differences
J behveen Swedish vowels, and infants from Stockholm were more likely
to notice the dlfferem.-e. between h'l-·oS\·ved ish vowels than the difference
twoJlnglish vowels (Kuhl et al., 1992). Twllng of perception for
consonan ts appears to take a bit longer to develop, but by the time infants
are 1 ye.u old, they have also begun to ign ore, jus t as their parents do,
6-8 S-10 10-12 n - 12 consonant distinctions no t used in their native language (Figure 11.20).
months months months
..._,_,
months O f course,, it is possible, w ith much training, to learn to perceive and
Eng.Ush infants Hindi infants
produce speec h sound distinctions that you've spent most of your life
igno ring. As yo u might expect, the longer a person uses only her first
FIG URE 11 .20 Hindi h as a dental language, the longer it takes to learn to produce and percei ve sounds
stop consonant produc&d w ith the from a second language (Flege, Bohn, and Jang. 1997i Imai, Flege, and Way-
tongue tip touc hing th9 t199th and a
retroflErx stop consonant that requires 1..'l nd, 2002). Many s tud.i es have been aimed at determining w ha t makes ne\>v
the tongue t o bend up and b ack in the distinctions hard or easy for second-la ng uage learners to pick up. Learning
rnouth. Adult speakers of English hear is most difficult w hen both of the sounds in the second lang uage .are similar
both o f thoo.e sounds as 't' because to a single sorn'd in the firs t langi.Mge (e.g., 'r' and 'I' for Japanese speakers
they are sirnilarto the English 't .' \'VhQn
Weirker and TMs (1 9 84) testoo infants
learning English). Learning is easier if the h\'O new soi.mds are both unlike
from English·spgaking famili9 s In Van- any sound in the native lang uage. Fo r example, na ti ve English listeners have
couver, the youngest (6-8 m onths old) no probletns distinguishing dick sounds from Zulu because Zulu clicks are so
re liably respondeid to the difference unlike any English sounds. Leaming a lso is easier if t\.,,.·o new so unds from a
betwwn thQiSQ two Hindi soun ds. By 1 new language diffe r in the same vvay thLlt two sounds from the firs t lan guage
year o f age, h owel/l'::f. the infants were
differ.
mimicking th9ir p arents' b 9havior by
ignoring the distinction betwee<l the Picking up the distinctions in a second languas;e is easiest if the second
two srunds. bnguage is learned a t the same time as the first. This is why language immersion
programs are becoming popula r in preschool and elemenhuy cturiculurns of
multicultural countries suc h as the United States. ll'l.e d o"vnside to this strategy
is tha t kids lea ming multiple languages at the same time us ually take a little
longer to m aster each of the languages than do children learning only a single
language (Bialystok and Hakuta, 1994). This is a natural consequence of having
to learn not one but two sets of n1les about when to ignore and w hen to pay
attention to speech sound diffet·ence.s (as well as learning two vocabularies,
two sets of grammatical rules, and so on).
LEARNING WORDS 1llus far in this chapter, '\ve've lea med many things about
the nah.m;: .and complexity of s peech sounds, and about how listeners perceive
consonants and. \-m,:els. But the w hole point of producing and percei ving these
speech sounds is to put them togethe r to fo rm words, w hich are the tmi ts o f
language tha t convey mea ning. H ow do novice language learners (infan ts)
rnake the leap f.rom s treams of meaningless speech s01.mds to the mea ningful
groups of speech so unds that we call words?
Fi rst1 le t's s ta te the obvious. Just like strings of musical n otes, no s tring of
speed' sounds is inhe rent! y meaningful. 1l1e s tring /d' -'aw'-'g' becomes mea n-
ingful to English-spea king infants, but to French-speaking, Spanish-speaking,
and German-s peaking infants th is s tring remains com pletely mea ning less
beca use thei r parents refe r to their fur ry house compani on as driw, perro,
and Hund, respectively. )nfants in different places in the world rnus t learn the
words tha t are s pecific to their native languages.

FIGURE 11.21 In the sentence '\ovhere are the silences

between words?• we cari SQQ that there are no breaks
between the sounds of one spoken word and the sounds
o f the next. Infants can use their experience with particu-
lar sequooces o f speech sounds to loom about boundar·
Jes between words.
Wherearethe e i I en c e s be t w een wo rd s 1
We've already seen how a series of consonants and vowels within a single
word tend to "run into'' one another because of coarticu.lation. It h.m1s out the
sihrati.on is n ot much better for a series of words fomtlng a sentence. TIUs fact is
easily seen in Figure 11.2.1 : without the lette rs at the bottom o f the figure, you
would have n o idea where the spoken word "where" ends and "are" begins.
Interestingly, our perception usually seem s at odds with
this acoustic reality. When we listen to someone talking
to us in o ur native language, indiv idual words seem to (a)
stand out quite clearly as separate entities. But listening tok ibugopi lagi kobatipol utokibu
to someone speak in a different language is o ften a very
different experience. Lis ten to the Chlnese a nd Arabic gopilatipolutokibugikobagopila
speakers in Web Activity 11 .4 : Word Breaks. Unless you
have experience with these languages, it probably sounds
gikobatokibugopilatipolugikoba
as if they include lots of really long words-som e as long tipolugikobatipolugopilatipolu
as whole English sentences. It may also seem as if people
in these cultures speak much faster than English speakers toki bugopi Iatipolutok ibugopi Ia
d o. Native Chinese and Arabic speakers probably say the
same things about tl10 English language and English speak-
tipolutokibugopil agikobatipolu
ers. This is the situation faced by infants the world over. tokibugopilagikobatipolugikoba
To study h ow infants learn words from the continuous
streams of speech that they encounter in their environment, tipolugikobatipolutokibugikoba
Saffran, Aslin, and Newport (1996) invented a n ove l lan- gopilatipolugikobatokibugopi la
guage. Each of the words of this new lang uage had three
syllable>-for example, tokib11,gopila,gikoba, and fipolu. Next
they created randomized sequences of these novel words, (b)
with word s running together just as they do in fluently tokibugopilagikobatipolutokibu
produced sentences (Figure 11 .22). A sample sequence
would be " tokib11gopilagikobatipolrtgopilatokib11tipol11g1kolxi." gopilatipolutokibugikobagopila
Eight-month-old infants listened to a 2-minute sequence gi kobatok ibugo pilatipolugikoba
such as this while sitting on their parents' laps. After this
ti pol ugikobatipol ugopilatipol u
FIGURE 11 .22: Eight-m onth-old infants can learn to pick
tokil>agopifatipolutl') · ugopila
out 1NOrds from streams of oontinuous speech based on the tipolutokibugopilagikt1batipolu
extent to \l\lhich successive syHables are predlctat)e or unpre-
dictable. While sitting on their parents' laps, infants heard tokibugopilagikobatipolugikoba
2-m!nute sequooces o f syllables (a). In the second part of the
QXperiment, Infants were familiar with thr&e·syftable sequences tipolugikobatipolutokibu gikoba
gopila. gkooa, tiPol.IJ that thQY had h<>ard beforo (b).
but they notCed that they had nsver heard other syllable com- gopi latipolu gikobatokibugopila
binations tpoitlto, bugopi, kd:iallj.

342 CHAPTER 11
brief period of 1eantlng, infants heard either "tupiro" (one of the novel words)
or "pabilur'' (a new combination of the same syllables used to produce the
"real" novel words). The infants listened longer to the nonwords than to the
words, indicating that after just 2 minutes of exposure, they had already begun
to learn the words in th is new "language.u How did they do it?
Saffran and her colleagues suggest that the infants in their study learned
the words by being sensitive to the statistics of the sequences of soWlds that
they had heard in the first part of the experiment. In the real world of language,
words are simply sequences of speech sotmds that tend to occur together. Other
sequences occtu together less often. For example, think about the sequence
'p'- 1r'- 1 ih' -'t'-'ee'-'b 1 -'ay'-'h'-'ee.' An infant will hear the sounds making up
the word pretty in many different contexts ("pretty dress/' "pretty good/' and
so on) and the so\U\ds making up the word baby in o ther contexts (e.g., "good
baby," "baby doll"). In contrast, the sequence 't'-'ee' -'b' -'ay' will almost never
be heard in any other context, because no English words have this sequence of
syllables. Saffran (2001, 2002) suggests that infants learn to pick words out of
the speech stream by accwnu.lating experience with sounds that tend to occur
togetherj these are words (at least to babies). Fo.r example, infants eventu..1.Uy
split the 11word 11 allgo"e into two as they acquire more linguistic experience.
When sounds that are rarely heard together OCClU in combination, that's a sign
that there is a break between two words.
Speech in the Brain

Our earliest understanding of the role of cerebral cortex for the perception of
speech and music was gained through unfortunate "natural" experiments in
which people had lost their ability to tmderstand Spe€ch following stroke or
other brain injuries. However, it is difficult to draw strong conclusions about
brain processes from brain injuries. Brain damage from stroke follows patterns
of blood vessels, no t brain function. Damage from stroke might cover just part
of a particular brain function, leaving some of the hmction Wldamagedj or
damage could cover a wide region that includes some or all of a particular
brain hmction, as well as all or part of other functions. Performance follow-
ing brain damage, along '.vi.th later experimental findings, has taught us that
different hemispheres of the brain are better at doing some types of tasks. For
mo6t peop).e, the left hemisphere is dominant for language proc-essing. The
development of techniques for brain imaging, such as positron emission to--
mography (PET) and hmctional magnetic resonance imaging (fMRI), has made
it possible for us to learn more about how speech is processed in the brain.
As we would expect on the basis of w ha t we learned in Chapter 9, hearing
Sylvian
sounds of any kind activates the primary auditory cortex (Al). Further, we
fisrn"'"-.. learned that the processing of complex sotmcls re.lies on a dditional areas of
cortex adjacent to Al. These nearby areas of auditory cortex, called belt and
parabelt areas, as shown in Figure 9.21 and Figure 11 .23, often are referred
Superi"'/ to as 11secon dary" or "association" areas. As one might expect, we see these
t.mpc•al
gyros Parabeltarea areas activated when listeners hear speech and music. A t these relatively
early levels o f cortical processing, acti'1ity in response to music, speech, and
FIGURE 11 .23 The superior tem-
poral gyrus aw:tands along the upper
o ther complex sounds is relative ly balanced across the tw"o hemispheres
surface o f each temporal lcbe. Primary (Binder, 2000).
auditory cortex {A1) and the adjacent Because we know tha t language is typically lateralized to one hernisphere-
b9lt area are hiddgn inside th9 SyMan usually the left side for right-handed people-processing of speech should
fissure, with onty the parabeilt a100 become more lateralized a t some point because perceiving speech is part of
expos9d tO'W'a.rd the post"n1or sup9ficr
temporal gyrus. Portkms of the super1or understanding language. One challenge for :researchers ls to create stimuli
temporal gyri (left and right) are actlYe that have all the com plex properties of speech without being heard as speech.
whm wg llshm to oornpl'9x sounds. We already learned that listeners are very good at understanding speech even

8230336 AMma Appa
und er adverse circumstances when some p.:1rts of the signal a re m issing or

distorta i , so this capabi.lity makes it pretty difficult to construct stinn.1Li that
are co1nplex like speech "''ithout being heard as speech.
Rosen et al. (2011) developed a particularly clever way to tease apart
cortical resp onses to acoustic complexity frorn responses to speech p er se. As
Figure 11.24 sh ows, they began with COff1plete sentences that included natural
chan ges in both arnplitude and frequency (Figure 1l .24a), altho ugh they re-
placed voicin g vibration wi th noise. Although u sin g n oise made the senten ces
sound whispered , th ey \.\.'ere pe rfectly intell igible. Next the resea rchers took
away d 1nn ges in frequenc;-; leaving be hi nd only changes in amplih1de (Figure
11 .24b). Then they took avvay chan ges in amplitude, leaving only ch.:""lnges in
frequency (Figure 1L24c)_Finally, to match the amount of acoustic complexity
in speech, they created hybrid "sentences'' by adding the amplitude changes
of one sentence (Figure ll.24e) to the frequency changes in a nother (Figure
1l .24a). l11ese h ybrid s (Figure ll .24d) were just as complex as the sentence in
Figure 11.24n, but they were completely unin telllg ible. 1l1e resea rchers played
these fou r types of sentences (Figure ·11 .24a--J) to listeners w hile their brains
were being sca nned using PET.
(a)
"The wife h e.lped her husb an d." Frequ ency chan ge15
Arn p litude changes
(b)
(c)
Frequ enC)' ch.anges
No amplitude changes
Hybrid
Frequ en cy ch ange15
\ \'rong arnplitude changes
FIGURE 11.24 Stimulli created t o measure cortical
resp onses to acoustic comploaxity versus respon ses
t o speech. (a) This $p€1Ctro;iram r€iJresents ru-i intact
intelligible sentence w ith natural chang€1s in frequency
(e) and amplitude. f/>--9} In thee.9 spgctrograms, the sam9
sentence is now uninte.lligit::te b9causg €lither frequEncy
c hang9S WGre removed IP) or amplitud8 chang.;;,s WfilfQ
removed (c). The spectrogram In (o'J has frequency
changes from t;a) but amplitude c hanges from (e). While
(dj Is QCIUal to (a} in acousticcornpl9Xity, it is uninte.lliglble
because o f the misrnatch between changes in ampli-
tude and fraqu9ncy.

344 CHAPTER 11
FIGURE 11.25 Substantial neural actMty was obs9tVed in both left and
right superio r temporal lobes (red) when listengs heard s.entenoes
such as the on9 shown in Rgur!:I 11 .24d, for 'Which amplitude and tre-
quEncy c hanggs came 1ro m different st:'lntEf'lces. An increase in activity in
only the languag0-dominant left anterbr superior temporal lobe (purple)
was evident ·wtu:in thc:a sentences were lntelligiblc:i (_figure with
matc hirq amplitude and frequency c hangoo.
8230336 Anma Appa
As Agure 11 .25 Ulustrates, neural activity in response to t.minteJ l.igible h ybrid

"sentences'' (see Figtue l l .24d) was fo und bilaternlly, with a little m o re activity
along the superior temp oral gyrus nmning alon g the top of the right tempo ral
lobe. Respon se.s in the left superior temporal gyrus becam e d ominant only
w hen sentences were intelligible because amplitude and frequency ch an ges
coincided properly (see Figure 11.2417). Fro m these find ings, it appears that
language-d ominant he misph ere resp onses depend o n lis teners using s peech
for linguistic tmdersrand ing, an d not from aco us tic complexity a lon e.
A deal of resea rch is being conducted to further our tmders tandin g
of how speech is processed in the brain on its way to becoming part of words
and sen tences. Fo r now, the evidence s uggests that as sounds becom e m o re
compl ex:, they are processed in m o re anteri or and ventral regions of the su-
perior tempor..11 cortex farther from A 1 {Patte rson and Johns rude.-2008;
Uppenkarnp et aL, 2006). When speech sounds become more d ea rly a part of
langu age, they are processed m ore anteriorly (forward) in the left temporal lobe.
This much appears true; hovvever, th is back-to-fron t pa:th may no t be all there
is to speed' perception in the brain. O ther brain regions are likely involved.
\.Vhat cou ld we lea rn if we could actually place electrodes right on top of
the brain? Prior to performing brain surgery to rem ove a tumor or to reduce
effects of very severe epilepsy, neu ros urgeons often place electrodes d irectly
on the s urface of the brain to localize regions of neural activity. Figure 11.26
illustrates an electrode grid placed over a region o f left superior p os terior
temporal lobe. Mesgarani et al. (2014) played 500 senten ces s poken by 400
diffe rent people to si.x human participants while recording directly o n the
s urfa ce of their superior temporal gy ri. At different s ingle elec trodes, they
fou nd that responses were selective for certain classes of speech sounds su ch
as fricatives, stop conson an ts, or vowels . In some cases, responses were e ven
rnore selective. For exarnple, responses to stop conson ants mi ght be selective
for only voiceless stops ('" p', ' t', 'k') or for on e p lace of ar ticulati on (e.g., 'd')
versus others ('b' and 'g').
Chang et al. (2010) p layed a series of synthesized sy llables va r}ing incre-
mentally from 'bah' to 'dah 1 to 'gah/ much like those in Figme 11.13, to patients
who were awaiting su rgery. The re.searchers found that, across populations of
tho usands of neu rons recorded by their surface electrodes, n eural responses
"'"·ere much like listening data for the sam e sy llables. Neura l responses were
very similar for pairs of stimuli tha t listeners would label as the sa me, as 'bah '

MUSIC AN D SPEECH PERCEPTION 345
FIGURE 1 1.26 Prior to perfoITTling brain surgery to

rernovg a tumor or to reduoo etfQCts o f wry sevge
.:pilepsy, neurosurgecns often place eectrodes dlrectty
o n the surface of th9 brain t o localize regbns of neural
activity. Before surggy, researchers C..."'m take advan -
tage of these electrod9s to record responses to sxperl-
mlfltal maw.rials. (Cc:urtesy of Rick L Jenison and
Richard A. R9'ale.)
or 'dah' or •'g ah'; responses were quite d istinct for pairs of syllables that were
acous tically separated by the same extent but '"'ere labeJed as different, as
'bah' and 1dah' or as 'dah' and 'gah.'
As with perception in generat cortical organization depends critically on
experience, and the perc;epti9n Of spee.:h ls trem·endously e.xperience-
dependent. Because \Ve kn ow that English a nd Hindi speake rs hea r sounds
like 'd' differently, we know that the way listeners discriminate 'bah' and ' da h '
depend s on experience. Chang et al. (2010) may have revea led a place in the !eh
temp oral lobe where experience with English 'b' and 'd ' s hapes neura l activity.
Cortical p roces.ses rein ted to speech perception could be distinguished fron1
brain processes that contribure to the perceptio n of other complex solmds in
hvo other ways. Listeners have a wealth of experience simultaneo usly hea ring
speech and viewing talkers' faces, and the tvkCurk e ffect (see Figure 11 .14) is
evidence of the prnfound effects th.at visual cues ca n have on the wny speech
sounds are pel'ceived. et al. (2007) also used grids of electrodes on the
s urface o f p osterior temporal cortex {see Figure 11 .26) in their experiments

ppa
346 CHAPTER 11
investigating cortical processin g of audio and visual speech They fo tmd

neural responses to auditory speech stimuli in the language-dominant hemi-
sphere were influenced substantially by sirnulta neous viewing of the lower
half of the face of a person eith er producing audible s peech or carrying out a
meaningless mouth m o tion .
Because vis ua l inform ation combin es wi th auditory experience when
we're perceiving speech, you may wonder what happens for people '"'·ho are
deaf. Zatorre (2001) s tudied a group of people \vi th cochlear implants who
pre·dously had been d eaf. Th.ese listeners exhibited increased brain acthdty
in the visual cortex when listening to speech . Zatorre hypo thesized that this
activation of visual a reas of the brain is the result of increased experience and
ability with lip-rea.ding for these previously d eaf indhiduals.
Finally, whenever people talk, they both he ar the solmds producing
and experience the rn ovements of speech production in their own vocal tracts.
One thing to le i.lm from future s tudies of brain acti vity is hmv these processes
of hearing and speaking interact. People h ave a lo t of experien ce with the
simultaneous activi ties of producing and perceivin g our own speech, so one
might expect to find brain regions wh ere these related activities combine. (See
Web Essay 11.1: Studying Brain Areas for Language Processing .)
Summary
1. Musical pitch has t...·o dimensions: tone height and tone chroma. Musical
Refer to the notes are combined to form chords. Notes and chords vary in duration and
Sensation and Perception are combined to form melodies.
Ccmpanlon Website 2. Melodies are learned psychological entities defined by patterns of rising
s1tes.s1nauer.comtwcife4e and falling musical pitches, 't'orith different durations and rhy thms.
for actMties. essays, stu:Jy 3. Rhytlun is important to music, and to auditory perception more broadly.
questions, and other study aids. TI1e process of perceiving sound sequences is biased to hear rhythm.
4. Humans evolved to be able to produce an extremely wide variety of sounds
that can be used by languages. 111e production of speech sounds has three
basic components: respirationTphona tion, and articulation . Speech sounds
vary in many dim ensions, including intensity, duration, periodicity, and
noisiness.
5. In tem1s of ar ticulation and acoustics, speech sounds vary according to
o ther speech sounds that precede and follow (coarticulat:ion). Because of
coarticula tion, listeners cam1ot use any single acoustic fea ture to identify a
vowel or consonant. Ins tead, liste ners must use multiple properties of the
s peech sign.."ll.
6. In general, listeners discriminate speech soun ds only as well as they can
label them. Dtis is categorical perception, \VhiCh also h as been shown for
the percep tion o f many other complex familiar auditory and visual s timul i.
7. How people perceive speech depends very much on their experience \vi.th
s peech sounds w ithin a language. This experience includes learning which
of the many acoustic features in speech tend to co-occur. Because of the role
of experience in how we hear speec h, it is often difficult to perceive and
produce new speech sounds fro m a second language fo llm\•ing experience
with a firs t language.
8. One of the ways that infants learn words is to use their experience with the
co-occ urrence of speech sounds.
9. Speech sounds are p rocessed in both hemispheres of the brain much li ke
o ther complex sounds, until they become par t of the linguistic message.
TI1en, speech is further processed in anterior and ventral regions, mostly
in the left supe1·ior temporal cortex, but also in superior posterior temporal
cortex.

lq
8230336 Anma Appa

CHAPTER "12
Shlnlchi fv1afl..zyama, i1, 2012

The Vestibular System and
Our Sense of Equilibrium
REMt:MBER WHEN YOU WERE A CHILO and you u sed to spin a round tmtil you
"veredizzy and cottldn' twalk s tra ight? Perhaps you even fell. \Vhy were you
dizzy? The sensations did not arise from one of the fi ve senses that Aristotle
recognized-vision , hearing, tou ch , taste, or s mell. YolU' dizziness a rru:e from
contributions of the vestibular organs to your sense of equilibrium.
11'1e vestibular o rgans a re a set of s pecialized sense organs loc<'lted in the
inner ear right next to the cochlea. Vestibular o rgans sense motion of the head,
as ""·ell as the orien tation of grav ity, <.'Ind rnake a pred ominant contribution to
om sense of tilt and our sense of self-motion . Taken togethe r, the senses of tilt
and self-motion comprise otu sense of spatial orientation.
FURTH ER DISCUSSION of the oochlea can be found in Chapter ll on

page349.
You may be askin g, nWait a mi nute, w hy didn't l lec1. m about the vestibular
system and ec:i uilibrium '"·hen I firs t learned a bo ut the five senses?" Good
question. Perhaps you should have. The vestibular "six th se nse" provides
hmdamenta1 contributions. that are often over looked . For example, the ves-
vestibular orgais The set of five
tibular system contribu tes to dear vision when we move and helps us maintain sense organs- three semicircular
b..'tlance w hen we s tand. And it is so crucial that some patients even report canals and two otdlth organs-
cognitive deficits when it fails. Yet desplte these essential contributions, the looated In each Inner ear that sense
ves tibular system toils in anonymity. Much of the tirne, we remain unaware head motion and head orientation with
of it until it stops wol'king p roperly. respect to gra>Aty.
The fundamenta l na ture of the vestibular sys tern is e1nphasized by the fact eq..1ilibrium our vestibular sense
that the vestibula r orga ns appeared very early in evolution ary history and have comprised of spatial orlentatlm per-
ceptlon-enccmpasslng our percep-
remaine d relatively tmchanged . The vertebra te fossil record s hows the presence tion of linear rnotlcn angular motion,
of distinct vestibul...1r organs in fis h at least 400 million years ago. 111e sys tem is and tllt- comblned With reftexive ves-
no t only an cient but largely automatic: vestibular perception is often relegated tlbt>ar responses like posture. vestlb-
to the attentional backg rm md, and many responses evoked by the vestibu lar ulo-autonomlc reftexes, and vestlbulo-
system a re reflexive. Though V..""e are a ll aware of th e n on11al hmdion of o ur ocular reflexes.
eyes and ears, only w he n we experie nce proble ms sud1 as dizziness, vertigo, spatial orientation A sense ocn-
spatial disorie ntation, imbalance, blurred vision, and /or i1lusory self-motion slstlng of three Interacting modalltles:
are we like ly to becorne a1."Utely a\.•:are o f our equilibrium sense. perception of Unear motion, angular
motion, and tilt.
Obviously, we can n o longer ask Aristotle-who is credited with first
ca taloging our sensory systems--w h y he did not include equilibrium or our vestibular system The vestibu-
lar organs and the neural pathWa)'S
ves tibular sense a mong the specialized sensory systems , but we can specu- directly assoo/ated with these sense
la te. It cer tainly is n ot beca use vertigo was u1Ucnmvn, since Aristo tle himself crgans.
described the vertigin ou s e ffects of alcohol. One expla na tion rna y be tha t it
vertigo A sensatbn of rotation
""·as not until the nine teenth cenhuy tha t scienti sts understood that the ves- a €pinning. The tenn Is often used
tibular system is a .specia lized set of sense o rgans. Until then, the vestibular more generally to mean any fcrm of
syste m had been considered an e ntrance to the coch le ct. In fa ct, the name dizziness.

350 CHAPTER 12
(a) FIG URE 12. 1 09tnonstration of th9 vestibulo-ocular

reflex. (a) As your fingErtlp m oves faster and faster in
front o f your fa09, thei fingGrtip begins to blur. l.p} When
you shake your head back and fcrth as if to say "no,"
Subject' s v iew the fing9rtip r" mains dear9f". (c} Text also remains
clearer during head shaking.
vestibtt!ar records th is error for posterity because

vestibt1le m ean s 1'entrance." But th is explana tion
is n o t e ntire ly satisfact o ry, si nce Aristotle h ad
cata loged other senses v-.<ithout <let.a iled anatomical
or physiological know led ge. (See Wade, 2000, for
a his torical review of vestibular knowledge prior
(b) to the nineteenth cen tury.}
Another expla1i..1tion be the inconspicuous
na ture o f our vestibular sen se. In fac t, as we'll
see, many responses evoked by the ves tibular
system are reflexive. For example, the vestibular
Subject' s view system helps us see clearly by reflexively ro tat-
ing the eyeballs in the sockets to compensa te for
head rotation-thereby h elping to keep \.isual
images s table on the retin a. This refl ex is cal led
the vesllbulo-ocular reflex (VOR).
To demons tra te th is to yourself, move your
han d a few inches back and forth in front of your
fuce (Figure 12.1a). Start slowly and then speed up
the m ovement. Focus on a fingertip and notice tha t
it starts to ap pear more and more blurry as your
hand m oves at a highe r frequency. This exercise
dem ons trn. tes the limits of smooth ptrrsuit, a form
of visu.."\l tracking that you learned about in C ha pter
Subject' s view 8. Now h old yo ur hand in &ont of your face and
s hake your head from side to side as if to say ''no"
(Figure 12.1b ). Again, start and gradually
increase the speed . At higher frequen cies of head
ro tation, you s hould no tice that each fingertip
stays in foc us more rea dily wh en you m ove your
head tha n w hen you move your hand. You ca n
co mpensa te fo r hea d m ove ment m ore readi ly
than hand m ovement because of a VOR that we
\.vi ll discuss in more detai l larer in this chapter.
Before \'Ve move on , th.ere' s one m ore lesson to
draw from this VOR exam ple. Rota te yom head
back and forth w hile you l'ead this tex t {Figure
12.1 c). Unti l now, beca use you were asked to a ttend to vision, you probably
were focused o n visual perception and didn't explicitly pe rceive your head
rotation. In fact, your hea d ofte n m oves arotmd, but you d o n ot ty pically
pe rceive your head motion, because the ••es tibular system usu ally performs
its job automati cally-with little, if any, conscious awareness. But now that
veslibulo-ocular reflex (VOR) A we've cal led yo m attention to it, as you continue to s hake your head, you can
s hort-latercy reflex that helps stabilize
vlslcn by counterrotatlru the eyoo pe rceive your head rotating, ca n 't you? Thus, it is n ot that yo u are unable to
when the vestlbular system senses perceive vestibular stimulation, but rather that vestibular s timulation is almost
hood movement. always relegated to the a ttenti ona l background (discussed in Cha:pter7).

THE VESTIBULAR SYSTIEM AND OUR SENSE OF EQUILIBRIUM 351
Evolution and Equilibrium

The vest ibular system is phylogentlcally old , >Mlich simply means that.
from an evolut ion pers).)ectiVe. It has been around a lc:ng time. All
mamm1"s have vestibular organs. In fact , 1"1 vertebrates, inc luding nsh,
amphibians , reptiles. and b#"ds have vest ibular crgans. Even sorne in-
vertebrates (e.g. , jellyfish) have gra\'iceptors, ard even bacteria need to
know up from do'lim to burrow dO"wnwards for SUIVival. All of these fac ts
suggest that equilibrium is rundarnent1"1y important.
But. relatlve to the vestibular organs of all other vertebrates, the
human vestibular organ s include the unique feature that the vertical ca-
nals are relattvety larger than in other sp:lc!es. This oontributes to higher
sensitivity, which is believed to yield enhanced head and eye stabiliza-
tion when running (S poor, Wood, and Zooneveld . 1994). This Is believed
to have contributed to enhanced exercise capacity (Bramble and Li-
eberman . 2004). wh ich, in tum, Is believed to hwe contributed to larger
hurnan brains (R1"chlen and Gcrdon. 2011). W ho knew the vestibular
system was so Important?
Vestibular Contributions to Equilibrium

The vestibular system p rovides the sensory foundation-alongside vision-fo r
spa tial o rientation, which includes perception of translation, ro tation, and tilt.
The vestibular system also makes crucia l con tributions to balance but d oes
not p rovide the sensory foundation for balance; kinesthesia d oes (see Chapter balance The neural prooesses of
13). While blind individuals and those \.v ithout vestibulnrsysterns can s tand , postural control by which weight Is
evenly distributed , enabling us to
individuals lacking kinesthesia typically canno t stand. In addition, the ves- remain upright and stable.
tibular system h e lps stabilize our eyes during hea d m otion . In this role, the
vestibular system makes crucial contributions to clarity of sight but does n ot kinesthetic Perception of the
position an:J movement of our limbs
provide the sensory fow1dation; m u eyes and visual system d o {see C hapters 2
Jn space.
through 8). The vestibula r system also helps maintain blood flow to the brain
via contributions to cardiac 01eart) regulation but is n ot foundational the re acttve sensing Sensln:J that
Inc ludes sett-generated probing of the
ei ther; sornatosensation is (see Chapter 13). environment. Besides cur sense o f
Also, as will be discussed in d etail later in this chapter, our sense of equi- equllibr1um, other active human senses
librium';is:'1at::tivt·, 1:ot By th.isr we mean that o ur e quilibrium sense Inc lude vision and touoh. Anlm1" active
(Figure 12.2) combines infonn<'.'ti6n flowing fro m our brain to our muscles sensing Includes the use of echoeo by
whales and bats, the use o f electrlcal
with information flovving inward to the brain from various sensory syste1ns,
elgnals by some l sh, and the use of
especially the kinesthetic, visual, and ,·estibular S)'"Stems. For example,signals whlskerafantennae by fish. Insects,
that tell o ur muscles to rotate our head provide information about h ead rota- and nocturnal rodents.
tion jus t a s vestibular s ignals d o. lnfornrntion fro m these various sou rces is
efferent commmds Information
combined to help improve our equilibrium. More generally, active sensing flowing a...ttward from the central ner-
balances information derived from efferent commands flowing outward from vous system to the perlphely. A com -
the brain to the periphery (e.g,, to muscles) wi th information from vari ous mon example is motor commands that
afferent signals fl owi ng from sensors imvard to the brain. regulate muscle contraction. The copy
In s ummary, the vestibular system contribu tes to our sense of e quilibrium, of suc h motor commands Is often
c alled an ccpy.
which is composed of m any fund amental reflexes a nd perceptual m oda li-
ties (Figm e 12.2), but the \'estibular system d oes n ot excl usively provide the afferent signals Information newing
Inward to the central navous system
sensory foundation for any perceptua l rnodality. Nonetheless, the of
from sensors In the periphery. Passr1e
the vestibular syste m' s contributions is pretty 3mazing (some might say it sensing would rely excluslvely on such
pro\.ides a" dizzying" array of contributions). When combined, these various sensory lnfbw, providing a traditional
percephla! and reflexive contributions are referred to as om sense of equilibrium view of sensation.

352 CHAPTER 12
Eye rotation
Visual sb.bility Balance Autonomic Spatial orientation
FIGURE 12.2 Our &quitlbrlum S9nS9 is com-

posed of multlple refla<es and multip le p9t"CGp-
tual m cdalltles that b eg in \f\llth the vestibular
organs In the Inner ear. EquHlbrium rQflQXes
ind ude vestibular contributbns to compgisa-
tory 9Y9 m ov&mQnts, callsd wstibulo-orular
reflexes. that rot ate our eygs to hgp ccmpen-
sate for head motion, b alance reflexes that help
u s stand, and 9V9n some autonomic reflQXQS
that help maintain bbod flow to our brain when
W9 rls& to a standing position. Spa.tlal orientation
includes perception of rotation, perception of
translatlon, and perception o f tilt, \Nhlch arie all
•pract1C9d• on playgrounds wery day.
because these various contributions, outlined in Figure 12.2, involve a balance

angular motion Rotational motion o f influences and / or a balance of forces-inatclling definitions of eqr1;libr;rm1
like the rotation o f a spinning top or you might find in a dictionary.
swinging saloon doors that rotate back
ardforth.
llnecr motion Translatlonal motion
Modalities and Qualities of Spatial Orientation
like the predominant movemoot of a O ur perception o f spatial orientation includes three sensory modalities: the
train car or bobblehead d oll.
senses of angular motion, linear motion, and tilt. Why do we call these 11mo-
tilt To attain a sloped pooltkxl like d.allties," as though they were different senses, rather than calling them "quali-
that of the Leaning Tower of Ftsa. ties'' ? For exanlple, vision and h earing are different modalities, but we would
transcllce To convert from one .say that color and brightness are different qr1alities1 n ot different moda lities.
form of energy to another (e.g., from The key lies in the energy 1ransduced. Color and brightness a re different in-
light to neural electncal erergy, or Iran
mechanical energy to neural electrlcal terpretations o f the same energy (light}-hence, qualities. Seein g and hearing
energy). involve different types of energy-light and pressure waves, respectively. As
semicircular canal Any of three
with seein g and hearing, perceiving angular motion, linear motion, and tilt
toroidal tubes In the vestibular system requires tha t three different stiinuli-angular acceleration, linear acceleration,
that sense angular motlm. and gra\oity, respectively-be transduced.
angular acceleration The rate of
c hange of angular velocity. Mathemati- Sensing Angular Motion, Linear Motion, and Tiit
c ally, the Integral of angular aocelera- These three s timulatio n energies are sensed by two types of vestibular sense
tlcn Is angular velocity, and the Integ ral organs: the semicircular canals and the otoHth organs. TI1e semicircular ca-
of angular velocity Is angular dlsptaoo- nals angular acceleration, which is a change in angular velocity; this
ment. Angular aoceleratlcn, ang<Aw
velocity, and angular dlsplaoement
signal makes a predominant contribution to our sense of angular m otion . To
all mathematically represent angU!ar experience your s ense of angular motion, simply close yotu eyes and rotate
motion. your h ead from side to side as if to say "n o." Because of contributions from

your vestibular system , you should a perception of angular otollth organ Either of two moohan-
ity tha t ro ug hly matc hes the true an gular velocity of yo ur head . lcal structures (utrlcle and sacculej In
The otolith orQans tmns duce both linear acceleration, which is a change the vestibular system that sense both
linear acceleration and gravity.
of linea r velocity, a nd gravity. The o tolith orga n s provid e a pred omina nt
contrib uti on to your sense of head tilt a nd a predorninant contributi on to your linear acceeratlon The rate of
sense of linear m o tio n , w hich is also referred to as your sense o f translation .
change of linear vekxlty. Mathemati-
cally, the Integral of linear acceleration
To experience your sense of tilt, simply pitch ymu he ad fonvard as if to 9'8.Y Is linear veloolty, ard the Integral of
"yes" and hold it there for severa l seconds; the n pitch your h ead backward Hnear velocity Is linear displacement,
and hold it th ere. You should ex'Pe rience a percepti on of tilt. Relative ly which Is also referred to as 'transla-
pure linear motion is more difficult to adl1eve but the experience of tion." Linear acceleration, linear veloc-
lty, and linear displacement all math-
riding in a car, trnin, or bus provides an example . Try the fo llo¥11i.ng when you ematically represent linear motion.
are a passenger in a car. 'With your eyes closed/ pay a ttention to your sense of
gravity A force that attracts a body
mo tion as the driver backs the car out of the garage, brings the car to a s top /
toward the center of the EMl1.
and then begins to accelerate forward . Jnitially, you should perceive backward
trans la tion (backward lin ear moti on) . You s hould also perceive the cessation sense of angulcr motion The per-
ceptual modaJlty that senses rotation.
of transla tion as the car comes to a s top, an d then forward translation as the
car accelera tes. sense of linear motion The
perc€!)tual tl1at senses
T\'\>'o different ty p es of sense organs-the semicirc ular c.:mals and the otolith
translation.
organs-establish a t leas t hvo sen sory m odalities. But if we have only two
types of sense organ s, why do we say the re are three m od(1lities? As noted sense of tilt The perceptual modal -
ity that senses l1ead Inclination with
<l iready, the otolith organ s transd u ce both gravity a nd linear accelern ti on . respect to gravity.
Perception of tilt results from the brain's estima te of orientation with respect
amplitude The size (increase or
to gravity, a nd perception o f lin ear mo tion resul ts from the brain's estimate decrease) of a head mcNement (e.g.,
of linea r acceleration. Pe rsonal experie n ce s u ggests that tilt perception is angular velocity, linear acceleration,
fw1d am enmlly different from tran.s bti on perception; these do not seem to be
different sensory qualities like color mid brightness. direction The line one moves along
'Nhy is this so? This key question leads to the hmda mental ration(11e for cr faces, with referaice to the point cr
two modalities that a rise from otolith signals. C lassical physics teaches that re;ilm one Is moving tCMtard or facing.
gravity and lineai· acceleration are distinct from one an other. Though Eins tein
hypothesized that g ravity and lin ear acceJe ration have e<p..riva le nt effects, he
d id not assert that they were the same. h1 fact, the brain d oes its best to separa te
the sign als from the otolith organ s into s ignals representative of gravity and
signals representative of linea r acceleration. (See Web Essay 12.1: Gravity
versus Unear Acceleration for tnore on th is equiva lence of grnvity and linear
acceleration.) ll1erefore1 '""'e assert three interacting sensory modalities-.." sense
of angular motion/a sense ot llnear motion, and a sense of tilt-p aralleling
the three different sou rces of stimulation energy- angu lar acceleration/ linear
acceleration, a nd gravi ty (Gu ed ry, 1974; Young, 1984).
Basic Qualities of Spatial Orientation: Amplitude and Direction

Each of our three spatial orientation modalities incl udes two qua lities:
tude and direction. As an example of a mplitude, the speed of our p e rceived
motion can be large (as in a jet ju.st before takeoff) or s mall (as for a baby craw l-
ing). As an example of direction, perceived linear mo tion might be forward,
up, o r to the left.
AMPLITUDE For linea r rnotion we ca n pe rceive tran sla tion h aving high ve-
loci ty (think o f a jet jus t before takeoff) or low veloci ty (think of a ca r inching
foru.•ard in a traffic j.9m). Similarly, we can perceive rotational velocity wi th high
ampli tude (think of vigorously shaking your head) or low a.m plitude (think of
the slow rotation of minute 0 1· h our hands on a clock). tilt amp litude
is also imp ortant. Ti lt amplitudes can be s mall (as w hen yo u gen tly n od your
head ) or laTge (as when you lie do\o\.'T\ or h<lng upside d own ).

354 CHAPTER 12
82
FIGURE 12.3 Movement of the head can b9 described in terms of a slmpe fixed
coordinate system: the x-axis atways points bward rglattve to the curr€f1t orienta-
tion of the head, they-axis always points out the left ear, and the z-axls always
pclnts out the top of the head. As part /lustrates, this coordinate systwn always
moves with the head.
DIRECTION To help clearly classify dlrection, we first define a simple Car-

tesian coordinate system that moves with the head (Figure 12.3). Since the
physical space that ""-e move in is three-dimensional, we need three axes. In
our defined system, the x-axis always points forward, they-axis always points
out the left ear, and the z-axis always points out the top of the head.
Tiuee directions define our sense of angular motion. That is, the head can
rotate in three independent ways, which can be represented (1) with a roll
angular velocity (Figure 12.4a); (2) with a pitch angular velocity, as when you
nod "yes" {FllJJre 12.4b); or (3) with a yaw angular velocity, as when you shake
your head 'jno" (Figure 12.4c). These three angular motions can be combined
to represent any three-dimensional head rotation.
(a) Rollo Rotation (b) Pitch: Rotation (c) Yaw: Rotation

around x-axis around y-axis around z-axi&
y
-r f
FIGURE 12.4 Rotating bodiQS can tnOV9 in threei directions. The head can turn with
a purg roll vQlocity around tM x-axis {a), a pure pitch wlodty around the y-axis p), or
a pure yaw velccity around 1:t'l9 z-axls {c).

THE VESTIBULAR SYSTEM AND OUR SENSE OF EQUILIBRIUM 355
(a) Positive x-axis (b) Po_sitive y-ilXis (c) Positive z-axi:s

translation translation translation
l=IGURE 12.5 Translating t::odigscan m OVQ in thrgg dlr9ctions. Th9 h98d c an trans-
latg forward and backward along the x- axis left and right abng the y-axis {b), or
up and down along th& z-axls (c).
There are also three directions for o ur sense of linear m o tion . Imagine (1)
stepping forward or backward along the x-axis (Rgure 12.5a), (2) slidin g fro m
right to left along the y-axis (Figure 12.Sb), and (3) translating up or down
along the z-axis (Figure 12.5c), which can also be combined to represent any
three-dime nsional linear m o tio n .
Finally,. each orientation has two tilt directions. For example, when you
are uprigh t, you might experience a pitch tilt forward or backward (Figure
12.6a). Or you might experience a roll tilt to the left or right (Rgure 12.61>).
What happened to the third dimension? The third rotation direction would be
a yaw rotation, but this would not yield a change in the tilt o f the head with
respect to gravity (see Figure 12.4c). When head rotations align with gravity,
there is n o d1ange in head tilt So there are three directions for a ng ular velocity
and translation, but only two directions for tilt.
(a) P•ch tilt (b) Roll tilt
F='IGURE 12.6 Tilting bodies can move ln tV'JO dlr9Ctlons. W'lan you are lnltlaly
upright with respect to gravity, there are two di"ectlons of tut. You can pitch-tilt for·
ward or backward (a), or you c an roll-tilt to the left or right (b).

356 CHAPTER 12
(a)
(b)
'\
1
Vestibulo-
cochlear
(VIIOnerve
FIG URE 12.7 (a) The vestibular

o rg..."10s occupy a membranous, fluid-
fllted sac that Is in a cavity in the tem- The Mammalian Vestibular System
poral bonGo, nQaf the cochl9a, and is
In this sec tion we look at w here and how rnotion signals are tr.. by
the nonhearing part of the Inner ear. (b)
The vestibular agans consist of three
the vestibular org ans. The vestibu lar organs are about the size of a large pea
semicircular c anals (antGrlor . postsrior, and c.Jn be found in the i1me r eM right nex t to the cochlea {Figure 12.7a). 1hey
and horlzcntaO arid twoo.tdltb organs respond prim.._uiJy to head rnotion-both linea r and angular-and hea d tilt
and sacCtJ!el Ol1 each side o f the \.-vi th respect to gravity. E...1ch inn.er ear h as one vestibular laby rinth, and each
head . The wstlbular organs are lnner- vestibular labyrinth includes five sense organs (Figure 12.7b): three semicir-
vattild by the vestibular nerve, which
jci ns the cochlear r1Qf'YQ to form cranial cular canals th.Jt sense rota tional m otion, and two o toHth orga ns (utricle and
nerve VIII, caned the vestltx..llocochlear saccule) tht"it sense gravity and linear acceleration. (See Web Activity 12.1: A
nelfve {ooe Flgu m 1.20 in Chapt€1r 1). Guided Tour of the Vestibular System .)
Note that neither the o tolith organs n or the semi circular ca nals resp ond
velocity The speed and dlrectlm
to constant velocity. Rather, they respond to chn11ges in velocity-called
In which moves. Math- celeration. The sensitivity o f the vestibular system to acceleration- an gula r
ematlcalty, va oclty ts the Integral of acceleration for the semicircular canals and linear acceleration for the otolith
acceleration. In words , linear va cclty organs-demonstra tes that the ves tibular system is principally sensi tive to
Is distance divided by time to traverse c1iai1ges in m otion. Constant motion, whether angular or linear, doe.s not res ult
that dlstanoe: angular vsoclty Is rota-
in vestibular signals that directly indicate m otion.
tlon angle dMded by time to traveise
that angle. As you will see again later, the oto lith o rgans transd.uce both linea r ac-
celerati on and gravity into a sin gle n eural sign.al sent to the brain. In fact,
acceleration A change In vsoclty.
Mathematically. acooeratbn Is the as Einstein asserted , no device can te ll the difference between gravity and
derivative of velocity. lnwords, llnear linear acceleration. Separating the o tolith measurement of g ravity and linea r
acoeteratlon indicates a change In lin- accelera tion into an estimate of gravity a nd an estima te of linear acceleration
ear vaoclt)O angular aooeleratlon Indi- is n ot easy and rnus t be important, since the brain expend s energy and effort
cates a d"lange In angular veloclty. to do so (Angelaki et a l., 1999; Merfeld, Zupan, a nd Pe terka, 1999). (See Web
hair cell Any c ell that has stereocllia Essay 12.2: Canal-Otollth Integration for m ore on how the brain dis tinguishes
for transduclng mechanical rnove- gravity from linear acceleration_)
rnent Jn the inner ear Into neural act1v1ty
sent to the brain: some hair OOls also
recave Inputs from the brain.
Hair Cells: Mechanical Transducers
Hair cells, which you read ab out in C hapter 9 when you began to le an1 about
mechanoreceptor A sensory
rea;ptor that responds to mechanl- h ea ring (see Fi gure 9.9), act as mechanoreceptors in each of the five vestibular
caJ stimulation (pressure. vlbraticn er org ans . Head m otion causes hair cell s tereodlia to deflect. Stereocilia deflection
movement). causes a change in the hair cell which alters neurotransmitter release,

(a) FIGURE 12.8 Hair oell ri96ponses. \i\tlen

not stimulated, the hair cel ls have a negative
voltage and release neurotransmitter at a con-
s tant rata, evoking a constant rats of action
potentials In the afferent nerves. W hen hair cell
bundles bend toward the kinoci lium and tallest
s tereocilia, the hair oolts bQcorne depolarizOO
and m ore neurotransmitter is reloosed (exci-
tation): vvhen they bend awaof, th0)' b9corn9
h)iperpolarized and les;s transmitter is rel9Med
Onhibltion). (b) Th is c ross-s.actlonaJ vliew of a h air
Nerve Resting Increased Oecrai;sed ooU array shows the direction o f dQflec tlon that
unpulses action action causes depolarlzatbn. tc) Top view of a hair ooll
"Afferent potential ra te patential rate array. The blac k arrows indicate the excitatory
directio n o f hair c 911 deflection.
Excitation Inhibition
whichr in hunr evokes action potentials in those vestibular-nerve fibers that

have one or rn ore syn.apses on the hair cell. These affe rent neurons carry these
action potentials to the brain
Le t's consi de r vestibular hair cell resp on ses in a bit more detail. In the
absence of stimulation, the hair ceJls have a negative voltage and release neu-
rotransmitter a t a constant rate1 evoking a cons tant rate of action p otentials in
the afferent n eurons (Fig ure 1 2.8a). Changes in hair cell voltage---caJled the
receptor potential-are p rop ortional to the bending of the hair cell bw1d.les
and control the ra te a t w h ich hair cells release netuotran.sm.itter to the afferent
neurons. 'When a hai r cell bends toward the tallest stereocilia (Fig ures 12.8b,c),
the hair cell voltage becomes less negative. TIUs voltage ch ange is also called a
d epolaTization bec:a use the hah cell becomes less polarized than the n ega tive
resting potential (see Figure 12.&). The h air cell d epolariza tion increases the
release of ne u rotransmitter1 causin g an. increase in the action pote ntial rate
(caUedexdta tion ). On the other hand, if the hair cell is bent nway frorn the talles t
ste reodlia, the cell potential becomes more negative (hyperpolarizes), causing a receptor potential A change In
voltage across the membrane of a
d ecrease in the !'eleaseof neurotransll"1itter and a decrease in the action potential sensory rec€!)tor cell On the vestibu-
rate (c<tlled inhibition). In Slffi1.mary, the rate of action p otentials t.rans nUtted by lar system , a hair cell) in response to
afferen t ne u rons increases or d ecreases followin g the hair re.II receptor p otential. stlmulatlon.

358 CHAPTER 12
FURTHER DISCUSSION of the receptor potential of hair cells can be

found in Chapter Q on pages 270-272.
Thinking back to the fact that amplitude is one quality of our vestibular
sense, we can begin to see how amplitude is encoded, since the rate of action
potentials is proportional to the receptor potential, w hid1 in h1rn is propor-
tional to the amount of hair cell de flection, w h ich in hlm is. proportiona l to
the a mpHtude of the m otion .
The fact th.a t the hair cells respond opposite ly for de flecti ons in oppo--
site directi ons (see Figure 12.8n) is also cru cia l for the codin g of vestibular
s timuli. For example, as we'11 d iscu ss in de tail in the following section, a yaw
rotation to the le ft w ill increase the hair cell receptor poten tia l fo r a hair cell
loca ted in the horizontal canal of the left ear, and a yaw rota tion to the right
will decrease the receptor potential for that same hair cell. A similar general
principle applies for the o tolith organs: acceleration in one direction increases
the receptor potentials of some hair cells, while acceleration in the opposi te
d irection decreases those receptor potentials.
Semicircular Canals
Each inner ear h as three se micircular canals-horizontal (or la teral), anterior
(or s uperior)1 and posterior (Figure 12.9) (the anterior and posterior canals are
C ilia
Ne Ne FIGURE 12.9 canals. The inner ear has three roughly cbugh·
fibers
nut·shaped semlcircular canals. Each canal hos a sW911ing miar the v,;ietibule
called an ampuUa, where angular motion is transducad by hair ciells in th9 crlsta
Wthin each crista, all o f the hair ci911s are aligned.

some times called the vertical canals). These cana ls are roughly orthogonal to ampulla An expansla1 of eadi
one a n other. The name semicirrnfnr canal loosely reflects the gross anatom.y of sernlci rc ular-canal duct that inciudes
this strudurer which has the circular sh ape o f a n incomplete toroid or d ough- that canal's cupula, crlsta. and haJr
cells, where transduction ccrurs.
nut. about tluee--fo urths of a toroid is fom1ed by a bony tube ap-
proximate ly 15 milUmeters (mm) lon gr with a cross section about 1.5 mm in crista M y o f thl spoolallzed detec-
d iam eter. This space----c.:1 lled the osseous (mea ning "bonyn) canal because it tors of angular rnotloo loca:ted In each
sernlcircular canal In a swelling called
is a canal carved out of the m..'lstoid bone- is colored tan in the figure and is treampulla.
fi lled with a fluid called pe rilyrnph. 1l'le remaining length of the toroid passes
through the vestibule, w hich is opened up in the figure to a llow a glimpse
inside. A second1 smaller toroid, appearing bl uish in the figure 1 is found inside
the larger toroid. This smaller toroid is formed by a mem brane filled wi th a
fluid called endoly mph, and in cross section it has a diame ter of about 0.3 nun,
w hich is just a little thicker than a very thick human hair.
The cross section for each ccmal swells substa ntially n ear where the canals
join the vestibule. Each swelling is ca lled an ampulla (plura l nmpulfrie) (see
Figure 12.9). Within theendolymph space of eachampulla, the angular-motion
d etectors me assembled into a sen.sory epithelium called the crlsta (p lural
cristae). Each crista consists o f a small ridge, '"·hich has an epitheJi um made
up of 7000 hair cells and is innervated by about 4000 nerve fibers. The
kinocilium s tereocilia (singular stereocilium) of ead1 hair cell p roject into
a jellylike cupu1a (plural cuprdae) that forms an e lasti c dam extending to the
opposite '"''all of the \-Vi th endoly1nph on both sid es of the dam .
When the head the inertia. of the endolymph causes it to lag behind
the motion of the head (Breuer, 1874; Brown, 1874; Mach, 1875/ 2001), leading
to deflection of the cupula and thereby to tiny defl ections of the s tereocilia in
the crista. As mentioned ea rlier, s uch de flections evoke changes in the
hair ce Us, wh ich in h.tm cause changes in the fil'ing rate of afferent neurons
(see Figure 12.8a). For each individual semicircular canal, all of the h(1ir cells
are aligned (see Figure 12.9). Tints, rotations in one direction yield i.ncr&1ses
in the receptor p otential of all ha.ir cells in that semicircular canal, as well as
concomitant increases in the action potential rate for all ne urons that inn erva te
canal. Rotations in the opposite dfrection yie ld d ecrec1ses
in the hair ce.U receptor poten als and concomitant decreases in the ra te of
action potentials.
The three semicircular can.:-US are maximally sensitive to rotations in differ-
ent planes, thus yielding direction coding for rotation. Specifically, each
canal is rnnx imally sensitive to rotations about the axis perpendicular to it,
and insensitive to rotations a.bout axes that fall in the
plane of that canal (Fig ure 12.10). 11-link of each canal }>.1a:ximum sensitivity
of horizontal canal
as a \vheel that can s pin about the axle. The canal is lnsen.siti\-e
maximall y sensi tive to rotations that circle the wheel 1
axle and insensitive to rotations in o ther planes.
HOW AMPLITUDE IS CODED IN THE SEMICIRCULAR

CANALS In the absence of any rota tionf many af-
ferent neurons from the semicircular canals respond
FIGURE 12. 10 Each semicircular canal is maximalty

SQflsitfve to rotations pgrpgndicular to the canal plan9.
Thinking of each semicircular canal as a 'Wheel. we can
Sl9Q that thi9 canals are maxlmalty sensitive to rotations
that align with the rotation axis. (shown in gr.een) and W"Q rotatiru1 axis
insensitive t o rotations. that fall In the plane of the sen1!cir- of horizontal
cular canal (e.g ., puipl.e and blue).

360 CHAPTER 12
FIGURE 12.11 Th9sernicircular

canals functbn in pairs that have a
push-pull r91ationship. (a)
s tirnulation of the hcrlzontaJ sernl -
cirrular canals as the head turns
to the tight (a yaw head rotation a<>
shown in F(lure 12.4c). This yBN
(a)
- Tu min& motion of he."l(i
Depolari.z.:1tion
ofh.tircells
rotation produC8s relative m ovan.ent

o f the erlcblyrnph in the horizontal
semicircular canals on both sid9S
o f the head. The movGoment of the
fluid bends the hair cell bundles
toward thei tallest st9r90Cilia on the
right side. which defXllarizfis thsse
hair cells and increases the rat9 o f
action potentials for neurons from
the right side. The hair cell bundles
rnow away from the tallest stereo-
c illa en the left, hyperpciarizing thr:i
hair 09lls and di9creasing the rate of
adiat potentials for left-siOO neuro11s.
This responS9 -an incr9aS9 on one
side couplOO w ith a decrease on the
other side-is ofto3n caJled a "push-
pull The two horizontal
canals fotm one push-p ull paJr. (b)
The axis o f maximum sensiti\'ity for
the right horizontal canal. (c) The right
anterior canal and the left posterior
canal form another pair. Note that the
rnaxirnum-sensltMty axes for the right
anterior and left JXlS1&r1or canals arQ
paTallB. This m eans that they maxj-
malty respond to rotations around
the same axis. The push-pull nature
o f this canal pair is aconsequencei of
the opposit9 directions of thQr max-
hnum-sensitMty rotations. as sho'!hfl
by the green arro'YVS.
Ylr'i th a nearly constant rate of action potentials (see Figure 12.8n); canal afferent
firing rntes at rest average about 100 action potentials (1'spikes") per second
(Goldberg and Fernandez, 1971), a rate that is hi gh relative to spontaneous
rates for nerve fibers for other sensory system s. For comparison, recall from
Chapter 2 that retinal ganglion cells fire s pontaneously a t a rate of abo ut one
spike per second in the dark.
The relatively high spontaneous firing rate of vestibular afferent n eurons
allows these neurons to d ecrease the firing rate for rotations in one direction
and increase th e firing rate for rotations in the opposi te direction. In
the semicircular cana ls are organized as hmctional pairs in what is ca lled a
push-pull a rra nge ment. The hvo horizontal canals-one on the right side of
the and one on the left-lie roughly in the sa_ir1e plane and form one
of the three functional can a l pairs (Figure 12.11 a,b} (C urthoys 1 Blanks, find
Markham, 1977; V. J. \Vilson and f\·1e lvill Jones, 1979). The h orizontal-cana l
afferent neurons on the right a ll increase their firing rate for yaw head tu.ms
to the right (shaking ''no'' ), and those on the left decrease their firing rnte.
For head turns to the left, the pattern is reverse d, with left-cana l afferent
neurons increasing their firing ra te and right-canal ne urons decreasing

their firing rate . The anteri o r and p osterior canals are minimally sensitive
to these yaw ro tatio ns.
In to the h orizo ntal-ca na l a rrangement, th e m irror synunetry of
the senUci rcular canals in the left and right ears yields hmctional pairs th a t
involve different vertical canals (Fig ure 12.11 c); the maxim um-sens itivity axis
of the anterior semicircular m nal on one sid e roughly parallels the maxi mum-
sens itivity axis of the p ooterio r semicircular cana l on the opposite side. So the
right an terior a nd left posterior cana ls fom1 one canal pair, as d o the left ante rior
and right posterior canals. These canal pairs work in a push-pull rnanner like
that previous ly d escribed for the horizontal canals.
The change in the firing rate is larger fo r large ch anges in th e angu lar
velocity of tlle head than for small d1anges (Goldberg a nd Fe rnandez, 1971).
Since the cha n ge in the firing rate is prop ortional to ang ular velocity, we can
think of the canals as some\•vhat similar to the speed ometer in a ca r, wi th the
cha n ge in neura l activity propo rtio nal to the angular \o-elocity of the head. As
a specific example, if the afferen t ne urons fro m a h orizontal canal suddenly
cha nge the ir firing rate, we know that the rotation includes a change in the
velocity compo nent a ligne d wi th the sens itive axis of that h o rizon tal canal.
The same is tru e for the o ther canals.
HOW DIRECTION IS CODED IN THE SEMICIRCULAR CANALS As previOLLsly

discussed , the three semi circular canals are maxirnally sensitive to rotati ons in oscillatory Refernng to back-and-
different planes; the result is d irection codin g of head rotations. Specifically, forth movement that has a constant
the hea d cn. n rotate about any arbitrary rotation axis made up of roll, pitd1, and rhythm.
yaw ro t:ltional velocity components. Each canal trans duces the component of
head velocity perpendicular to its plane. TI1e brain then combines these signals
to sense the rotation direction of the head movement.
SEMICIRCULAR-CANAL DYNAMICS If you sudde nly

begin to rota te a t a consta nt velocity, the semici rcular
canals sensitive to that ro tation "'rill by causing a (fl) Stimulus
sudden change in afferent ne ural activ ity. But as the rota-
tion continues at a conshmt velocity, the afferent neura.l
activity ""ill decay back to near zero after about 15seconds_ Aa:eleration Constant Deceleration
If you then suddenly d ecelerate to a s top, the canals w ill
show a large response in the opp osite direction (Figure
\ ve locity - . ./
12.12). 111e afferent neural activity \o•lil l then decay with

a time com se similar to that during the constant-velocity
rotation (Goldbe rg and Femand ez, 1971).
W hat happens for more natural head rotati ons-- (b) Neuron activity
like tbe oscillatory back-and-for th m ovem ent when
you s hake your hea d "no"? (U y·ou d on' t have a solid 120 336 Amma Appa
FIGU RE 12. 12 R99p0nse o f a semicirculru-·canal neuron

to constant-v91ocity rotatio n. (a) The stimulus is a rotation
that first accelerates t o a constant ang ular vtilocity, then
maintains tha.t vG!ocity, and th9n d909l0fat9s the head to a
stop. (b} During the initial the c upula defltcis,
causing the neuron actMty to increase. During the constant
angular velocity, th9 cupula returns to its nondeflecfQd
position, oo the. neuron activity returns t o the bas-='ine ratf:i
after about 15 seconds of constant-velocity rotation. During
deceleration, the c upllla Is defl9cted in the opposite direc-
tion, causing a transient decrease in the firing rate. (After
40 80 1'1l
Gok:H:>9rg and Femardez, 1Q7 1.) Ttme (s)

362 CHAPTER 12
sinusoidal Referring to any oscil- understanding of frequency analysis, now may be a good time to review the
lation, such as a sound wave or "Fourier Analysis" section in Chapter I. See also Web AcUVlty 12.2: Slnusoldlal
rotational motlcn, whose waveform Is Motion.) F1gure 12.1 3a shO\vs a sinusoidal m otion trajectory at a frequency of
that of a sine curve. The penoo of a
slnusddal oscillation Is the time that It
takes for one full back-and-forth cycle
(a)
of the motion to occur. The frequarcy
of a elnusoldal oscillation Is defined as +SO
the nurner,; 1 drJkJed by the period. +60
:r
_g -20
.. -40
-bO
"" -80
2J 40 60 80 100 l:ll 140 160

Time(s)
(b)
180
23 336 m a f. pa
,"'
160
140
120
]:
100
80 I
..
-ii
Q
60
40
20
0
0 2J 40 60 80 100 !)'.) 140 160
'Time (s)
(<)
"'
1.3
!' 0.9
• 0.
05
H 0.1
d 0.01 0.1
FM:iuency (Hz)
FIG URE 12. 13 Sinusoidal m otion trajectories. Stimulus (a) and responsei ft>) for a
si11gh3 semK;;ircutar-canal afferent n.euror1 at 0.05 Hz. Note that the neural fir1ng rate
inc r98S'9s and decreasEas nearly in tandem with the oscillating stimulus. (c) When the
sensitMty calculation is repeated at differi:int stimulus frequ€<'1Cies, it can be seen that
the sensitivity of the neiuron changes v....ith frequency. For this seornicircular-can.al neu-
ron, the response SGnsiti\.'ity averaged about 1.3 spikes/second PQr degrQQ/s.econd
for frequenc ies above 0. 1 Hz. The response sensitivity decrEia.Ses substantiaJly for fre-
quenci91S bebw about 0. 1 Hz. (Data from FemandBZ and Goldberg. 1Q71.}

0.05 he rtz repeatin g back-and-forth motion tha t takes 20 secon ds to Fourier analysis A mathematical
cmnple te (like slm-v1y s hakin g your hea d ). Fig ure 12.13b shows the oscillatory procedure by which any signal -In ti>s
neural response-in an afferent neuron evoke..-l by that sinusoid a l m otion . The case motion trajectories as a func-
tion of time-can be separated Into
firing rate increases and d ecreases as the angular veloci ty of th e h ead increases component sine wavoo at different fre-
or decreases; that is, the ch a nge in firing rate h as the same frequency as the quencies. Combining these sine wm.res
head velocity. \Ve can calculate fill estitn.;ite of afferent neuron sensitivity by will reproduce the original rnotion
taking the peak-to-peak a rnplih.lde of the change in the firing rate (from Figure trajectay.
12.l3b) and dividing it by the peak-t<>-peak velocity (from Figure 12.13a). The u1ricle Ona of thl two otolith
inaxinnun firing rate is about 170 s pikes/second; the minimum, abou t 10 crgans. A sacllke stn..ct ure that con-
spikes per second. So the pea k-to-pea k a mplitude o f the change in the firing tains the utricular macula. Also called
utrl:;ul.Js.
rate is about 160 spikes/second. The peak-to-peak tl mp litude of the cha nge in
motion is 160 degrees /second (80 degrees/second to the left and 80 degrees / saccule One of the otolith
second to the righ t). 1l1e result is a sensitivi ty of about 1 spike/second per crgans. Asacllke structure that con-
tains the sacx::ular macula AJso called
d egree/second of motion. Experimentally, this process is o ften repeated a t sacculus.
several different head rota tion frequencies, yielding a number representing
macula Any of the specialized
the neural sensitivity a t ead1 freque ncy tested. detectcrs of linear acceleration and
Semicircular canals are n ot equally sensitive to all frequencies of rotation. gra'lity found In each otollt11 organ.
Figure 12.1 3c s hows a plot of response sensitivity as frequency is vmied for
a typical neuron innervating a semicircular canal; this plot indicates how the
osci.1.la tory-canal afferent firing rate changes ,..,.·ith head rotation frequency. The
data show that theampLitudeof the canal afferent neuron's response is nearly
constant for rota tion frequen cies above abou t 0.1 Hz. But when the frequency
is less than about 0.1 Hz, the neuron responds less and less. For examp le, the
response sensitivity ata freq uency of 0.02 Hz (one cycle in 50 seconds) is on ly
about0.5 spike/ second per degree/ .second-more than a 50% reduc tion from
that observed a t 1 Hz (one cycle in one second).
WHY SINE WAVE MOTIONS? Now that we have d escribed cana l afferen t
responses to .sinusoid al motion in some detail, you may wonder w hy an yone
would care. We rnrely motion in the real world;
we experience complex thai vary w1tH tlme. \ Vh.it do responses to
sinusoidal motion a t different frequencies tell us about how we.sense motion?
One a nswer is th:l.t, al th ough ;'pure" sine motions may be ra.re in the
real world, patterns of oscilfatory motions thilt have a predominant freq uency
are quite common: think again of .shaking your head to s,i.y "no" or nod ding
your head to say "yes." More generally, Fourier analysis tells us that any
coroplex: rnotion rn n be broken down into some number of single-frequen cy
components. Therefore, if we know responses to sin gle frequencies, we kn ow
a good deal about responses to more complex mo tion . lf this sounds familiar,
that's because the same argumen t was made in Chapter 3 (.see "W h y Sine
Wave Gratings?'' on page 61), as well as in Chapter 9 w hen we talked about
hearing (see /(Sine Waves and Comp lex Sotmds" on page 264).
Otolith Organs
.l.\s we mentioned earlier, the sensing of gravity and linear accelera ti on relie.s; on
two struch ues in each ear called the otolith organs: the utrlcle (or utriculus) .:md
the saccule (or sacculus). Each of these organs consists of a small, oval-shaped,
fluid-filled sac th..i.t is about3 mm long in the longes t direction and includes an
area rn lled the macula (plural mnculae), \vhich is where actual sensory trans-
duction occurs (Figure 12 .1 4a). Ead1 human utricular macula contains about
30,(X)O hair cells; each sa.ccular rnacula, abo ut 16,0CKl The utricular and saccular
ma culae a re inneivated by about 4000 neurons each . Each macula is roughly
planar m1d is sensitive to shear forces- forces para llel to the macular
plane. Perpendicular forces have little influence on neural response. Small move-

364 CHAPTER 12
(a)
(b)
Otoconia
Otolith.ic
nwmbl'ane,
gelatinous:
layer
I-fair ce lls Afferent neurons
FIGURE 12. 14 The otolith organs. {a) Orientation o fthi9 utrk::ular and saccular rnac -
ulae in thg head. Thg sacculoo ore oriented more or less verticalty: the utriciQS, more
or less horizontally. The striola Is a structural landmark that dMdes each otolith organ.
The tallGSt stareocilia point tOYVard the strio la in the utric ular rnacula and from
it in the saccular m acula, as th.:ci arro'VY'S indk::ate. Because hair c ells depolarize if thfiy
dafl9Ct toward the tallest st9reocilia and hyperpolarize if they dr&fl9Ct from them,
the wrO'WS also show the movement direction that causes maxjmal neural excitation
at each location on the m aculae. Note that. given one utrlcle and one saccu le on
ead1 side o f the head, th€<e is a continuous representation of all directions for s,e.ns-
ing gravity and lineiar acoeleratic;n. (,p) Cross secticn of the rnacula of an otolith organ.
Hair bunctlQS project into a gelatinous lay4i!r. (c) Th9 otoconia sho wn in this scanning
el9Ctron mic rograph come from the utricular macula o f a cat. C rystats. are between
0.5and 10 mlcrornet9rS tong. (Part c from Lindeman. 1973.)
ment.s caused by parallel shear forces, due to gravity and / or linear acceleration,
deflect the hair cells and produce chan ges in the firin g rate of afferent neurons.
TI1e cilia o f the otolith hair cells enmsed in a gelatinous s h·uctme (Flgure
12.14b ) tha t contains calcium c ub onate crystals called otoconla (sin gular
otoconi11111) (Figure 12.14c). There are liternll v milli ons of these otoconia in the
otoconla Tlrry calcium carbonate utricle and saccu.le. The:se s mall ghajh otolith prg•m s their name: oto
stones in the ear that provide Inertial
mass for the otollth organs, enabllrq mea ns "ear/ and litlios m ea ns "stone " in Greek, so otolith tr.:ins lates as "ear
them to sen se gravity and linear s tone." The otoconia are d enser than the s urrounding fluid . Like any dense
acoe1€<aUon. object, they are pulled by both grav itational force and inertial force d ue to

linear accele ration . The resLLlting displaceme nt of the o toconia drags the
gelatinous layer, the reby m.oving the hair cell stereocilia, leading to ch anges
in the hair cell receptor potential, w hich in turn ca use changes in the rate of
action potentials in the afferent ne urons.
HOW AMPLITUDE IS CODED IN THE OTOLITH ORGANS As in the semi-

circular canals, one aspect of amplitud e coding in the otolith organs can be
foun d in the response of a single n euron or s ingle hair cell. Recall that (l)
hair cell recep tor p otentials increase when hair bundle tips m ove toward
th e largest stereocilia, (2) recepto r p ote ntia ls d ecrease for m ovements in the
opposite direction (see Figure 12.8), and (3) the direction of rotation is coded
by exci tation frorn a sernicircu1ar canal on on e side and inhibition from the
other s ide.
Similar m echanisrns are at work in each otolith
organ macula . Populations of !"t.:ilicells on ead1 rnacula
have their s tereocilla oriented in opposite directi ons (•)
(see Figure 12.1411). Bo th the utricle a nd the sa ccule
include a central band c..<J. lled the striola (plural striolae).
On opposite s ides o f the striola1 hair cells-are oriented Starttih End tih
in opposite directions. Re me mber tha t movem ent
towa rd the ta ll est s tereoci lia exci tes the hair cells 1
and m ovement in the opposi te direction inhibits the
hair cells. Since the n euronal response arises from
synapses to the hair cells1 tilts (or linear ::.Kcelerations)
in op posite directions cause opposite d)..."lnges in firing l:U
rate (Figure 12.1 5). So a tilt or an accelerrition that
maximally excites a hair cell and afferent ne uron on
one s ide of the s triola will maximally in hibit a hai r
cell and afferent neu ron on the opposite side.
Larger accelerations {or larger gravitational sheCtr
forces) move the otolith org;.ms' otoconia more . This
moveme nt1 in tum, leCtds to greater d eflection of the
hair cell bundles, which causes larger ch anges in the
hair cell receptor potentfals. The recep tor potential
evoked in a given hair cell is prop ortional to the com-
ponent of gravity or linear accele ration that is aligned 40 80 120 160 200
with the sensitive axis of tha t hair ceH. Larger d1anges T1.1ne (s)
in the hair cell receptor potential lead to larger dianges
in the rate of action po tentials sent to the brain via (I>)
afferent neurons.
Sta1ttilt End tilt
HOW DIRECTION IS CODED IN THE OTOUTH ORGANS
\.-
Direction coding in the otolith organs arises, in part,
from their anatomical orientation. The plane of the
utricular macula is horizontal; the plan e of the sac-
\ Constant tilt J
cular m acula1 vertical (see Figure 12.1411). The macu-
FIGURE 12.15 Activity of a vestib.1lar neuron Innervat-

ing the (an otolith organ). '3) A changQ in Nad tilt
is the stimulus (top). As sho VYfl in th.e histogram o f the
discharge rats (bottom), the neuron's activity inc r98.S9S in
respcrise to tilt in a particular direc tion. (b) ThQ same neu-
ron decreases its .octlvity in response to tilt in tt1e opposite
directio n. (,A.ftQr Goldberg and F9rnando9Z, 1976.) Tune(s)

366 CHAPTER 12
lae are exquisi te ly sensitive to gravity and linear accelern ti on in the plane of
the macu1a and insensitive to gravity and acceleration perpend icular to the
macula. Therefore, wi th the hea d near upright, the utricle \vill be sensitive
prin1arily to any Earth-horizontal linear accele ration, while the sacc ule will
be sens iti ve to vertical linear acceleration ..
Th e o ther component of direction cod ing a ri ses from va riati ons in the
orientati on o f the h air ce lls 1 ri9ht). Different ha ir cells
respond m axi mally to diffe re nt m o\·e ment ions, with the direction of
maxi mal sensi ti vity varyin g sys tematically across the plane of ead1 marula .
For exarn.ple, h air cel ls in one region of the utricular macula will be maximally
sensitive to forvvard-backward acceleration, while cells in another region will
be maximally .sensitive to side-to-5ide lin ear acceleration. In behveen, cells wil l
maxim.ally respond to linea r acceleration that has both forward-bllckward and
side-to-si de comp onents.
Spatia l Orientat ion Perception

Long after it was known that \'\>'e see with o ur eyes and hear \>1r-i th o ur ears, the
primary som ce for o ur perception of spa tial orien tation rem ain ed a mys tery.
In fuct, in the eigh teenth cenhlI)'i gross fluid shifts in the head were accepted
as an ex plan.a ti on for the source of otlI sense of spatial orien ta tion. O n ce it was
established that this sense degmd es w hen the vestibulu system. is dam aged, it
became dear that the vestibular system p rovides crud.11 information regard-
ing spatial orientation. For ex ample, w hen patients with vestibular loss are
rnoved in the dark, they have a mud1 m ore difficu lt time correctly perceiving
the ir motion than do peop le w ith normal vestibu lar function.
Today, three different techniques are frequently used to investigate s patial
orientation perception: thresholds, magnltude estima ti on, and matching. For
a threshold study, we ca n ask, \Vht' t is the minimum m o tio n (the threshold)
required for cor rectly perceiv ing the direction we are m oved? Note that this
is different from simply reporting whether we've m oved , sin ce vibration can
pro·dde a mo tion cue wi thout info rmin g .about the d irection of th e m o ti on .
The vestibular system tells us m ore th an w hether motion is p resent or n ot;
it actually informs us of the direction of m otion- fo r example, \'\>·hether we
moved to the left or to the right.
In a magnitude estimation stud )i subj ects might be asked to give verbal
reports of h ow much they tilt, rotate, or trans late, usin g ph ysica l mUts like
the num ber o f degrees they rota ted . Alterna tively, magnitude est im ation may
utilize arbitrary scaling. For example, subjects may be trained to rotate a knob
in proportion to their perceived velocity o r provide a verbal indicator of their
velocity on a scale of 1-10.
In a matching task, s ubj ects might be asked to align a visu al line w ith
perceived Earth-vertical (wlUd1 wny is "down''?). [n such a task, called the
"subjective visual vertical" the in vestigator could produce a vestibular
sti mulus by tilting the s ubject. That s ubject would be provided with a visible
line in otherwise dark s urrotmd.ings and with the ability to rota te the line to
the perceived Earth-vertical. Alternative!); haptic sensation, which you \vill
learn n10re about in Ompter l3r could be utilized ins tead of vision. For this
technique, s ubjects might be asked to use their sense of limb p osition to align
a bar tha t they h old- but cannot see-with perceived ver tical.
FURTHER DISCUSSION of similar matching in connection with the

sense o f touch
can be found in Olapter 13 on pages 417-418.

FIGURE 12. 16 The re d line on this graph s hows the

angular velocity of a person at rest, with gyes closed,
w ho was suddenly rotated at a coostant speecl for 60
seconds and then was abruptt>,r return9d to r.eet. Initially,
/Actu al
,·elocity th9 subject's estimate of angular velocity was accurate
(purpl9 line at time 0). but then the percaiwd vefoc·
ity dropped to near 0 after about 20 S€1Conds.
stopped. the subject perrOOved an abrupt w loclty In the
opposite directioo, which mimic k.9::1 the initial P9f'C9ption
"'SemicircuJ,1r but in the opposite diroctlon. For comparison , WQ shOIN
can al respon se a reprBSentatlon o f the canal rooponse in bll1e, slmllar
to that ::.tiown in Flgise 12. 12. No te thefast9r return to
for the canal response. (Aftef Young, 1984 .)
10 '1l 3IJ 40 50 70 I'll
Tune (s)
Rotation Perception
U yo u are spun o n a barstool in the dark a t a nearly co nstant velocity, you w ill
initially perceive an angula r velocity tha t is roughly the same as the actua l
rot.."ltion. H owever, if the constant-velocity rota tio n lasts more than a second
or two, you wlll p erceive tha t you are s lowing d own (Figure 12.16). If the
consta nt-ve locity rotation continues for more than 60 seconds o r so, you w ill
perceive that you are no longer rota ting.
This descrip tion may rem ind you of the way the response of the semicircular
canals d ecays during consta nt-velocity rotation (see Fig u re 12.12), a nd it is
another example of how the vestibular sys te m is a ttuned to changes in mo tion.
Interestingly, th ough, the time course of the perceprual d ecay is m ore gradual
than th.a t of the semicircular-canal s ignal sent to the br(1m. TIUs effect is so1ne-
times ca ll ed ;'velocity storage" because the perception of rotati on persists after
the a fferent signa l from. the semicircular c.:1 nals h as dissipated . The veloci ty
storage phenomen on is interesting and importan t because it shows that the
brain has improved on the incoming sensory informati on to yield a rotn tion
perception that-while far from perfect- is closer to the actual rotation than
if the pe rception s im p ly fo llowed th e time course of the semicircular-canal
afferent signal (Berto lini et al., 2011).
Later, if you are abruptly brought to a stop fo llowing ex tended rotation,
you ,.... ill perceive an an gular velocity opposite the one you experien ced w hile
rotatin g (see Figure 12.16). This rotation illusion is one that m any of us p layed
with as children wh en we would spin ourselves for a w hile and then s uddenly
stop spinning and try to stand or walk. (A nd som e amusement park jtmkies
still play with this illusion as you 'll see at the end of th is d1apterl) The diz-
ziness and Imbalance th at we experien ced when. we stopped rotating were
due to an illusion o f self-rotation caused by the se1nicircular-cana l response.
One can develop an intuitive tmders tanding of the reason fo r th is illusion
by considering the analogy of riding in a car. When you 're riding at a constant
velocity, you and the car are moving together. But when the car suddenly stops,
you are throvvn forv•.rru-d beca use you have momenhun and keep m ov ing even
thou gh the car has s topped. When you're rotating at a constant velocity, there
is little o r no hair cell deflec tion, because the endolymph and cupula are mov-
ing together. \ Vhen the rotation is sudd enly h alted, h owever, the cu.pula s tops
dizziness A commonly used lay
moving quickly but the endoly mph has m omenhun and tend s to keep rnovi.ng. term that rK:flspedfically ln:llcates any
The hair cells are therefore d eflected, the direction of the hai r cell resp onse form of pEfceti.red spatial dlsorlenta·
is opposite the one measured when the cons tant-velocity rotation bega n. tlon, with or wlt11out lnstablllty.
H ow sensitive we to rotation? Direction recognition threshold s for ya\.v Imbalance Lack of balance;
rot..1tion have been m easu red for rotational motion frequencies ranging from 0.05 urstead lness; rearly faHlrr;i over.

368 CHAPTER 12
FIGURE 12.17 Mean v elocity thresh-

old as a function of frequency for seven
subjects. Threshold wlocity was the
peak vo:iloclty a::::hieved during a single
cycle of slnusddal accel9t"ation at
which subjiects correctly recognized
the direction o f rno tbn most o f th.e
time. The plotted cu rv e shows a rnod'111
1
! 4
-2
Velocity
frt to the data. The inset shows the 31

angular acoel9ratbn, ang ular w loc-
ity, and angular displacement for a
single cycle of slm.JSOidaJ acceleration
j
at 0.5 1-tz. For the QXafOpte sho wn, the
peak magnituOO of the aci:geration is
1.57 degrees/s2, the p::iak velocity is 1 0.5 1.5
degreek., and the peak dispacerrient Is
1 degree. t;After Grabherr et al., 2008.)
0.02 005 0.1 0.2 0.5

Frequency (Hz)
Hz {one back-and-forth cycle of y m. . ·accele rntion in 20 second s) to 5 Hz (five

back-and-forth cycles of acceleration in l second ). For frequencies above
1 Hz {one back-and-forth accelera tion cycle in 1 second), direction recognition
thresholds are rou ghly constant; your head has to be m ovin g at a sp eed o f
just a little below 1 degree pel' second , w hich a t 5 Hz corresponds to a head
an gular di splacement o f just 0. 1 degree or ju.st 0.02 of a minute on a dockface!
C learly, we are very se ns itive to rotation. Por freq uencies below 0.5 Hz (one
back-and-forth oscilla tion in 2 seconds), thresholds increase wi th d ecreasin g
frequ ency (Ftgure 12.17). The important poin t here is to recognize that rotation
thresholds vary as the freq ue ncy of the angulM acceleration stimulus varies.
(Recall tl1at hearing tluesholds changed wi tl1 frequency too [see Figure 9.22]
tho ugh the cause of the c hanges is different for th e tvvo modalities.}
Translation Perception
When subjects a re p assively translated sh or t d istances while seated in a chair
in the dark and then asked, w hile s till seated in the chair, to u se a joystick to
actively move the di.air to reproduce the distance that they had been passively
h'anslated, they do so accurately. But even though n ot C\Sked to do so, they also
reprod uce the 7.'elocity of the passive-motion trajectory (Be rthoz et al., 1995).
The unrequested replication of velocity s uggests that the brain remembers
and replicates the velocity trajectory. E..1.rlier we sai d that the o to lith org.:'l n.s
tra nsduce linear acceleration, which is the change in linea r velocity. Therefore,
replication of the velocity trajectory m eans tha t the brain also see ms to math-
ematically Integrate the acceleration signal provided by the otolith organs to
y ield a p erception of linear velocity. Th.is apparen t calcula ti on suggests that
w hile otolith organs .sense linear acceleration , our brains tum this i.nfom,ation
int o a perception of linear velocity.
mathematlcal integration Com- As for yaw rota tion, directi on recognition thresholds for s ide- to-side (y-
puting an Integral-me of the two nxis) transla tion h ave been measured for m oti on freque ncies ranging fro m
main operations In calculus (the other, 0.05 Hz fo•)e b..ldc-ond-forth cyde of in 20 seconds) to S Hz (fi ve
the Inverse operntion, Is differentiation).
Veklclty Is the Integral of acceleration . back-anCl-forth cycles of ac le rafion m 1 second). For frequencies above 1 H z
Change of position Is the integral of (on e back-a nd-for th cycle in 1 second), direction recogn ition thresholds are
velocity. ro ughlyconstnnt. To sense tmnsl1tion direction youl' head has to be

moving at least a t a sp eed of 5.0 mm pe r second, \·vhich a t 5 H z corresponds

to a head an gular d isplaceme nt of jus t 0.5 mm! Clearly, we are also very s en-
s itive to trans btion, As for yaw rotation, for frequencies below 05 H z {on e
back-and-forth oscillation in 2 second s), thresholds increase \vi th d ecreasing
freque ncy (Va lko et a l., 2012).
77/t Perception
How wel l d o we perceive OlU' tilt w hen we are s la nted away from tn1 e Earth-
vertical? For tilt angles less than 90 degrees--thatis, body orien tations ben.-.·een
standing up (0 degrees) and ly ing d0\¥n (90 degrees)- we .:'lre pretty good ac-
cording to a va riety of magnitude esti rnation techniques. Observers p rodu ce
relia ble a nd consistent answers if they indica te their perceived tilt verbally or
ii they align a handheld probe with perceived vertical (Figure 12.18).
We are not perfec t, thoug h. Some con sistent e rrors appear, especially in
the s ubjective visual vertical task described nea r the beginning o f the ''Spatial
Orientation Perception " section. Back in 1861, He rmann Rud olf Aubert fm md
that w h en he roll-tilted his head to the left or right whiJe loo king at a ve rtical
streak of light, the vertical Line appeared to tilt in the direction opposite his
head tilt. For tilt angles less than 90 d egrees, the illusory tilt e rror is typically
a bout 10 d egrees, but the app aren t tilt of the vis ua l line cc1 n be as large as
45 d egrees w hen the hea d is tilted 135 d egrees. You can investigate this i!Ju-
IE-OL
sion in a completely d..1rk room by lea vin g th e d oor open jus t a crack-jus t
(11") Pitch Hapticallyindicated tilt

Backw,ud
,;..... 100 0
] ..-. 50 Q
j 0
- lOO
•
,,
Back- - 100 -50 50 100 For-
ward ward
FIG UR E 12.18 Subjects ar9 gen-

(c) Y.lw orally pretty good at Indicating how
much they are tilted. Data (opM
c lrc lf'<S) show tilt perc eptio n provided
by subjects u sing a handheld haptic
indicator (shown schernatk;;ally at
loft) for s tatic pitch (a), ro ll fl:>). and
yaw (c) tilts. P€1rc9v9d tilt roughty"
reflects the actual tilt (solid lines) for
Right -50 0 50 Left
aH three tilt directions. (Aft.er Bor·
Actual tilt (degrees) tolaml ot al.. 2006.)

370 CHAPTER 12
enough to see a line of light in the doonvay without illuminating the room at
all-while you hold your head in a tilted position. Does the line appear to tilt
w hen your head is tilted? Does it move in the direction of your tilted head or
in the opposite direction?
Titresh olds for recogniz ing the direction of tilt show that normal s ubjects
correctly re port the direction of a static tilt w he n tilted about 1 d egree off ver-
tica l in the dark. This sensitivity serves our ability to s tand upright1 because
the farther we are tilted from upright, the m ore difficult standing up is. For
example, the amount of muscular effort required to maintain posture roughly
d oubles if we a.re tilted 2 degrees in.stead of 1 degree, so just imagine the effort
of trying to stand tilted 20 d egrees away fro n1 upright[
Sensory Integration
1he senses d o n o t opera te independently. Ins tead , the brain combines signals
from differen t sensory sys te ms via neural pr0t."e.Sses of sensory Integration.
Fo r example, visua l cues influen ce sound localiza tio n-an effect u sed by
\o"entriloquists. Vestibula r signals combine wi th information from nWTi erous
sensory syste ms to p rovide us with a n understandin g of the p osi tion a nd
movem ents o f the a nd body.
FU RT HER DISa.JSSION of a remarkable example of sensory integra·

tion-the McGurk effect on speech perception-can be fourd in Chapter 11
en page 336.
Visual-Vestibular Integration
Most of us have experien ced illusions of self-motion ca used by moving vis ual
1..-ues. Perhaps you 've perceived self-motion w h.ilewatchin gan IM...l\X movie. Or
perh ap s you've felt as if you were moving backward w he n you were stationary
but the car (or train or bus) next to you began to 1nove forward. Or perhaps you
felt unsteady w hen standing on a bridge looking at the water fl owing beneath.
A ll of these situa tions le."td to pen:"eptions of illusoiy self-motion called vec.
tlon. \'ection can be very compe!Ling. For example, drivers stopped in traffic
often press harder on the brake pedal w he n they perceive that they and their
stationary car are moving, even though it is the cars around them that are moving.
To cons ider hO\·V vection contributes to spatial orienta tion , imagine a per-
son s tanding upright w hile vievving the ins ide o f a sph ere rotating about an
Earth-horizontal axis (Figure 12.19a). At first, subjective perceptions match
reality; human s initially p erceive that th ey a re s tationary and that the s phere
is rotating. But if they continue to observe the rotating visual d isplay for 10
.secon ds or so, they us ually begi n to perreive that they're rotating in the direction
opp osite the sph ere rotation (Figure 12.19b). lhis illusory ro tational vection
sensory integration The process
demonstrates the crucial contributi ons of vision to our sense of self-rotation .
of comblnlr.g different serroory signals.
Twlcaly, combining several signals ln fact, signals rela ted to vision con verge with th e se1nicircular-canal signals
yields more accurate and/or more pre- in the vestibula r nuclei, w hich is the firs t place in the brain tha t vestibular
cise Information than can be obtained infor mation reaches. (Web Essay 12.2: Canal·Otollth Integration discusses a
Iran lrdMdual sensory signals. This s inUlar interactio n of s igna ls from the semicircular cetn.:'1ls and otolith organs.)
Is oot the mathematical process of
Paradoxically, subjects experiencing such ro tational vection almost never
Integration learned ln calculus (e.g., the
Integral of acceleration Is VelocltY). report that they are tumbling head ove r heels as they wo ul d if they truly were
rotating to the extent that they perceive. In fact, s ubjects typically experience a
vection An Illusory srose of self-
motion caused by moving visual simulta.neo tts illusory sensation of tilt that graduaUy builds up to a relatively
cues when one is rot, In fact. actualty cons tant level (Rgure 12_.1 9c). Theseperceptions--experienced as a sensation of
moving . motion \11i--ithou t getting O:Ul}""--Vh ere--are rontrad ictory,since we canno t be rotating

THE \IESTIBULAR SYSTEM AND OUR SENSE OF EQUILIBRIUM 371
(a)
FIGURE 12.19 Rotational vectlon . A

subject views a \Jlsual dlspla'f rotati ng
relative to gravity (as suggested by the visual a.1es) while also maintaining a In roll. For demonstration purposes, the
constant orientation with respect to gravity (as indicated by the otolith organs}. visual display Is shoW1 hQrg as trans·
ln addition to exemplifying sensory integraticu, this sensation of motion without parent. '8) lnitlaHy, the subjoct correctty
getting anywhere demonstrates that our sense of spatial orientation is not con- senses tha.t she Is stationary and the
strained to combinations of motion and orientation that are physically possible. visual display Is rotating. '1:;i) The subj9ct
begins to peroetve vectlon, a sense o f
In this example, the role played by the vestibular system is to put the brakes self·rotation in the dlrectlon opposite
on visually induced vection. Patients suffering from severe vestibular damage the rotation of the visual dlspla'f. (c) Roll
generally report greater vection than do normal s ubjects.Astronauts experienc- V9Ctlon ls often accompanied by an
ing rotational vection in space-in the absence of gravitational signals--report Hlusion of roll tilt that is induC8d by the
a head-ovet'-'h eels twnbling sensation that is a bsent on Earth. These findings peroetved roll rotation. The d!rectlcn
o f the percelVQd tilt Is consistent 'With
are explained by the fact tha t neither the patients n or the astronauts receive the tilt direction that 1NOuld occur if thQ
normal gravitational cues fr om the o to lith organs to contradict their visual subject Wf!f9 truly rotating In roll. CAfter
rotational cu es. (Web Essay 12.3: Space Motion Sickness discusses another Young, Sh&lhamer, and Mod&stino,
aspect of spaceflight that many astronauts experience.) Since illusory motion 1986.)
is greater when there are no o tolith cues to contradict the visual cues, '\o\"e can
infer tha t, under nonnal circumstances, infonnation from the vestibular system
is combined w ith visual information to }';eld a "consensus" about our sense
of spatial orientation.
Active Sensing
Our sensory systems are simultaneously activa ted as the result of our ow n
actiol\s and changes in the external world. Indeed, m ost of our sensory experi- sensory reafference Olange In
ences are gained by active exploration of the world resulting from locomotion, afference caused by self-generated
eye m ovements, tou ching, and other interactive activities. Our brain's ability activity. For the vestibular system, ves -
to d istinguish sensory events that are self-generated , sometimes called sen- tibular afference evoked by an active
sory reafference, from those that arise externally, sometimes called sensory head motion wruld
yield smsory reaffermce.
exafference, is essential for perceptual stability and accurate m o to r control.
For example, w hen our eyes move, the im.age of the world m oves across our sensory exatterence O'lange
In afferenoe ca.used by external
retinas; yet we do n ot pe.rcei ve the image of the world as m oving.
strnull. Fe< the vestlt:>tJar system,
In order to avoid responding to sensory inputs that arise from self-generated vestibular afference evoked by pas -
actions, the sensory system needs to know what the motor system has d one. sive head motion would yield sensory
Based on their observations, Von Holst and Mittelstaedt (1950) proposed the exafferenoe.

372 CHAPTER 12
(a)
Vestibular Estimated Estimated

.Ufe1l:'nce - vestibular = veslibufar
Estimated reufference exafferenoe
vestibul.u
reafference
(c)
Active
Ve&tibular Estimat.ed. E'sti.nlat<i! d

,lfference - vestibular = vestibular
re.:ifferen ce exaffere noe
FIGURE 12.20 Sim plified rQPrQSQntation of the actiVQ VQstibular S9nsing nGtwork.
(a) When a command Is sent to the ntlCk musclEi to rotate the head vla efferent motor
n'11...lrons, a copy of thoS9 mot or command sgnals- cal led an "eff9renoe copy• - is
sent In to the brain. This signal ts processed by a nrural network. call&d an
"internal model," that pr.;;idlcts the 8Xp9cied vestib.1lar afferenca due to the self-
gengrated m otio n, which is call.ad vestibular rsaffergnce, • This estimat ed
reaffen:nc e is subtrnc ted from the signals from the vestibular affo;irmt neurons to yield
estimated vestibular QXaff"Qr9flCQ, the of the affer9nt signal that is no t dlJQ
to motion. ft;) When thie hea.j un09rgo eis passive rotatlcn, the head
ro tation is accuratQ!y encodQd by the vestibular afference (blue). No neck m otor com-
mand is gener.ited duting passive ro klllon . so the passive expgc ted vestibular
enoe signal is nat {green). c orresponding to no sSf-generated m otion. The d iffa,renc e
b9tween th9SQ )'iG!ds estimated vestibular which, for passive rrotion ,
Is identical to the vestibular affgent signal. (c) WhEf'I the head is acttvely rotated, the
head rot ation is accuratQ!y tilncoded by the vestibular affiar0nc9 (blue). In th is cas9, a
neck motor command is generated to evoke the passive ro tation , so a mo tor com-
mand ls sent to the neck muscles and a oopy is S9nt to an Internal m odel that
dic ts the vestibular afftfQllCQ anticipatg,d for th9 actlV9 h 9ad rotation OOng g9nQFat9d
(gre€n}. For some ve:;tibutar nudel neurons, tho;i differenc e between th9 vestlt:ular
afferQflCe and th9 estimated r9afferenoe y ieilds 9Stimated QXaffe-t;!f')Ce, w hich is n9arty
flat. (Parts b.c after OJlleo , 2011 .)
principle of reafference. In today's version of this principle (Figure 1220a),

a copy of motor commands called an efference cop y is generated to h elp the
brain p redict the expected sensory results of motor commands. TI1is p redicted
sensory activity is called sensory reafference an d the brain 1s estimate of sensoiy
reafferen ce is s ubtracted from the sensory afferent s ignal to elimin ate reafferen t
information. In certain model systems, il\duding the electrosensory systerns
of electric fish, self-generated sensory information is selective ly suppressed
at th e level of afferent fibers.
Signals fr om the vestibular a fferent neurons d o n ot distinguish between
ac ti ve and J;tead movemen ts; afferent ne urons respond iden-
tically to sell-genera ted and externally a pplied self-motion (Figure 12.20b ,c).
But differential processing of vestibu lar signals is evident at the n ex t s tage of
processing in the vesti bular nuclei. The mod ul ation o f so me vestibular nuclei
n eurons, wh.ich receive direct inpu ts from the vestibu lar afferen ts, is dramati-
ca lly atten uated in response to vestibular inputs that res ult fro m se lf-generated
movem en ts. The drruna tic di£fere1'\ce the responges of these vestibular
nuclei ne urons demonstrates a dear example of active sens ing.. sll1ce this brain

signa l d irectly depends upon w he the r the m otion is self-genera te d. (acti ve)
or not (p assive). \Ve e mph asize that this distinction between self-generated
ne ural acti vi ty and externally gene rated ne ural activity a rises at the first central
sy napse in the ves tibular nuclei for three vestibular pathways tha t contribu te
to balance and perception. This shm . .·s tha t this fundamental dis tinction occurs
very early in the processing of sorne ves tibular informa tion. This s u ggests that
active vestibular sensiltion, the ability to dis ting uish between self-generated
and externally gene rated neural activ ity, is htndarnental.
Reflexive Vestibular Responses

As m enti on e d e arlie r, som e crucia l contributi on s o f the ves tibula r sys te m balance system The sensory sys-
are a utom a tic, o r reflexive, d oing th eir work o u tside of conscio us aw areness. tems, neural processes, and muscles
For example, there is a set of automati c responses called vestibulo-ocular re- tliat contribute to postural control.
Specllc components Include the ves-
flexes--VORs fo r shorL \Ve started the wi th one of these. Remember tlbutar crgans, klnoott1eels, vestlbulo-
lookin g a t your finger as you sh oo k your head ? (lf not, now might be a good splnal patt1ways, skeletal bones, and
time to go back and try this de monstration again; see F'igure 12. 1.) TI-tls task p::E>tural control rnusdes. Because of
illustrates how the \ · oR contributes to vis ual s tabili ty b}'· coun terrotatin g the Its cruc191 contributions to balance,
eyes in the head during head m otion sensed by the vestibular system . some even informally refer to the ves-
tlbUlar sys-tern as the 1 balanoe
1here is set of vestibulo-autonomic l'e flexes til.'lt rontribute to auton omic and the vootlbular organs as the ' bal-
responses like tl"""' tlIBt regulate blood pressure and help 1mtln Wn adeq uate blood ance organs... But the balance system
flow to the brain. Other vestibul o-auton om ic reflexes lead to motion sickness. Is much more than 1ust the vootlbular
Finally, vestibulo-spinal reflexes contribute to postural control via the balance system, and the vestibular system
system. To demonstrate vestibul ar contributions to balance, tty s tanding on contributes to more than Just balance.
one foot '"'ith your eyes closed. (When you s ta rt to feel \.ms teady, immedi atel
open your eyes and put both feet on the ground !) Mos t hea lthy people with a
normal balancesystem-inc:lud_ing nom1al vesti.bufar fun ction--can do this for
at leas t 5.secondsa nd often mud 1 longer. Patie.11ts without vestibula r function,
however, cannot stand on one foot in the d ark fo r more than an instant.
Vestibu/o-Ocular Responses
The angular VOR, the eye rota tion tha t helps compensate fo r angular rotations
of the head, is a robust reflex. In fact, this reflex is so robus t tha t it is a st:md ard
part of clinica l examinations of vestibula r fun ction. Furtherm ore, the VOR is
certainly amon g the best-studied reflexes, d ating as far bac k as the late 1700s
w ith rep or ts by William Ch arles Wel ls (1792).
The ang ular VO R is th e compen satory eye rotation evoked by the semi-
circula r cana ls when they sense head rotation. For exa mple, w hen the head
ro tates (yaws) to the left, the reflex pa thways ca use the eye to rotate to the
right with respect to the head, to compen sa te-at least in pa rt-fo r the head
tum. \ Vh en observing th e eye, we see this eye rotation as m ovemen t of the
pupil to the right. Of cou rse, s in ce the eye is rea lly a spherical eyeball, it is
really rotating in the eye soc ket, no t movin g latera lly.
Recall that six ocu lar 1nuscles--called "oculom otor mus d es"- rot..1.te the
eyeball (see Figure8.16). Muscles can o nly pul l; they canno t push Muscles are
paired to pull in opposi te d irections and a re therefore called agonist-antngonis t
muscle pairs. For example, m ovin g the left eye h orizontally req\Lires a coordi-
nation of the la teral rectus that pulls the eye to the left a nd the m ed ial rectus
that pulls the eye to the right. To m a ke an eye n10vem ent to the right, the
latera l rectus is inhibited a nd thus re laxes its pull, while the medial rech1 s is
excited and thus increases its pulL The six oculom o tor musc:les are orga n ized
in three pairs tha t rotate the eye in ead1 of three direc tions (see We b Activity
12.3: Observing Torsiona l Eye Move m ent). Eye m ovem ents result from a
coordi nated inhibition Lllld excitation of the eye muscles.

374 CHAPTER 12
TMget
FIGURE 12.21 Contribution of the angular VOA to \lisual stablllty. '8) the
"1'(QS do not c ount0rrotate, the image of the object Is not fowated during head rota·
tion. VVhen thQ ef'/9S counterrotate during head rotation -with the aroount of eye
rotation roughty equal to the amount of head rotation-the Image of an object can
r9main fowa.t9d even during hQB.d rotation. Because th9 Q)'QS counterrotatg in the
head In (b) but not In they continue to look at you in but not in (a}. This effect
demonstrates how the angular VOR oontribuh;1s to vlsual acuity, by helping to k9Bp
otject imaggs stationary on the retina, thiereby reducing
\Vhen the VOR is working effectively, it actively rotates the eye in the
head sud1 that the rotation of the eye roughly compensates for the rotation
of the head in space. Figure 12.21 demonstrates this with a simple example.
In Figure 12.21a, the eyes rotate with the head; they do not counterrotate in
tl>e head . ln Figure 12.21b, the eyes counterrotate in the head, which helps
stabilize the visual field on the retina. If the eyes do not counterrotate in the
head, the retinal image tends to blur during head rotation. (Remember how
your fingers blurred when you rapidly moved them back and forth?) The
COWlterrotation of the eye in the head helps reduce this blur by reducing the
motion of the image of an object across the retina. Note that because of eye
counterrotation, the actual rotation of the eyes with respect to the external
world is much less than that of the head.
The most direct neural path for the VO Rs consists of an arc of three neurons
that yields reflexive eye responses with a latency of less than 10 milliseconds

Left eye
'
. Right eye FIG URE 12.22 Neural pathways for
Superior the thr.ee-neuron arc of the angular
Supeno'---.,
.
""tus VOR. Eac h type of neuron Is
""""'
Lateral / Llteral
using a different ooto r. Eac h of the
vestibular afferent neurons (btue) - one
""'"'' - branch for each of th9 three canals -

projects to one of thrge vestibular
nuc lei: superior, mediaJ . or lateral. Ntlll-
rons that oonn9Ct aff91"'9f"lt to Q'ff9rient
nrurons. caJled intern;:o.urons (green),
project from the vestibular nuclei to the
thrQQ ocular m otor nuc lei: abdUCGns,
trochloor, and oculornotor. (Only the
nuc leus, which is O M
of the three ocular n10tor nuclei, is
sho'M'I.) The six eff9rent oculomotor
nieurons (red) project frcrn these thr99
ocular motor nuclei to the six oculotno-
tor musd.as.; lnferbr oblique, superior
inftiflor rec tus, superior rgc tus ,
lateral rectus, and m9dial roc.tus. Recal l
that you previousty W9re introduc9d t o
the orutomotor muscles in Agure 8.16
In C haptEM" 8.
Vestibular
nuclei:
Superior
Medi.al
L1teral
Inferio r -
Pons
I
Vestibular organs
(ms) b etween the start of head motion and the eye movement. (That is really
fast. Try doing anything else in less than 10 ins!) The first ne urons in the arc
are the afferent n eurons {bottom right in Figure 12.22); these ne urons transrnit
information from the vestibular periphe ry to the vestibular nuclei. the
afferent neurons synapse on intemeurons. These intemeurons synapse oneffer-

376 CHAPTER 12
(a) O.Dl Hz (b) 0.05 Hz (c) 1.0 H,
Tnne(s) Tnne(s) Time{s)
FIG URE 1 2.23 VO R responses in the dark at thr99 Thie s timulus

(sinusoidal head rotation; top row} is shown for (a) 0.01, {b) 0.05, and {c) 1.0 Hz.
S inusoidal rnotio n evokQs an oscillatory VOR (bottom row'), \Nhlch o ppos9S the hoBad
rotation suc h that head rotations to the right are acoompanied by eye rotation s to
the left. If the VO R weKe perfec t, It would have the same pMk am plitude (labeled
•A" in the figure) as the head rotation , m ean ing that thg, eye rota ted <:lt the sam e
velocity as the he.ad rotation. However, the p.eak. compensatory eye velocity shown
for 0.05 and 1.0 Hz in th€! dark is about 70% o f the peak head velocity. We would
saf that this has a gajn of about 0.7. Wh€f.si gain is a d imensionless ratio of the eye
velocity d i\lided by the head velocity. A gain of 0 would indicate that the eyes d id
not rotate with respect to the head• .3S in Agu re 12.2 1a, A gain o f 1 would Indicate
that the eyes were counterrotating with the sam e amplitude as the head mo tion, as
sugg9stQd by Agure 12.2 1b. The rem aining 30% is usually m ade up by v isual oo n-
tri butlons. At lower frequencies, the VOA Is smaller. for exarnple, at 0.0 1 Hz it is onty
about 50% o f the peak head velocity (a gain of about 0.5).
ent oculom otorneuron.s in the ocular motor n uclei. These oculom otor neurons
synapse \.Vith the ocul omo tor rnuscles to rota te the eyes "";th respect to the head.
As you rota te your head from s ide to side, the ang ular VOR h as properties
sirn.ila r to the charac teris tics that were d iscussed earlier for the semicircular
ca nals, as ill ustrated in Figures 12.23 a nd 12.24. The VOR d emons trates a
8230 high amplitude at high frequencies, b ut it gets sm aller and
smaller as the frequency drops below abo ut 0.05 H z (Figure 12.23). Though
the freq uency characteristics of the VOR and can al afferents are qualita tive ly
simila r, there is an interesting d ifference. As Pigu re 12.24 shows, the response
of canal afferent neurons declines for frequencies below 0.2 Hz. ln compa rison,
the VOR declines for frequencies below 0.05 Hz.
TI1at difference is in teresting because it means that, in some sense,, the VOR
is responding more accurate ly than a simp le read ing of its inp ut signa l would
provide. This effec t is an alogous to what we saw earlier fo r rotation percep-
tion (see Fig ure 12.16) and called "velocity s torage." It s hmvs that the brain is
not si mp ly a relay station that passes sensory inform ation to the muscles tha t
yield the refle.xive action. Instead, the b rain helps sha pe the reflexes to be as
effective as possible 1 given the avai lable sensory in.form ati on . In thi s specific
case, centra l p rocessing increases the effecti ve VOR fre'l u ency range (Raph an,
M ats u o, and Cohen, 1977; Ro bin son , 1977).

FIG URE 12.24 Dynamics of the VOR lblaci< cu!V9)

ID at"€1 such that the response has a nea.11y constant
0.9 gain- roughly again o f0.7, as calculat ed in Figure
OB 12.23 -at frequencies 00tw9e0 about 0.05 ond 2.0
07 Hz. In fact, whlle not shown in this figure. the VOR galn
0.6 VOR roliwf'lains rnmrly c:onstant for fre qu8nci9S up to 25-30
Hz (Hutere< and C ullen . 2002: R. Ramachandran and
05
Lisberger. 2005). Perfectly oompensatory response-
-...... Sem idrcul.u canal
·"&, 0.4 afferent neu rons
having a gain o f 1 - is shO'M1 in r9d. SQe Figure
12.21 for examples of how this gain is calculated at
"'
0
> OJ
each frequency. Bebw 0.06 Hz, the gain of th9 VO R
decreases. For ccmparison, the ncnnalizOO frequency
rooponse of semicircular-canal neurons that we sav-1 in
Figure 12. 13 is indicat.ed by th9 gr9811 c urv9. Sln09 the
0.2
Qreoen curve rep-esmts the inforrnatton pro\lided to the
brain and the black line represents the VOR respon se
at different frequencigs, the d ltferenc.e between the
black and grgen curves rspresents nrural compMsa-
tion performed b y the brain.
0.05 0. 1 0. 2 0.5 1.0 2.0
Frequency (Hz)
\¥e also h.n:e a translational VOR that is evoked when the o tolith organs
se nse head especia ll y h igh-fre quency h ead translation . Th is
translational VOR helps us keep our eyes pointed a t an object w hen the head
tra n.sl'-'l tes in one direction or the o ther.
Vestibulo-Autonomic Responses
The vestibu.lar .system also m akes contributions to responses o f the autonomic autonorric nervous system The
nervous system. Perhaps the most vivid of these responses is v1otion Si.¥e l Pf'!1 at the nEilVOUS system that Inner-
a vestibu.lo-autonom.ic ordeal that m.any of us wish we had never experienced. vates glands, heart, system,
and so on. ard that O! responsble for
Severe sy mptoms of motion sickness i.ndude nausea and vomiting. Motion reguatlng many Involuntary actlora.
sickness typical.ly results when there is a disagreem ent between the motion
and orienta tion signals p rovided by the semicircular canals, otoHth orgt.ms, and
vision (Oman , 1990; Reason and Brand, 1975). For example, if you nre below
deck on a boat, your vestibular system w i.11 <1C'C'l.1rately record the motion of
th e boat while vision su ggests no relative m otion of the world, sin ce you <ind
the boat are m ovin g together.
\Vhat is this resp onse good for? One of severa l hypotheses is that it is a
d efense again.st some dasse.s of p oisons (M. Treisman, 1977). lf you have been
poisoned, you want to get rid of the poison before it gets rid of you. But h ow
d o you know if you have been poisoned? lf the poison is a neurotoxin, disrup-
tion of the sensory systems is a good hint. How do you know tha t your senses
have been d isrupted? One way i.s to check whether senses that norrna lly agree
w ith one ano ther have s topped d oing .so. Nonnally, if you move, you r visual
system and your vestibula r system both register th.."1. t fact. If the vestibular
syste m says one thing and the visu al .syste m says a nother, the brain may
d ecide that it is time to rid the bod y of a possible cause of the d isagreement.
This response could be a lifesaver if yo u just ate a bad mushroom. It is less
d esirable when you have to pull out the mo tion sickn ess bag during choppy
weather at 35,000 feet.
Other vestibule-autonomic responses are less spectacular and generally take
the form of compensatory contributions. For exarnple, cons ider the proble m
of regulating b lQOl..1. pressure. The heart ptrn1ps blood throughout the body,
bu t m aintaining oxygenation of the brain blood fl ow is especially critical,
because you will Wack out f.n just a few seconds if your brain d oes n ot receive

378 CHAPTER 12
(a)
(b)
:r
..
1l
zS- -5
- no vi&lJ<li CUe!I l Appa
e g_
- 10 -- Lesioned,. n o visual cues
20 30 40
Tune following whole-body tilt {s)
FIGURE 12.25 Vestibular inftuenoes on blood pressure. (a) This trace represents
blo::>d pr9SSllr9 In the h9r0.d vGrsus tirTI9 in respons9 to a nOS9·up tilt in a w ith
a lesloned vestibular sys.-tem. In the absence of wstibular contrlbutlcns. a transioot
decrease In blood pressure can 00 observed immediately after the tilt. (b) C hange In
blood pressure is much greater for the subject vvithout a functional vestibular system
Qesioned) than for the subject with a nom1al vestibu lar system, d.:monstrating that the
vestibular h9ps maintain m ore constant blood pressurie during whola-body
tllts. (After Yates and Miiie', 1900.)
adeq uate oxygen. Gravity pulls blood downward, so in the nonnal upri ght
posture your hea.rt has to work to maintain bl ood flow to the brain . When
you 're lying down, it takt>S mud\ less work to pump blood to the brain. Now
s uppose you rapidly stand u p. TI1e cardimr.ftSt.""'UJar system has to s uddenly
change the regulation of blood flow to maintain adequate oxygen supply to
the brain. If these mechanisms fail, you vv'ill experience light-headedness or,
in extreme cases, blackout By in.formb1g the relevan t parts of th e autono mic
nervous sys tem about the position and rno tion of the head, the vestibular
system contributes to the regulation of blood flow to the brain. Asa result1 one
consequen ce of destmctive lesions of the vestibular organs is that blood pressure
regulation during whole-body til ts becomes much less stable (Figure 12.25).
Vestibulo-Spinal Responses
In the early 18QOs, the Fren ch physiologist Jean Pierre Flourens reported ab-
nom1al head movements in pigeons after he had cut individual semici rcutu
canals. TI1e head movements he reported were in the plane of the lesioned
canal. These s tudies demons trated the presence of w ha t we tod ay consider to
be the vestibular influences on posture con trot and they initiated the formal
s tudy of vestibular influences on balance.
Vestibular reHexes keep us from falling over. \Vitho ut ves tibul o-spinal
reflexes our ba lance would be severe ly degraded . We would be unable to
s tm1d in thedmk. Prut of the reason is that w hen ,.\,:e s tand, we are inherently
unsta ble. As tvo.·o-legged crea hues, we ca n be thou ght of as being composed of
a series of inverted pendulmns--penduh.1n1s b1. which the mass is above th e
pivot point. Bala ncing a pencil or broom on your palm demonstrates a simple

(a) FIGURE 12.26 The contributions of the vestibular

10 $'.{Stern to balance are demonstrated by the compari-
son of postural responses of subjects with normal
Vo9stibukat" func tbn to r€!1Spo nses of patients suffGiring
swere bilateral vestibular loss. Subjects We<e asked t o
maintain therTISQt.•as upright in thei presen09 of small
anglilar displacements of their feet that were d9slgned
to chaJOOge their balance ccintro l. Fcr the purposes
o f illustratioo, a platform tilt of 20 dQgr99S is shown:
in typicaJ experiments the tilt '#OUld be much smaJler
82" lottl«MS<> tho might fal offt). I!>) Th<> values on
the y-axis (the gain) represent the ampNtude o f the sub-
- Nonnal ,iect's responSQ d\1decl by the amptitude of displace-
fTlS'lt. (Gakl In this c ontext Is defnQd as tilt of the s ub-
-- Vestibu Llr-IOGs p atients
ject di\'ided by tilt o f the platform.) (b) If the subject did
0.1 the task p«factly, th9 rQISUlt VV<>lMd bQ a gain Of o ('019).
If the sub)ect remained perpendicular to the platform,
O.o! 0.1 10 the gain would be 1 (919). If the subject tilt€icl more than
Frequen cy of platfotm tiJt m otion (Hz) the patfonn. the gain would be greater than 1. The
response gains for normal subjects were atways near
(b) or below 1. On the other hand. vestibular-loss patients
Subject tilt Subject tilt Subject till had responSQ gains that often 9X09Qd9d 1 , indicating
=CJ' = 20:- > 200 that their b ody tilt exceeded the platform tilt distur-
bance. Patients vvlth vestibular loss exhit)ted much
greater sway than normal subj9Cts, dearty deimonstrat-
ing the fundameirrtal contr1butlons o f the vestibular sys -
tem to balanCQ contrd. (After Plilterka. 2002.)
Platfonn Platform Plltform

tilt :: 2C1' till :: 21Y tilt ::20&
Cain :: 0 Cain : 1 Caln ;i. l
inverted pendulum. These challenging tasks can be mas tered with practice,
but imagine trying to s tabilize a series of at le.:1st four brooms or pencils on
top of one an other. At its essence, this is the task faced by our balance systein,
which works to keep the head, torso, thighs, and calves- each an inverted
pendulmn- upright w ith respect to grav;ty.
A th orough discussion of the dynamics of the entire p osture control sys tem
is beyond the scope of this book. To s tudy the poshtral system experi.Jnentally,
investigators have simplified th e dynamics by s trapping the head and body to
a rigid boa rd that converts the complex multi joint body into a single inverted
pendulum pitdring fonvard and backward about the ankle joint. \Vhen the foo t
plate that the subject s tands on is gently rocked fon,:ard and backward while
the subject's eyes are dosed, nom1al s ubjects dem ons trate body sw·ay that is less
than the amplih1de of the applied rocking m ovement (Figure 12.26), indicating
that the vestibular system is helping to compensate for the applied- movement
d isturbance. The \·estibular system measures the movement of the hea d and
sends commands to the poshual contro l system that help red uce the amo\.lnt
of body sway. In contras t, patients \v:i th severe vestibular loss demonstrate
body sway that exceeds the amplih.1de of the dis turbance. This difference
behov'een p oshtral responses of people with n ormal vestibular ftmction and

380 CHAPTER 12
,,,..,
Ro.stral \ ..
medulla
(oontrolling r;(
neck muscles)
Medial vestibulo-splnal
fibersinmedi..al
longitudinal fas:iruli
Lateral
vestibulo-Eipinal
!nod
Lower part
ofcervic.U
spinal rord
Lateral
\'estibulo-
spinal tract
:;;;:,:,,
Lumbar
spinal cord r Mc:itatory synapt'e
To flexor muscles
To extensor mua:k>s
FIG UR E 12.27 NQUfal pathways for vestibu lo-splnal reflexes. Most vgistlbular affer-
ent neurons (blue} synapse In cnt1 of the vestibular nuclei, but some project to the
C8"eb4illlum. From thQ wstibular nudel, desCQndlng lnterneurons (green) carry the
inforrnatlcn via wstibulo-splnaJ tracts downward through thrci brain stem and spinal
cord until they synapse on efferent neurons (red) that activate the musdes that con-
trol balanca. (After Nettgr, 1983.)
those of patients with vestibular loss is a clear indication of the importance of

the vestibular system to balance and posture control (see Figure 12.26). A little
later in the chapter, we' ll discuss mal de debarquement syndrome, an unusual
spaUal dsorientatlon Impair -
ment of spatial ooentatlon. More spe- disorder that causes imbalance and/ or spatial disorientation.
clflcally. any Impairment of our sense The vestibule-spinal response can be thought of as a whole family of
of llnear motlonr angular rootlon, cr tilt. reflexes (Figure 12.27 ). In the vestibular nuclei, the primary afferent neu-

ron s synapse o n d escending interne urons tha t carry information thro ug h

the lateral and medial vestibule -s pinal trncts. H ow far these n e u rons carry
information d o'W11 the s pinal cord depends o n their contribution to the bal-
an ce system . If the interneuron synapses o nto a n e uron that contro ls a leg
musd er the information is tran smitted beyond the bottom of the spinal cord.
If the information con tributes to p os tural contro l of th e headr it is tr.:i.n..smitted
only as for as the neck.
Spatial Orientation Cortex

We have visual cortex and cortex. Do we have a vestibular cortex?
Some areas of the cortex certainly do respond to vestibular input, but these
areas tend to include a varie ty of sensory and mo to r signals. There does no t
appear to be a n area of the cortex exclusively dedica ted to vestibular signals.
This is no t shocking. sin ce we learned earlier tha t pe rception o f body mo ti on
and tilt result fr om multisen sory convergence, includin g pred ominant con-
tributions by th e vestibular system and vision.
It may be that there is simply no good reason fo r the cortex to
ves tibular information in isolation w hen o ther sensory informati on is avai l-
able. For example, the visual system is respon sive to constant-ve locity visual
motion-like that experien ced during rotation in the light as you see the room
spin around you-while the vestibub:r system responds primarily to ch anges
in velocity a nd is relatively in se nsi tive to con stant-veloci ty 1no ti on . More
specifically, reca ll that w he n rotated a t n constant ve locity in the dark, after
'1bout 30seconds htm1ans perceive that they are not rotating (see Figure 12.16).
Therefore, when rotatin g with a nearly con sta nt ,·elocity in the light, it seems
reasonable that the brain would utilize the vis ual information indica ting mo -
tion to bette r e.s tiulate self-m otion . It thus makes sense that areas of the cortex
related to the percep tion of tilt a nd .self-motion d em onstu1te a convergence o f
visual and vestibular in.forma tion, as well as information from other sensory
systems tha t contribute to s patial orientation.
Vestibular Thalamocortical Pathways

Vestibular information read\es the co rtex via w hat are ca lled the '' tha lamo-
cortical pathways" (R g ure 12.28). This m ean s s i.mply tha t the ves tibular
information , like most o ther sen sory information, reaches the cortex via the
thalamu s. Ne urons fro m the vestibular nudei carry vestibular information to
the thalamus, w here that in.formation is processed and relayed to the cortex.
Cornpare this, for exa mple, to the visual system., w here the retinal ganglion
cells synap se in the lateral geniculate nucle us of the thalamus (see Figure 3.16
and "The Lateral Geniculate Nucleus" section in.Chapte r 3). There the infor-
mation is processed and p assed on to visua l cortex.
Evidence s uggests that the temporo--parieto-insular cortex is involved in
spatial orientation perception. 1his area of the cortex receives input from both
the semicircular canals and the o tolith organs. Fu.rthe r1nore, inunediately after
this part of the cortex is lesioned by a stroke, many patients report illusory tilts
and/ or ill usory translation . l11ough mo re rare, rotation al vertigo-an illuso1y
sense of spinnin g-is reported by some o f these patients.
Though m ore direc t vestibular projections may exist, there is a vestibular
pathway that leads to the h.ippocampus thro ugh the cortex, and nemons i.n
the hippocampal fo nnation respond to vestibular s timuli (Horii e t al, 2004).
These include a set of ne m on s called 11 head direction cells'' becau se they
tend to spike vigorously w h en an a nimal's h ead is p ointed toward a speci fic
direction (Taube, 2007).

382 CHAPTER 12
FIGURE 12.28 A$00nding veistlbular CentM1

(thalarnocortical) patt-iways pass fro m fis&ure
th9 VQstibular nuclei t o thQ thalamus
Paiietal
lobe
on their way cortQX. The braln is
\oiewed from the right side. CortIDC acti·
vation patt€W·ns shown at the top are
PET (positron emission tomograph)?
scans (the on th9 left is oolorlze::i)
obtained during stimulation of the ves-
tibular system on the right side. RQd
and yellow indicate activation. which
occurs in the ternporo-parieto-insular
areas of both hemispheres. Unlike
roos t oth€11' sensory system s, much o f
the vestibular activation is not 'Jisible
on the outermost su 11ace of the cortex
but is found deeper In the brain, in
the Insular cortex. (Brain from
uh
Dieterich and Brandt, 2008; cou rtesy
o f Marianne Diet'1irich.)
Vestibular
nuclei
'/
8230336 AmMa Appa
/
I
Vestibular
cirga ns
Cortical Influences
1lle areas of the cortex tha t receive projections from the vestib ular system.
also project back to the ves tibular nuclei. The exis tence of these pathways
suggests th at feedback from areas of the cortex that resp ond to vestibular
s timulati on likely rnod ulates low-level vestibular processing in the brain
stem.. A specific role fo r the:se projections h as n ot been but it is known

tha t higher cogn..itive knmvle dge can affect bo th perceptions and reflexive
responses. For example, imaginin g whether an unsee n visu a l targe t ro tates
with you alters the VOR evoked hy ro tatio n . As is approprh1te, the VOR is
s uppressed w h en s ubjects imagine that the targe t m oves \vi th them but is
relatively la rge whe n s ubjects imagine that the target is Earth-fixed (Barr,
Schttltheis, and Robinson, 1976).
As another example of higher-order cortical influences, knowledge of
the motion capa bilities of a s pecific d evice can influence motion perceptions
(Rader, Oman, and Merfeld, 2011; I Verthei m, Mesland, a nd Bies, 2001). For
exa rnple, if you '"'ere blindfolded and then taken to a merry-go- round that
you had never ridden before, your perception of tilt and motion might differ
from. wha t you would expe rience if you were riding a merry-go-round tha t
you had ridden m an y times before. lli.e vestibular stimuli might be the same,
but your knowledge and expectations--both of which are higher-level fun c-
tions-wo uld be different and could alter yom spatial orientation perceptions. FIGURE 12.29 Some symptoms of
vestibular dysfunction, like Imbalance
causing falls . mirror th,., &ff9cts o f alco·
When the Vestibular System Goes Bad hoMc beveragBS. This butto n highlights
how the symptom s of V9Stlbular disor-
We have lea rned that the vestibular system makes fundam ental contribu-
ders could be mislnteq:::o;i.ted.
tions to our sense of s patial orientation and also contributes to a number o f
reflexive responses. W ha t h appens w hen the vestibubr system fails? Reflect-
ing the \vi.d espread influence of the vestibular system, the bad news is that a
lot of problems develop (Baloh and Halmagyi, 1996). Many patients develop
spatial disorientation. Many experience imbalance. cannot see clea rly
unless they make an effort to hold thei r heads absolute ly still Many d evelop
tnotion sickness tha t can lead to or even vomiting. Some even develo p
cognitive p roblems. Because we are usually tm.:1'"'·are of the vestibula r systern 's
contributions, it is possible to misinterpret the actions of people suffering from
vestibular disorde rs (Ftgure 12.29).
The good n ews is that m ost patients partially to the situa tio n . For
example, many patients quickly learn to curtail activities that lead to problems.
This strategy is effective, but 1.."'Urtailliig o ne's lifesty le is not an entirely st'l tis fuc-
tory soluti on. Forhrrmtely, in addition, m ost patients also lea rn to utilize other
sensory information. As \vi th the exquisite of hearing that iss metimes
reported in blind patients, other sensory systems fill in to nelp beh.:w-
i oral deficits ca used by vestibular problems. Th is adaptation, coupled wi th
physical rehabilitation and a curt-ailing of motion-related activities, yields an
altered lifestyle that helps patients acclimate to their disability. Let's consider
just two vestibular d isord ers.
Mal de Debarquement Syndrome

Most travelers feel a little tmbalanced after disembarking fro rn a large ship
fo llowing an extended cruise. Th is imbalance is sometimes accompanied by
m oti on s ickness and by s\11/aying, rocking, or tilting perceptions. You may
have experienced these sensations after spending jus t a few hours o n a boat.
After disembarking, as you lay in your bed, you might have felt the gentle
roc king o f the waves, even though you knew you \Vere pe rfectly s till. These
symptom s are bo the rsome but typica lly dissipate within a few h ours. It is
genera lly believed that these perceptions are an aftereffect of ad ap tation. Spe-
dficall)'i you adapt to the rocking m otion experien ced w hi.J e on the boat. TI1is
adaptati on-" gettin g your sea legs"-is appropriate w h.ile you're on board
the boat, but it is in.appropriate once you're back on fim-.. ground, leading to
transient perceptions of disorient::.11tion, imbalance, and rocking that appear
""' hen you firs t disembark and the-n dissipate as you readapt to firm ground.

384 CHAPTER 12
Relatively rarely, people are tmab le to readily readapt, and the condition
some times le:i'ds to a dinical syndrome called mal de debarquement syndrome
(meaning "disembarking sickness"). For these patients, the symptoms of
spiltial disorientation, imbalance, and rocking last a month or more after they
disembark. In extreme c..1.ses, the syinptoms c.an las t for years and can be very
d ebi li tating. It remains a mys tery why some individua ls are una b le toreadapt
when they leave the ship.
Meniere's Syndrome
Lnag:ine suddenly experiencing dizziness, imbalan ce, and spatial disorie nta-
tion so severe that either you have to lie down (quickly!) or you fall d own .
Lnagine tha t severe m otion sickness en sues and leads to repeate d vo mitin g.
Now imagine that these sy rnp toms can occur su d denly and more or less un-
expectedly at a ny time. Th is is th e p li ght expe rienced by some patients suf-
fering from Mtfniere's syndrome, named after Pro.sper Mffiihe, the Frend1
physician who first described the syndrome in 1861.
Meruere's syndrome afflicts about one in 500 people. It rnay s trike at any
age, bu ty pically the .syndrome first occu rs in mid adulthood. Other symp -
tot in.:Jude tinhit'us (an illusory ringing sound }, hea ring lo.ss, and a feeling
of pain or fullness in the ear-makin g this a n inner-ear disorder and no t just
a vestib ular disorde r.
111e vestibular symptoms of d izziness,. imbalance, and disorientation a:re so
debilitating that patients often are incapacitated w he n experiencing a MhU€:re's
attack. Furthermore, the concern that these symptoms might rehun at any time
can be terrifyin g; some patients become afraid to leave their homes e ven when
they sy mptom-free. Making matters worse, a lthough the vestibular system
is knm•m to be the source for the spatia l disorientation s uffered, the s pecifi c
ca use of the symptoms remains elus ive. Wllilesome think that excess fl uid in
the inne r ear cause5 Me n.iere's syndrome, o thers think tha t several diffe rent
inne r-ear disorders yield this co ns tellation of symptoms.
Treatments of th e disease includ e medications to lower pressure in the in-
n er ear, implanted d evices that provide h'anstympank mi crop ressure pulses,
and some times p rocedures th.at destroy the ves tibulM appara tus. Stop for a
moment and imag ine TI1e transient Mhlihe's syndrome sympto ms are
so.severe that patients (and their p hysicians) a re vv;Uing to ind u ce a permanent
disability just to be rid of th e sy mptoms. For those of LIS lucky enough to have
neve r experienced severe ves tibular proble m s, this provides a small hint at
how disabling the symptom s can be when the vestibular syste m_malfunc-
tions, as well as a glimpse at the fundamental contributio ns provided by the
vestibular system.
Amusement Park Rides- Vest ibular Physics Is Fun

The canals are not very good t ransducers of lmv-fre::::tLJency rotations -
an effect t hat amusement park rides use to great advantage. \l!Jhat
makes playground and amusement park rides fun? C learly these are
designed t o be sensory exp! riences. Certainly hearing, taste, and sm ell
are not the predominant sensory contributors. and w hile visio n may
play a role in the overall experi ence. most of these rides do not require
vision to create the sensatio ns people feel. In fact, some roll er coaster

THE VESTIBULAR SYSTEM AND DUR SENSE DF EQUILIBRIUM 385
patrons enjoy the expertence rnore with their eyes closed ! Playground
and amusement park rides are designed, in large part , to stimulate the
vestibular system. In fact , much of the enjoyment from a geed amuse-
ment park ride derives from tricking the vestibular system in some 'Nay;
typlcal ty, the designer of a good amusement park rtd e Is playing w ith
one or m ore of t11e fundamental characteristics of the vestibular systern .
As a first example, let's consider the simple child-powered merry-
go -round fo und in playground s (see Figure 12.2). These d evices typi-
cally have a great deal of m ass. especially compared t o the mass of
the child ren. The large device mass means that it takes a substantial
amount of time- say 10 seconds or more-for it to speed up or slow
do'lm. Such gradual changes have low-frequency components that
trick the ::::.emlcircular canals into incorrectty sensing angular velocity.
At the same time, the combination of both the radius from the rotaticn
axis at the center of the ri de and the angular velocity at the edge yield a
centripetal acooleration with lov.r-frequency oomponents that is sm sed
by the otd lth organs. Lmv-frequency accelerations trick the brain into
perceiving self-tilt even h1 the absence of actual tut. This divergence of
percept ion from reality is the definition of an Illusion. Such illusions seem
to yield at least some of the fun expertenced when r1dlng amusem ent
park rtdes but can also lead to motion sickness.
Now let s oonsld er the roller coaster (Figure 12.30). Altho ugh part
of the fun of a roller ooaster oomes from the thrill of moving at a high
sp eed, t11e t wists and turns of a ro ller ooaster rnlnlmally change the
sp ood of the carriage. These twists and turns are t here pr1martly to yield
vestibular st1mulation wen beyond that typically experienced in nonnal
Usually, the turns are located where the carrtage travels with near-
maximal speeds-thereby )1 elding hi gh angular velociti es transduced by
sernlclrcular c anals and high linear accelerations transduced by the oto-
llth crgans. These extreme vesti bular stimuli add to the thrtll experienced
during roller ooaster rid es .
FIGURE 12.30 Many, but not all, enjoy the thrill of vestibular stimuli. Much
of the thrill Q)(p0fl9nc.:d while rlding a ro ller ooaster arJs9s due to vestibular
stimulation ,
8230336 Arr1m.:1 App.:1

386 CHAPTER 12
Summary
1. Tlie vestibular organs are the itulef'-ear organs that sense head mo tion and
Refer lo the gravity and contribute to o ur equilibrium sense.
Sensation and Perception 2. The vestibular organs include three semicircular canals (hori zontal1 ante-
ComP'!rion Website rior, and posterio r), wh ich sense angular mo tion , and h vo oto lith o rgans
sltes.slnauer.comlwoHe4e (utride and saccule), which sense both gravity and linear acceleration.
fer acti'-'les, essa,s. study 3. Hair cells are the mechanorecep to rs that convert both orientation ' "·ith
qt.eStions, and other sl.Jdy aJds. respect to gravity and head motion in to signals that are sent to the brain.
4. Our eq uilibrium sense incl udes spatial orientation perception as w ell as
p ostural, vestibule -autono mic, and vestibulo"'cular reflexes.
5. We d o not h ave a vestibular sense, but o ur vestibular system makes a pre-
d ominant contributio n to our sense of equilibrium. More specifica lly, o ur
equilibrium sense u ses inform a tion fro m multiple sensory systems--\'•"ith
the vestibular and visual sys tems making predominant contributions.
6. Spatial orientation includes three perceptual moda lities: linea r mo tion,
ang ular mo tion, and tilt Direction and amplitude are qualities tha t de fine
each of the.se three perceptual modalities.
7. The vestibular sys tem contributes to spa tial o rientation percep tion, but the
brain processes the vestibul ar in.formation to yield p ercep ti ons tha t differ
su bstantially from the signals found on the afferent neurons .
8. We are exquisitely sensitive to head mo tion even in the d ark, recognizing
the directions of rota tio n, linea r motion, and tilt a t very low thresh olds.
9. T11e vestibular sys tem contributes to d ynam ic visual acuity by evoking com-
pens..'l.tory vestibulo-ocular reflexes (VORs).
10. The vestibular sys tem helps maintain balance v ia postura l reflexes.
11. Vestibula r problems a re l\'idespread, and treatments are limited. For
M€niere's synd rome p atients, for example, the symptom s may become so
disabling that pa tients accept treatments that yield perma nent d isab ility ju st
to be rid of the symp toms.
8230336 Amm.:i Appa

Jr 8230336 Aroma Appa

CHAPTER "13
1a Appa
Kim.Joon,parfy-k,u1's '11..i!ion, 2007

Touch
HAVING STUDIED THE PERCEPTUAL SYSTEMS FOR VISION AND HEARING, you
should recognize a number of questions that c urious minds want asked and
an swered about any sense. ln this chapter we consid er the follmving questions
reg..'\ rding the sense o f to uch:
• What a re the physical s tirnuli fo r touch ? More p reciseli whatJQ1;ms o f
sti mulation en ergy lead to the sensati on of being touched ? nma App
• What is touch good for? What might the evoluti onary ''niche" of this
sensory modality be?
• What is the sensory appara hLS for touch, and how do these structures
ch ange touch stimuli into ne ural signals?
• Which n europ hysiological pathways co1mect toud1 receptors to higher-
orde r perception a nd cognition in the brain ?
• How d o we use touch input to detennine the identity (w11at tasks) and
loca tion (where tasks) of objects in the world?
• How d oes the sense o f toud1 compare wi th and interact y,,r:i th other
sen sory modalities?
Let's start with the first hvo of these questions. By its m os t n arrow defi-
nition as a sense, the tem1 touch is used to re fer to the sensa tions cau sed by
mechanical displacements of the skin. These d ispbcem ents occur w hen you
are p oked by your4-year-old nephew, licke d by your d og, or kissed by your
significa nt othe r, They occu r any time you grasp, wie ld, or o then<\ise nMke
contt'lc t w ith an object. \ Ve w ill use the term tactile (the adjective form of touch)
to refer to these mecha nical interactions and will expand the definition of
tor1dr to include the perception of temperature d Kmges (thermal sensation );
the sensation of pain, w hich occ urs w hen mu body tis.sue.s are drunaged (or
potentially damaged) in some way; itdU.ness; and the internal sensations arising
from muscles, tendons, and joints that infom1 us o f the p osi tions m ove-
ments of our limbs in space, tecl1nically called klnesthesla. Several od1er terms
arise in connection wi th the sensory modality of tou ch, varying in indusive-
ness. Proprioception (from the Latin for ''one's own ") incorporates sensory
input from locations internal to the body, such as your stomach, along with
klnesthesia Perception of the posi-
kinesthesia. Somatosensatlon (somn is the Greek word for 1'body '') further
tion and movement of our limbs In
encompasses the input from touch rec-eptors in the skin as well as the pro- space.
prioceptive syste m . As was n oted in Chapter 12, the veshbular system a dds
proprtocepllon Perceptlai me:Jl-
to proprioceptive infomut tion in controlling balance and to sornatosen sation ated by kinesthetic am Internal
in controlling the autonomic nervous syste m. receptors.
lt is difficult to conceive o f our species survi ving without a sense of touch . somatosensation conectlvely, sen-
Pain serves as a sophisticated wt>1rning system tha t tell s u s '"'·hen some tlU.ng sory signals from the skin, rnusdes.
might be in ternally wrong o r w he n an external s timulus might be dangerous, tendons. Joints. and Internal roceptors.

390 CHAPTER 13
enabling us to defend our bodies as quickly as p ossible {e.g., by rapidly moving

away from tbe noxious stimulus). Ternperature sensa tions enable us to seek or
create-a thermallysafeen vi ronrnent. Mechanical sensa tions play an important
role in our intimate sexual a nd reproductive activities, and they p rovide a
p owerful means of communicating om thoughts and e motions nonverbally.
On a m ore fund amental level, touch is important because we can use it to
identi fy and manipulate objects tha t cannot be seen or hea rd. Blindfold your-
sel f for a t least 10 minutes and try d oin g some routine tasks, like makin g a
sandwich, ge tting dressed, or taking a sho\'\:-er. The first thing you v.ill notice
while doing this exercise is just how much our species n ormally relies on vi-
sion to inform us about the world arm.md us. But you s hould also discover
that touch ca n substitute for vision to a surprisin g d egree: You probably won't
have as 1nuch tro uble distinguishing the peanut butter jar fr om the je lly jar as
you rn.ightthink. And if yo u p<1y attention, you will find that you d on 't actually
use vision, or an y sense other tha n touch, very much a t all for some tasks (e.g.,
FIG URE 13.1 CQf"amicJar. Qing buttoning your sh irt, brushing your teeth, opening that jar of peanut butter).
period, O"iina, mid-eighteenth c entury.
TI1ere is on e more thing you may become acute ly aware of during your
experiment wi th the blindfold: O ur eyes and ears can perceive signals from
objects tlmt urs- .fM JrJ;?Q.,l the bod_J,. but we must ulmost a lways be in direct
contactwi tl1 an ob]ect to percerve ·rt by t uch (exceptions to this rule include
a jackh ammer, wh ose on t11e street outsi de we can feel; and the
s un_,. from w hich '\\fe feel wa rmth even though it is milli ons of miles
Therefore, to use touch to learn abo ut the world , we must act. If we want to
know the weig ht of a bea utiful like tl1eone in Flgure 13.1, we pick
it up. We might also run o ur fingers a long its raised a mtours,cu p its rounded
sh ape behveen ou r hands, o r press it to o ur fo rehea d to feel its coolness. In
S llln, touch involves action, argu ably to a greater degree than any of our o ther
senses d o. Fo r further discussion of h ow the sight of an object might invite
you to touch it, see We b Activity 13.1: Need for Touch.
Touch Physiology
Touch Receptors in the Skin
TI1e sites of our sensin g equipment for vision, a uditio n, olfaction, and gusta-
tion are all located in organs (the eyes, ears, n ose, a nd m out11, respectively)
that are more or less dedicated to sensory proc-essing. Some other aninwls have
analogous appendages: antennae. You might think that, for touch, humans
d o n ot have a readily apparent sense organ. In fact, the hum an .sense o f toud1
is m ostly housed in w hat is actu ally the largest and heaviest of the sense or-
gans, the skin , w hich covers an area of approxima tely 1.8 squa.re mete rs and
weighs about 4 kilograms. Touch receptors are em bedded all over the body;.
in both h..1irless and h airy skin. They are also found within our mouths and
our muscles, tend ons, and joints.
Although the ex terna l quality of the skin varies across different parts of the
body (it is dlicker in some parts and thinner in others, smootl...erin sorne regions
and coarser in others, and so on), most skin includes the basic s ubstrnctures
shm-vn in Figure 13.2. The tactile receptors are embedded in both the outer layer,
epidermis Tlie o..rter of two maja called the epidermis / and the t.mderly ing layer, knmvn as d1e dermis . Just as the
layers of skin. eye h as its rods and three types of con es, the skin has multiple types of tactile
receptors. These receptors form the basis fo r multiple ''channels," sp ecialized
dermis The innEf of two major lay-
ers of skin, consist Ing of nutritive and information-processing .s ubsystems that ead1 contribute to tl1e overall sense
connective tissues, within which lie the of touch. For example, if we wrap om fingers around a cube of ice_, different
rnechanoreceptors. channels conve}'· inforrnation about its temperature, its shape, and its texture.

TOUCH 391
FIGURE 13.2 A cross s9Ctio n o f hair·

less skin of the human hand , schematic ally
d erronstrating the locations o f the four
t)'?e:s of rnechano recgptors -Meissn.;.r
corpuscles. cell neurite cornpexes,
Ruffini i;mdings, -and Pacinian corpuscl9s -
:and illustrating the two major ·1ayers of
Epidermis human skin-the epidennis and the dermis.
The subcutis, w hich underliGS th'9S9 outer
two layers as shown here. Is not usually
Included in th9 formal definition o f •skin .•
(After R. S. Johansson and Vallbo, 1983.)
a llneurite
complex
Dermis
8230336 Amma Appa
1
lsubrnhs
We will d iscuss the receptors for these various channels in d etail within
the sections th at fo ll ow (see a lso Web Ac tivity 13.2: Somatosensory Recep.
to rs). In genera11 however, each type of receptor can be characterized by three
attributes:
1. Type of stimulation t.o which the receptor Receptors res p ond
to diffe rent s tirnulus events--for example, to pressu re, \oibration, or
temperatu re cha nges.
2. Size of tire rt.>cepth1e field. Receptors are acti vated when s timulation is
a pplied to a particular area o f the body, which constitutes the
n>ceptive field (recall that the sa me te.r m is used for the recepto rs ot other
sensory systems). The size of the receptive field is the extent of the bo dy
area that e licits a recep to r response.
FURTHER DISCUSSION of recept We fields as they relate to vision can be

found in Chapter 2 on pages 44-47, and throughout D'iapter 3.
3. Rate of adaptation (fast "'""·"slow). A fas t-adap ting (FA) receptor

responds w ith burs ts of action p o tenti<'l ls, fi rst when its p referred
s timulus is applied and then aga in when the s timulus is rem oved.
It does not respond during the stead y s tate between s timulus onse t
a nd offset. [n contrast, a s lowly adapting (SA) receptor remains acti ve
through out the perio d during whid1 the s timulus is in contact with its
recep tive field.

392 CHAPTER 13
mechanorec: eptor A sensory TACTILE RECEPTORS Tacn l e receptors are called mechanoreceptors b e-
receptor that responds to mechani- cause they respond to .m echanical s tilnul ::\tion or p ressure. A tactile receptor
cal stlmulatlon (pressure, vibration, cr consists o f a "nerve fibe r" and an associate d expand ed ending. 111e nerve
movement).
fiber of a recep tor is composed of its axon and rnyelin sheath, if present. All
A-beta fiber A wide-diameter, tactile nerve fibers fall into a m yeli.nated class ca lled fibers, whi ch have
myellnated sensory nerve lber that
transmits signals from mechanical relatively wide diameters that permit very fast neural conduction. llle fo ur
stimulation. populations of tactile receptors that a re found in th e hairless (glabrous) skin
on the palms are s hown in Fig me 13.2 The n erve fibers of the various types
glabrous In reference to skin, lack-
ing hair.
of tactile receptor are assumed to terminate in different expanded endings.
You may be s mpri sed to lea rn that the specific kind of expan ded ending for
Meissner corpuscle A speclaUzOO ead1 type of ner ve fiber has p rovoked consid erable controversy among tactile
neive ending assodatoo w1t11 fast-
adaptln;i (FA I) fibers that have small scientists for over a cen tury n ow, bec.:i use observin g the physical connections
receptive fields. d ireetly h as proved ei<\J'<\mely d ifficu lt.
Merkel cell neurile complex A TI1e expanded enclin!;s f the t n:lifferen f popttlations of tactile fibers in
specialized nerve ending assoolated the hairless s kin of the hand are named after the anatomists who first d escribed
with adapting (SA ij fib€fs that Meissner corpusc les, Merkel cell neurtte complexes, Paclnian
have smaU receptive fields. puscles, a nd Ruffini endings. The endings of Meissner and Merkel receptors
Paclnlan corpuscle A specialized are at the junction of th e epidem1is and dermis, whereas the Pa d nian
naive ending associated fast- and Ruffini receptors are embedded more deeply in the dermis and underly-
adapting (FA II) fibers that have large ing s ubcutaneous tissue . ll1e fiber of a tactile mechanoreceptor ex tends all the
receptive fields.
\Vay from its terminal at the expa nded ending to the d orsal root of the spinal
Ruf11nl ending A specialized nerve cord, and it can someti mes be very long (s tretching fro m the sole of your foot
end'1g associated 1,;t11 slowly adapting to yo ur backbone, for example).
(SA IO fibefs tl1at have large rooeptlve
fields. Tile fo ur types of mec ll.:1noreceptors can be independently classified accord-
ing to their adaptation rates and tl1e sizes of their receptive fields, as measured
from activity o f their tactile ne rve fibers. ll1ese two dimensions lead to a second
set of labe ls for the mechanoreceptor types, lis ted in Table 13.1 . E.ad1 type of
bctUe fiber is particularly sensitive to certain features of mechailiml stimulation,
rendering it s uitable for particula r types of hinctions, as shown in Table 13.2.
• SA l fibers respond best to stead y downward pressure, fine spatial
d etails, and very low-freq uency vibrations of less th an about 5 Hz.
TI1ey are especially importa nt fo r texture and patte m perception. Some
acti vities particularly d ependent on this touch channel indude rea ding
Braille rmd d etermining the location and orientation of the slot on the
hea d of a screw that we can feel but n ot see. \ Vhen a sin gle SA I fiber is
stimLilated , people report fee ling 11 p ressure.'1 These fibers are assumed
to terminate in Merkel cel l neurite complexes, but it is n ot as yet clear
whether the.se endings play a direct role in generating the afferent
response or V\-·hether, in contrast, they ftmction on.ly to enable the fiber
itself to develop response capabilities.
• SA 11 fibers in the skin (as well as in all fib rous tissues in the body) re-
spond to s ustained d ownward press m e, a nd pa rticula rly to lateral skin
TABLE 13. 1
Response characteristics of the four mechanoreceptor populations
Size of receptive field
Adaptation rate Small Large
Slow SAi {MerkeQ SA II (Ruffini)
Fast FA I (Meissne(,I FA II (Pacinian)
Note: FA I= faM-adaptlng typg t FA 11=1ast-aoapt1ng type 11: SA I= slowly adapting type
1; SA 11: slowly adapting type 11. Tllet91TT1lnaJ ending associated w1th each t)o'?e or tactile
nerve Tibsr Is shown In parentheses.

TOUCH 393
TABLE 13.2
Mechanoreceptors: Feature sensitivity and associated function
Mechanoreceptor population Maximum feature sensitivity Primary function(s)
SAi Sustained pressure. very low frequency Texture perception
(< - 5 Hz) Patterntform perception
Spatial deformation
SAii Sustained downward pressure (low Finger position
sensitivity to vibration across frequencies)
Lateral skin stretch
FAI Temporal changes in skin deformation Low-frequency vibration detection
(- HO Hz) Stable grasp
Skin slip
FA II Temporal changes in skin deformation High-frequency vibration detection
(- 5CHOO Hz) Fine texture perception
stretd1, which occurs, for example_, w hen we grasp an object \¥hen you
reach out fo r your coffee cup, the SA Il fi bers help determine w hen your
fin gers are shaped p roperly for picking up the cup . S..A. Il fi bers ter mi-
nating in the folds of s kin around the nails convey forces on the finger-
tips as they in teract with objects (Birznieks et al, 2009). Scientists have
s ho\"'11 that when a single SA lJ fiber is s timula ted, people experience
no tactile sensation at all; for s timulation to be detec tabl e, more than
on e SA II fi ber m us t be stimulated. Althou gh SA Il fibers in the s kin are
assumed to terminate in Ruffin i endin gs, recent research h as question ed
w hether these expanded terrnin.."1.ls are as numerous as tra dition ally
believed.
• FA I fibe rs res pond best to low-frequency vibra ti ons from about 5 to 8230336 Amma Appa
50 Hz. lf your \."'Offee cup is heavier tha n you expected and begins to
s lip across your fin gers, this motion across the skin will cause jus t su ch
v ibra tions, a nd FA r fibers w ill help you con ect your g rip before your
coffee spills all over yo LL When a lone FA I fi ber is s timulated, people
rep ort .n. very localized sensation that they d escri be as 11wobble" or
"-flutter." These fibe rs are assum ed to term ina te in Meissner corpuscles.
• FA H fibers respond bes t to high-fr equency v ibrations fro m about 50
to 700 H z (the highest frequen cy tested to date). Such vibrations occur
whenever an object fir.s t rnakes contact wi th the s kin, as, for exa rnple,
w hen a m osquito lands on your arm. Such vibrations a re also gener-
a ted wh en a n object that you 're h old ing contacts another object, so FA
II fibers help you de tennine h ow ha.rd you 're tapping your pencil on
your d esk as you try to cram all this in fonnation into your brain. \Vhen
a s ing le FA [] fiber is s tirnulated, people repor t a 1nore d iffuse sensation
in the s kin th"'t corresponds to a "bu zz." These fibers have been sh own
to termin ate in Pad.n.ian corpuscles.
The four types o f m echa noreceptors are a lways working toge ther to
info rm us a bout every individual object we touch. TI1e SA I and FA I fibers,
in pa rticular, act a nalogously to cones and rods, res pectively, one affording
acui ty and the o ther sensitivity to low-inte11sity stimulation. K. 0 . Johnson
(2002) gives the ex.a rnple of opening a d oor wi th a key. Feeling th e shape of
your key in your p ocket requires the SA I (and maybe also the FA I) channel.
Shaping your fingers to grasp the key involves the SA I1 channel. As you insert
the key into the lock, your grip fo rce increases so the key does not slip,
than ks to you.r FA l cha1mel. FinalJy, your FA Il channel te lls you when the
key hos hit tlle end of the keyhole.

394 CHAPTER 13
FIGURE 13.3 Thermal n;iceptivity 10

functions. showing the response o f
warmth and cok:t fib€irs to diffarant t94n·
peraturoo. (,After Guyton , 100 1.) Cold
fibers
10 )) (J)
Nerve fiber temperature {°C)
11"1e majority of research on toud1 recep tors has con centrated on the hair-
less parts of the body, but m ore recent research acknO\vledges the importance
of h airy s k.i.n, such as the forearm, for tou ch perception. Understandably, this
area of s tudy has focused on manunals wi th more hair tha n humans have,
s uch as the cat. The mechanoreceptors in hairy skin exhibit a va riety of end-
ings and associated fiber types, as with hairless areas, but additional recep tors
are fo und near the follicles of the ha ir itself_ [n hrnnans, hairy ski n appears
to play a unique and important role in pleasant touch, to be discussed below.
THERMORECEPTORS Thermoreceptors locared in both the epidem1al a nd

thermoreceptor A sensory reoep- dermal layers of the skin (see Figure 13.2) infonn us :lbout cha nges in s kin
tor that signals Information about temperatu re. There are two distinct populations of thennorecep tors (Fig ure
changes In skin temperature. 13 .3): Warmth ftbers fire w hen the te mperature o f the skin s tu·rounding the
warmth fiber A sensory nerve fibers rises. Cold fibers (w hich outnumber war mth fibers by a ratio of about
fiber that fires Wien skin temperature 30:1) fire in resp onse to decreases in s kin te mperature:. The neu ral fibers that
Increases.
mediate cold and wa rmth include unmye lina ted,nnd hen ce reL1tive ly slowiy
cold nber A sensory na-ve fiber conducting, C fibers and faster-cond ucting, myelina ted A - delta ftbers. Both
that fires when skin temperature types are s malle r in diam e ter th an those coming from. the mechanoreceptors
dea;eases.
in the skin-the fi bers kno\.vn as type A-beta- and they lack
C ftber Anarrow·dlameter, unrny· specialized endings.
elinated sensory netVe Iba< that trarn-
mlts pain a'1d temperature signals. Our bodies nre cons tantly workin g to regula te thei r internal temperature,
so unde r nom1al conditions the skin is kept between 3o='C and 36°C (86°F and
A-delta fiber An lntermeclate-
slzed, myellnated sensory nerve fibe<
96°F), and neither cold no r warmth fibers respond much w hile s kin tempera-
that transmits pain and temperature ture rem11i ns within this range. If you bundle up in your long underwear and
signals. sn owsuit but then si t ins ide in front of the fire, your skin temperature w ill
probably rise above36°C, and your warm th fibers w ill begin to fire. If you then
take the snowsujt o ff and walk out into the s now, your s kin tern perature will
rapid ly begi n to fu ll, and as soon as it goes below 30°C, your cold receptors
will start firing.
TI1enn oreceptors also k ick into gea r when \Ve make contact with an object
thatiswarrne1· or colder tha n our skin. Objects in the environment are typica lly
cooler it is usually the cold fibers that te ll us about the object. For
example, s teel conducts h eat more efficiently than stone. Your cold fibers will
thus fire less rapidly and fo r a shorter period of time when you touch a s teel
object than w he n yo u to uch a stone object the steel object \.vill wam1
more q u ickly to match your ski n tempemh1 re). lf you've had prior expe ri-
ence \vi th steel and s tone, you can make use of the diffe rent them1orecep tor
responses to make this distincti on.
NOCICEPTORS Pain is the realm of to uch that hi.15 the dubious ho nor of be-
ing ho1n e to the sensations we like the least. We may find some vis ua l stimuli

TOUCH 395
revolting and some olfactory or gus tatory stimuli d isg usting, but of all the nociceptor A sensay receptor that
sen sations, it is pain that we take the m ost drastic actions to a void. Pain begins responds to painful Input, such as
with signals from nociooptors, touch receptors that h ave bare n erve endings extreme heat or pressure.
and that respond to various fo rms of or to s timuli that have the c tacttle {Cl) afferent A narrow-
potentia l to darn.age tissue (including extreme skin tempe ratures lmver than dlarneter. unrnyellnated sensOfY nffve
fiber that transmits signals frcm pleas-
15°C or higher than 45°C). (Pain :receptors are also folmd in internal organs,
ant touch.
but we ·will confine this d iscussion to toudt.)
Like thermoreceptors, n ociceptors b ck specia li zed endings and ca n be
div id ed into two types by their nerve fibers. f\·1yelinated A-delta fibers respond
pri1narily to s trong pressm e or heat, and unmyelinated C fibers respond to
intense sti mulation of various sorts: pressure, hea t or cold, or noxious chemi-
cals. Many painful events seem to occu r in two s tages: a quick, sharp burst
of pain, followed by a throbbing sensation. These tvi.ro stages may reflect the
onset of s ignals firs t from the A-delta fibers and then from the C fibers (Price
e t al., 1977).
It might seem tha t pain perception has no upside, but consider w ha t would
happen if \Ve had n o nodceptors. We would n' t be able to sense d angero usly
sharp or hot objects. Lacking alarms, we might soon lack fingers! Som e diseases,
sud1 as Hansen's disease (lep rosy) and diabetes, a re cha racterized by the loss
of pain sensation and provide rea l-life exa mples of the consequen ces. TI1ecase
of "Miss C,'' reported by Melzack and Wall (1988), sh m."·s what can happen to
people born with insensitivity to pain Not only did MissC lack pain sensa tion,
but she did not sneeze, cough, gag, or protect her eyes re.tlexively. She s uffered
childhood injuries from bu ming herself on a radiator and biting her tongue
w hile chewing food. As an adult, s he developed p roblem s in her joints that
were a ttributed to lack of discomfort, for example, from standing too long in
the sa me position. She died a t age 29 from infections tha t cou ld probably have
been prevented in someone w ho was alerted to injury by painful sensations.
PLEASANT TOUCH RECEPTORS The tradition al way o f classifyin g different

bodily sensations, introduced at the beginning of th is chapter, is in terms of
tactile, therm..'1t pain, and itch experiences. Collectively, these classic sensations
are known as d iscri minative toud1. More recently, however, scientists tmcovered
a fifth component th.,1 t they named 11pleasant'' or 11emotional" touch (McClone
et a.l., 2007). The em otional p roperties of nonpainful bod ily toud1 appear to be
m ediated in large pa rt by· a class of tmmyelinated (and thus, relatively s low)
peripheral C fibers knmvn as C tactile afferents (CT afferents for sho rt) tha t
a.re not rel;;;ted to either pa in or itch. This type of C fiber preferably responds
to mechanical stimulation in the form of s lowly moving, lightly applied forces
(like petting!). The adaptation rate of CT fibers is intermediate between those
of the myelinated SA and FA mech anoreceptors,
Isolating attributes o f the pleasant touch system is rather difficult because
ordinarily stimulation of the CT fibers also induces responses from the my-
elinated A-beta fibe rs that respond to general mechanica l stim ulation.. This
isola tion has been made possible, however, by studies of individuals wh o lack
A-beta fibers because o f a rare disorder. One s uch indi vidual, G L, feels n o
sensation of touch below the n ose and cannot feel pleasant to ud1 either, 'vhen
stimulated on hairless skin. Yet when stroked on hairy skin \vi th a bn:1s h.. s he
can detect and coarsely loca lize the source, which s.h@ vagu:eJy Rleas.."mt
(Bjornsd otte r et al., 2009; Olausson e t al., 2008).
Researchers believe that CT afferents are loca ted only in ha iry s kin . It has
been suggested that these mechanical uni ts form part o f a neural s ubsys tem
that integrates the body wi th i ts sensory and social environment. In everyday
behavior, the C T system could p rovi de or s upport emotional, homlonal, and

396 CHAPTER 13
FIGURE 13.4 A muscle spindl9

emb9dded In main (ex.trafusal) muscle
fibers contains inn'1ir Qntrafusaf) fibi9r s.
VVhen the inner fibers contract, a sen-
sory response from the spindle Is sent
back to the central nervous system,
corwe)'ing informaticn about muscle
and thus. rQQu1ating muscli9
tension.
behavioral res ponses to skin-to-skin contilct '"'·ith conspecifics, s uch as being

s t roked on the back of the ha nd, arm , or fa ce by a g irlfrie nd or boyfiiend.
There are indications that pleasa nt bodily contact promotes e ndorphin a nd
oxytcx:in responses, which contribute to feelin gs of well-being, confidence,
and calmness. Pearhtlness can be l'e1.1 uced in rats by h1ctile s timula tio n s uch
as strokin g wi th a bnish, and tou ch in human infan ts
has similarly been fo und to have beneficial effects (T. M. Field e t al., 1986).
Kinesthetic Receptors
kinesthetic Refernng to perception l.n addition to the tactile m echan oreceptor s in the .skin, yet other types o f
Involving sensory rnechancfeceptcrs In mechanorecep tors lie within muscles, tendons, and joints. These are collec-
muscles. tendons, and Joints. tively refe rred to as kinesthetic receptors, and they play an important rol e
muscle spindle A sensory recep- in sen.sin g where our limbs are and w hat kinds of moveme nts we're makin g
tor locatoo In a muscle that senses Its (C lark and Horch, 1986i LA. Jo nes, l 099). The a ngle for med by a limb a t cJ
tension.
joint is perceived primmily through muscle recep to rs called muscle spindles
(Figure 13.4), w hic h convey the rate at w hich the muscle fibers are chang ing in
length. Recep tors in the tendons prodd e signals abou t the tens ion in muscles
attached to the tendons, and recepto.rs d irectly in the joints them selves come
into play particularly wh en a joint is bent to an extretne angle.
TI)e importance of kinesthetic receptors is graphically illus trated by the
striking case of a neurological patient named Ian Waterman (read Pride and
a Daily Marathon [1991] by Jonathan Cole for m ore about th is in teresting in-
dividual). The cutaneous ne rves that connecte d \ Va terman's kines thetic an d
o the r mechano recepto rs to his brain \"Vere des troyed by a v iral infection when
he was l 0 years o ld. Lackin g kinesthetic senses1 \Vatem)an is n ow completely
d ependent on vision to tell him about the p ositions of his limbs in space. lf the
ligh ts are turned off, he canno t tie h is sh oes, w alk up or dow n s tairs, or even
clap his hand s, because he ha s n o idea where his hands and feet arel Ca ught
in an eleva tor when the li ghts '"'ent out, he \ '\'as w1able to remain standing
and could n ot rise again until the illumina tion returned (For addition.ti de--
tai ls about Waterman 's d)al lenges, n <:t the arru · gtit'grt!e to which he has
compensated for his lack of kines thetic recep tors, see We b Essay 13 .1: Llvtng
without Kinesthesis.)
From Skin to Brain

Initially the axons of variotts tactile recep tors are combined into single nerve
tnmks, in much the same way that retinal axons converge in the optic
nerve (see 0 1apter 2) and coch.lear hair cells converge in the au ditory n erve (see

TOUCH 397
Chapter 9). This analogy omits important differences, however: First, whereas labeled lines A theory of sensory
there are only two optic nerves and two auditory nerves, there area number of cod ing In which each nerve lbe< car-
somatosensory nerve trunks, arising in the hands, arms, feet, legs, and other nes a particular stimulus quality.
areas of the s kin. Second, the tactile ne.rves mus t carry their messages consider- dorsal hom A reglm at the rear of
ably farther. Because the receptors for sights, sounds, tastes, and smells are all tt-e sp'1BI cord that reoowes Inputs
located in the skull, the pathways that deliver information from these receptors fran receptors In the skin.
to the brain are fairly short. Touch messages, on the other hand, must travel as somatotyplcal H1Mrg noonal
far as 2 meters to get from the skin and muscles of the feet to the brain. somatooensa:tlon.
To cross this distance, the information must move up through the spinal
cord. The cord is far more than a handy transmission pipe; its neural structure
makes important contributions to the perceptual outcomes of touch. As described
thus far, the nerve fibers arising from the skin might seem to constitute labeled
lines; that is, each fiber type codes a particular touch sensation. Beyond the
peripheral neural layer, however, the lines become interconnected, making it
possible for complex patterns to emerge. The s pinal cord is the site at which
this cross comm unication is initiated.
The cell bodies associa ted w ith the tactile neural fibers a re bundled into
a cascade of ganglia, lumpy bodies that lie jus t outside the cord. The axons
themselves enter the spinal cord a t the dorsal horn, which is toward the back
of the spinal column. The horn is organized into multiple layers, or laminae,
as s hown in Figure 13.5. Every skin mechanoreceptor projects into the horn,
although that m ay n ot be its only projection. The inputs to the cord are orga-
nized somatotyplcally. Somatotopy is analogous to the topographical spatial
representation of events on the retina found in vision (see Dlapter 3); adjacent
areas on the skin are ultimate1y-corutected. t'o adjacent areas in the cord. What
may be surprising is th.a t these inputs constitute only a small part of the neural
structure o f the dorsal horn1 as most of its neurons lie entirely within the spinal
cord and serve as local connections, like the intermediate layers o f the retina.
In a review of dorsal hom function, Abraira and Ginty (2013) propose that
it jg this connectivity that provides the sense of touch with its rich canvas of
effects, from caresses to p okes.
h=u -.'I { /
FIGURE 13.5 NeuraJ projections

to the dorsal horn of the spinal cord.
baS9d on th9 cat. rat, and monkay. The
horn ls divided into layers (laminae-o f
which layer II is the subs tantla gelatl-
nosa). Afferent typgis terminat9 with in
the horn In a regular pattern, as shown.
(From Todd. 2010.)

398 CHAPTER 13
(a) Spinothal.amicp.:ithway
Toce:rebr.1-I
Po,-··d- \)
(b) Dorsal column- medial lemniscal p.1thway
gyrus of
oottex
spinothalamic
tract
temperaru1-e Rb.?rsfor pressure,

and pain vibration, position sense
FIGURE 13.6 Pathways from skin to

cortax. ta) Spino thaJamic pathway. (/J)
Dorsal column-medial lemniscal p ath· Once in the s pinal cord, to uch infor matio n proceeds up ward toward the
Wa-J. (After Levine, 2000.) brain \.;a two m.ajor pathways, as s hown in Fig ure 1 3 .6. The evohJti onMily
olde r splnothalamlc pathway (Figure 13.6n) is the s lower of the two nn d
ca rries mos t of the info rma tion fro m thermoreceptors .md n ock-ep tors. The
dorsal columi>-medlal temnlscal (DCML) pathway (Figure 13.6b) ind>1des
M d er-d iameter axons and fewer syn apses and therefore con veys infonnation
more quickly to the brain. Tactile and kinesthetic infonnation carried a long this
pathway is used for planning a nd execu ting rapid m overnents, '"'·here quick
feedback is a mus t. The DC.tvfL pathway not only includes fibers ascending
directly from the mechan orecep tors, but also is densely popula ted by fibe rs
&om neurons o rigina ting in the d orsal hom , p resumably conveying the output
of the ne ural acti vity w ithin the spinal cord.
Neurons in the DCML pa thway first syna pse in the cuneate and gracile
nuclei, near the base of the brain (see Figu re 13.£..b). Activity is then passed on
spinothalamic pathway The roLrte to neurons that syn apse in the ventral posterior nude us of the thala mus. Recall
frcm the spinal cord to the brain that from Chap ters 3 and 9 that the visual and auditory pa thways also pass through
carries moot of the Information about
the thalamLLs, each syn apsing in its 0\-\<11 modali ty-specific nudeus. Because
skin temperature and pain.
th.is p or tion of the brain is large ly shut do\.\rn when we are asleep, the brain
dorsal column-medial lemniscal d oes not register {and therefore does n ot attempt to respond to) the relatively
(DCML) pathway The route from
gentle toud1 sensa tions tha t occur, for ex:a.m ple1 when \"'"e roll m·er in our s leep).
the spinal cord to the brain that carries
signals frcm skin, muscles. tendons, From the thalan1us, much of the touch information is carried up to the cortex
and joints. (Figure 13.7) into somatosensory area 1 (8 1), located in the parie tal lobe jus t
somatosensory area 1 (S1) The behin d the postcentral gy rus. In ternlS of its posi tion in the transmission chain
pr1mary receMng area for touch In the fro m the periphery to the brainrSl is analogous to Vl in vision (seeCh apter 3).
cortex. With respect to.responses of cells in 51 to spatial pattemsr the analogy is closer to

TOUCH 399
SI FIG URE 13.7 Primmyoo;natos"10·

sory roc;eiving areas in the brain. S1
includes Brodmann areas 1 , 2, 3a.,
and 3b, Areas 6 and 7 are immediately
postarior to S 1. 82 lies 'With in tha lateral
sulcus.
somatosensory area 2 {S2) The

secondary rooelVlr-,g area for touch In
the <X'.fte.x.
the retina. When monkeys scan raised-dot patterns wi th their fingertips, recep- somatotoplc Spatially mapped '1
tive fields found in SI tend to talce the form of region with adjacent the somatosensory cortex In corres·
inhibitory areas, rese mbling the ON-center retinal gang)ion cells {DiCarlo, IXf)dence to spatial events en the
skin.
Johnson, and Hsiao, 1998). Neurons in Sl commlulicate with somatosensory
area 2 (S2), \·vhlch lies in the upper ba.n.k of the lateral sulcus, a nd vd th other homt.11culus A mapllke representa·
lion of reglors of the body In the brain.
cortica l areas. The m oto r areas of the cortex, w hi ch control m ovements of body
parts, are located just in front of the central s ulcus. This adjacency enhances
conununication behveen thesmnatosensory a nd m o tor control systems. FIG URE 13.8 The sen soryhomunc u·
Touchsensa.tions that result from stimul ation of the skin arespn tially repre- lus, stowing brain regions that respond
to stimulation o f different parts of the
sented in area Sl 1 and to.some extent beyond, somatotoplcally. Similar to the
body. (a) Multiple maps exist in primary
skin's projection to the spinal cord, adjacentareas on the skin luwea rollftection,
to adjacen t areas in the brain (Figure 13.Sa). As a re.su it, the somatosensory
cortex is organized into a spatial map (or as we see below, multiple maps) of one withh S2. A schematic o f the
the layout of the skin. Each m ap h as been ca lled a sensory homunculus (plural relative distribution of body parts h S1,
os orlgin<llly dartvsd by P"'1ield and
_,,,.,,.,.., (1 950).
(•) (b} Soma!osensory map
Prnn'''l' { D Am I
somatosen.so1y D Area 2
oortex (S1)
D Area3
Secondary L• g
somatoSli!nsor}' • Foot
cortex(S2)
Toe;
Genitalia
Thurnb- - - L----i--i'1',
Eyes----!;;= ± :
Nooo - --;..- " -
Face
Chin - Lateral Medill -

400 CHAPTER 13
body Image The lmpresslm of our homm1Culi) (Fig ure 13.Bb) and actually has a hvin ho munculus because the re
txxlles In space. are corresponding spatial maps in the left a nd right h e1nisphe res.
phantom limb Sensation perceived The sensory hornw1 culus is derived largely from the work of Canadian
fran a physbally amputated limb of neurosurgeon Wilder Penfie ldf who charted the somatotopic map with the
the body. aid of pa tients und ergoing brain s urgel'y to allevia te epilepsy. Because there
are no pain re1..""eptors in the brain, the patien ts did n ot need to be anesthetized
and could rem illn responsive. During the operati on, Dr. Penfield systemati-
ca lly stimulated different parts of a patient's somatosensory cortex with an
electrod e. As the probe was moved from one location in Sl to another, the
patient reported feeling sensa tions in the anns, legs, face, and so on. The cor-
respondence between the s timulati on and the sensation gave rise to a map
o f the body in the brain (see Web Activity 13.3: The Sensory Homunculus).
In fact, brain contains multiple sensory maps of the body. Separate maps
are now known to exist in the differen t subareas of 51, an.d additiona l maps
exist in secondary areas, as shown in Figure 13.&.
Like the retinotopic map in area VJ (see Figure3.16 in Chaptel' 3), Penfield's
.somatotopic m ap in Sl is distorted. The thumb, for example, g rabs a big
piece of real estate relative to its size. ln con trast, sensations fr om the leg a re
processed in a relath·ely small portion o f Sl . ln the visual sys tem, the foveal
is overrepresen ted in Vl (cortical magnification; see C h apter 3) because
there are many more photoreceptors in the than in peripheral parts of
the retina . Similarly, a Larger d 1unk of Sl is d edica ted to p rocessin g infom1a-
tion from the Bps than from the neck bec,1use tactile receptors :lre much more
heavily concentrated in the lips than they are in the neck.
1l1e distortion in the brain's m ap is ech oed in how people pe rceive their
own bodies, r their body Image, Fuentes, Longo, and Haggard (2013) had
8230336 people draw maps of their OV\>TI bodies on a computer, by clicking on locations
o f body parts relative to th e on-screen image of a head . People's body image
proves to be systematically distorted toward top-heaviness, w ith expand ed
shoulders and upper arms but red uced-size lower arms and legs (Figure 13.9).
The body image d1anges with the body it5elf1 as was s how n by the results
A\!erage AV'2'r<l£o?
of a s urgical procedure to eJongate arms shorten ed by dwarfism (Cinunino
true body body image et :ri l., 2013}. Wi thin 6 months, the patient's body image of her upper lim.bs
chan ged from being shortened to being con gruent "'11th normal con trols, while
the lower portion of her body im age reni.:1-i ned und1anged . (See Web Activity
13.4: The Rubber Hand Illusion fo r an experience of displacing your bo dy
image, and see Web Essay 13.2: Body Image to see how the impression ""'e
have of our bodies in space [body image] is highly changeable.) l11e relatively
tight correspondence between body parts and areas of Sl can have unfortunate
sid e effects in cases of limb amputation. Lf an amputee's left ann is mis.sing.
obviously n o mechanorecep tors are sending tou ch signals from that arm.
H owever, s poradic activity can continue in the area of the amputee's right
51 corresponding to the arm, leading to the perception o f a phantom limb.
11
At times, patien ts may perceive their ph'1ntom limbs to be in lmcomfort,1ble
positions, leading to persistent (and very real) pain.
1l1e psychologist Vi laya nur Ramachandran made the astonishing observLt-
tion that amputees often report feelin g sensa tions in thei r phantom arms and
FIGURE 13.9 ShaP<> of the body ha nds w he n their faces or remaining limbs are touched (Figure 13.1 0)_ The
as determined from locations o f parts som ce of this soma to.sensory confosion can be traced to an idiosyncrasy in the
(dots) for pe::>pl9's truQ OOdy (left) and h omunculus. Note in Fig m e 13.8 that the area responding to the face is located
locations they report relative to the
(some\-vhat arbitra rily) adjacent to the area responding to the hand and arm.
h98.d, which form thSr body image
(right). Multiple maps are aver;:qed by
Apparently the hand and arm areas of Sl are, to some extent1 ltinvaded '' by
scaling each one r9atlve to the person's neurons carryin g information from touch recep tors in the face. However, other
body height. tA,ftQr Fuentes et al.. 20 13.) parts of the brai n Listening to the hand and ann meas are not fully aware of

TOUCH 401
these altered connections, a nd the refore they a ttribute activi ty in these areas to
stimulation from the m issing limb. (You can read m ore about R..'Ul1achandran's
fascinating s tudies on phantom limbs in Web Essay 13.3: Phantom Limbs.)
Projections from Sl form the basis for further analysis of objects and s mfaces
by the cortex of the brain. The results of some recent s tudies s uggest that, like
vision (see Chapter 4), the sense of touch may s how a di vision be hveen what
and where system s in higher corti cal renters. A patient s tudied by Reed, Caselli,
and Farah (1996) s howed an im.p ainnent in he r a bility to recognize objects by
touch (w/rnt), but s he showed no abil,il)' nw ). Anotl1er
patient could locate -a nd manipul a te objects by touch wi thout recognizing
the m (Rossetti, Rode, a nd Boisson, 1995). Activation of the observed
w ith fMRI (see G1apter 1), h as been fo und in different areas, depending on
whether the task is to locate an object or to recognize it tactually. And , as in
vision, there is relatively more d orsa l activation for locating objects a nd more
ven tral activa tion for recognizin g objects (Reed, Klatzky, and Halgren , 2005;
Reed, Sho ha m, and Halgren ,2004) . You can lea rn more abou t the 1.oliat versus
where systems for touch by tapping into a lively deba te on this topic am ong
scien tists interested i.n the nemal circuitry tha t to object p rocessin g
(see Dijkem'li'ln a nd de Haan, 2007).
1l1us fur we ha\.""e described the pa th of discriminative touch to the brain;
pleasant touch appears to fo llow a different trajectory. Whereas primary so-
m atosensory cortex is activated by the physical aspects of the stimulus (e.g .,
by the pressure it produces on the skin or a drn:nge in skin temperature), the
CT system is associa ted \vi th another brain area, the insula, wh1d1 is thought
to pl1y a role in regulating the body and linking sen sory to em otional systems.
When G L, w ho has CT fibers but la cks input from tactile mecha noreceptors,
was s timu la ted by bmsh ing hairy skin, brain activation was found in the
FIGURE 13.10 Phantom lim bs may
insu la but not the sornatose11sory ilreas (BjOm sd otter e t a l., 2009). A no ther b9 on the facQ and stump
area of the frontal lobes kn ow n to be involved in emotion-the orbitofrontal subsequent to amputation. Amputees
cortex-is also activa ted by p leasant mech anical s timulation (S. T. Francis e t report feeling the arnp...rtatoo hand
al., 1999). As described in C hapter 15, both ins ula <md o rbitofrontal cortex wMn thgir fa09 or remaining limbs are
receive dired p rojections from the taste receptors, and as we w W find later in stimulated . S. Ramachandran,
1003.)
this chapte r1 the in sula plays a role in social connectedness and the relief we
get from scra tchi ng a n itd1!
FUR™ ER DISCUSSION of the orbitofrontal ex::> rt ex in the oontext of olfaJ-

tion and taste c an be found in Chapters 14 and 15, respectively.
Lest you conclude tha t th e nemal organi zation of our brain s for to u ch is
permanen t and unchanging, consider an experiment performed by Pascua l-
Leone and Ha milton {2001). These neu.roscien tislo; de prived no rmal, sighted
volunteers of visu al s timulation by h aving the m \•Vear a blindfold for 5 days.
On each day the volunteers participated in an tNfRI study d uring w hich pairs
of Brnille p atte rns were presented to the right index finger. Subjects were
required to jud ge w he ther the two patterns vvithin ead1 pair were the same or
different. Brain imagin g found th..i.t on the first day, the Braille task activa ted
only area Sl on the left side of the brain (whic h, because of the ne ural pathways
from the skin, is ac tivated by touchin g the right sid e o f the body). But as the
d ays progressed, the amount of activation d eclined in Sl whi le in creasing in
VJ. Apparently area Vl , w hich we thin_k of as d edicated to vision, took over
processin g the spatial pa tterns introduced through the sense of touch . Th is
cha nge in \ '1 was tr.msient: removing the blindfold resulted in a full rehirn to
the ne uTaJ hmctionin g that had been observed before blindfolding .

402 CHAPTER 13
neural plasticity The abillty of Such results revea l the ren\arkable neural plasticity o f the so1natosensory
neural circuits to undergo changes In system . Plasticity is a rec urring the me in sensory systems . \,Ve saw some thing
function a organization as a result of \;ery similar at the end of Chapter 3 in the d iscussion of visual developme nt
pre.,cus actMty.
and s tra bis mus, in w h ich abnorma l exp erie nce alters the wiring of the visual
substantia gelallnosa A jellylike system . Note that the exampl e vve're discussing h ere s hows th at plasticity is a
regbn of Interconnecting neurons In
property of the adult brain a nd is not limited to the immahue nervous syste m .
the dorsal horn of the spinal cord.
TI1e philosopher Wiiiiam Molyneaux ""'"as probably not thinking neural
gate control theory A doocrlptlon
of the pain-transmitting system that
plasticity w hen1 in 1688, he posed the question to John Locke as to whether a
lnocrporates modulating signals from blind person who was s uddenly a ble to see would recognize objects previously
the brain. known only by to ud1 . Medical interventions evenh1a1ly m ad e it possi ble to
answer this question by prodding sight to congenitally blind individuals . It
was d emon strated tha t although visual objects a re n ot connected immediately
to their touched equiva lents , only a few days of visual experience are n eed ed
to do so (Held et aJ., 2011). Thesmprisin g find ing th at the connec tion is m ade
so quickly, given long-term experience wi thout vision, tmderscores the plastic
nah1re of the perceptual system .
TI1e pathways from the skin to the brnin tell just one part of the s tory of
the tran smission of signa ls in touch. Downward pathways from the brain can
alter the sell.S.l horu; tha -iitfu1uJating the periphe ry produces. Some o f the m ost
s urprisin g effects o f these d ownward patlnvays rela te to the fee ling o f pain1
"''hich we discuss next.
Pain
We te nd to thin k of pain as a n inevitab le o f s tress on or damage
to om bo dies, flowin g from sensory levels to the conscious feeling of "ouch.''
1l1e scientific s tudy of pai n, however, reveals it to be a highly s ubjective srnte
with d istingu ishablecornp onents. Wha t we think of as p ain a rises at multiple
levels-sensory, emotional, and cogni ti ve (Price, interact to cre-
ate a conscious exp erience.
MULTIPLE LEVELS OF PAIN Pain sensa.tions are triggered by the nocirep-

tors (descri bed ea rli er in this chapter). Ne urons ca rrying n ocicep tive signals
arrive at the dorsal hom of the spin al cord in its outermost layers, particular-ly
the second, called the substantla gelatlnosa (see Figure 13.5). Neurons there
receive infom1ation from the brain, and they for m synapses w ith the ne urons
tha t are conveyin g sen sory infonnation from nociceptors to the brain (see Fig-
ure 13.6). According to the very influe ntial gate control theory (Melzack and
Wa ll, 1988), the bottom-up pain signals from the nociceptors can be blocked
via a circuit located in the s pinal cord . Neurons in the d orsal horn actively
inhibit pain trans mission, and what is transmitted to som atosensory areas in
the brain is the combined ou tput of pain excitation fr om the nociceptors an d
this inhibition. The inhibitory neurons in the dorsal horn receive input signals
from h vo quite disparate sources: the large-diameter A-beta fibers coming from
the skin, w hich respond to benign touch rather than pain, and the top-do\'\>"ll
pathways from the brain. Gate con trol theory getsitsname from the idea that
the trans mission of the pain acts like a gate tha t is pushed open by exci ta tory
pain s ignals but closed by inhibitory inputs.
Pa in signa ls arising a t Sl a nd 52 don't tell the w ho le s tory, h ovv·ever.
Imaging m e thods identify other a reas o f the brain that corresp o nd to the
emo tiona l aspects of painful expe rie n ces. In one sh1d y (Rainvill e et al. 1 1997)
(Figure 13 .11 lt participan ts \.\.o·ere hypnotized and their hands were placed in
luke\va rm or very hot water (activating the rmal noc::iceptors). The participants
"'"·ere someti mes told that the impleasanb1ess from the wnte r \Vas in creasing
or decreasing, and their brains were imaged dtrring th ese periods by positron

TOUCH 403
(a) (b)
FIGURE 13. 11 PET signals showing the effect o f hypnosis

on the brain , as obsE;<Ved by Rainvile et al. (1007}. (a) Primary
som atosensory cortQX (8 1) w as no t aff9c1:9d by thg SLQgQSt-
gd unpleasantness of pain (high on the left , low oo the rlglt).
p) The ante rior cingulate oortax (/'..CC, d rdOO) . hO\<VWQI',
showQd significantly d iffen;int activ.:itb n , dgpGnding on sug-
gested unpleasantness.
emission tomography (P ET; see C ha pte r 1). The primary sensory area s of the
cortex- 51 and 52-vlere ac tiva ted by the hot vva ter_,. but the s ug gesti on of
greater unpleasantness d id not increase the ir respo nse relative to the s ug-
ges tion of decreased unpleasanh1ess. In contras t, another area, the anterior
clngulate cortex (ACC ), did resp ond differentially to the two hypnotic sug-
gestions, by increasing or d ecreasing its activity according to the s uggestion
of increased or decreased unpleasantness. The researchers concluded tha t the
ACC processes raw sensory d ata from Sl a.nd S2 in s uch a way as to p roduce
cm e motiona l resp onse.
A t a highe r le vel s till, p ain can prod uce wha t Pri ce (2000) called "secondary
pllin affect," w hi ch reflects the inflltence o f cognition . Seconda ry pain affect is
the em otion al response associated w ith long-te rm suffering that occ urs w hen
painfu_l events are ima gin ed or reme mbered . For ca ncer pa tients
w ho face a second round o f d1emotherapy m ay remember the fi rst and d read
what is forth coming . This component of pain is assocfated l'Yith the prefrontal
cortex, an a rea concerned \.vi th cogn ition and executive control.
It ma y seem odd to assocfote pain with laughte r, but at least some o f the
resp onse people have to tickling seems to depend o n nodceptors (Zotte r-
man, 1939). And, just as signals fro m the brain can control pain perception,
they appear to come into play "'rhen we try to tickle ourselves. Self-induced
tickling no t only prod uces less la ughter, it also p ro duces less activity in the
som atosen.so ry cortex, beca use of can celing s ignals from othe r brain areas
that know w he re the tickJing stimulation is coming from (S.-J. Blakem ore,
Wolpert, and Frith, 1998).
Som e people may claim tha t itchiness is as grea t a p ain as pa.in itself. In one
extreme case, Mary Ellen Nilsen, over a period of a year after experiencing an
o utbreak o f shingles (the same virus th a t causes chic ken p ox), scratched he r
scalp so inte nsively that she broke tluough the bone and damaged brain tissue.
anterior clngciate cortex (ACq A
Pain and itch are cl osely connected in a nelUaJ sense as well as a p erceptual regbl of thl brain associated v.;th the
one. Som e itch fibe rs respond to both pain and itch, w he reas others, called p:irca ved unpleasantness of a paJn
pmriceptors, are itch -selective. The sep aration of pain and itch is shmvn by m ke sensation.
w hose itch neu rons in the spinal cord were e limina ted . They didn' t scratch, pretrontal cortex A region of the
even w hen injected w ith s tuff tha t drove norma l mice into scratching frenzies; brain concerned 'Nlth cog nition and
the a nimals' response to p ain, in contrast, was una ffected (Sun et al., 2009). executwe control.

404 CHAPTER 13
analgesia Decreasln;J pain sersa- As for scratching, d espite w hat you may have been told about 1'making it
tlon dunng ca1aclous experience. worse," it ben efidaUy decreases activation at itch sites as early as the .spinal
endogenous opiate A chemical cord (5. Davidson et al, 2009). At the leve l of the brain, scratd1ing regulates
released by the body tha! blocks the responses of the anterior dngul.1te cortex and the insula, w hich, you rn.ay
release or uptake of neurotransmitters recall, respectively respond to e1notion..1l pain and pleasant to uch (Papoiu e t
necessary to transmit pain sensations
al., 2014). And tml.ike tickling, w hich is mo re effective when d one by someone
to the brain.
else, scratching yourself is more pleasurable and itch-re lieving than being
placebo effect Decreasln;i pain
sensation when people think they're
scratched (Papoiu et a l., 2014).
taking an analgesic drug but actually
are not. MODERATING PAJN Pain experiences are the complex result of sensory signals
interacting with many other factors that have mode rating effects. Damping o f
pain sensations (without losing consciousness) is called analgesia. Resp onses
to noxious stimulation can be affected by analgesic drngs, of course, but per-
haps more surprising are the a ttenuating effects of anticipation, religious belief,
prior experience, or exci te me nt. Studies also highlight the importance of inte r-
p e rsona l and bro..1 der social influences on the em otional compon ent of pain .
Pain can be rnodernted to some extent by benign cotmterstiff1ulation- for
example, nibbing th e s kin near a stubbed toe. Thi s res ults from at
the level of the spina l cord ben-vee.n the large-d ia1neter fibers and the nocicep-
tors, as described by the gate control mo de l.
A mo re drns tic measure is "cotmterirritation " or "diffuse n oxious inhibi-
8230336 Amma Appa tory control"----extreme p ressure, cold, or other noxious stimulati on applied
to another si te distant from the source o f the pain. For exainple, pain fr om
electricall y s timulating a tooth can be reduced by noxio us s timulati o11 of the
h..i.nd (Motohashi and Um.ino, 2001). Painful s timulation can also s uppress the
se nsati on of itch, a ltho ugh you m ight prefer the itchiness. As with the gate
control model of pain regulati on , the explanation invo lves inhibitory interac-
tions in the s pinal cord, as shown in Figure 13.12.
TI1ere are m.'l.ny s tories of soldiers in battle w ho did not feel painfuJ wmmd s
until the stress V•ta.s over. The analgesic effect in su ch cases is probably caused
by endogenous opiates, dlemicals released by the bo dy that block tlle release
or uptake o f neurotransmitte rs necessary to transmit pain sensations to the
brain. Differences between indiv;duals with respect to pain resp on siveness
(tha t is, pain 11 thresho lds ") may reflect differences in their baseline levels of
these substances. Externally produce d s ubs tances s uch as morphine, heroin,
and codeine are similar in che mical structme to these endogenous opiates, and
thus they have si milar an a lgesic effects. O ther drugs, such as acetaminophen
and ibuprofen, a lleviate pain a t its source by counteracting chemicals th.,t
would oth erwise start the n ociceptors firing.
\Vhat about pain reduction in the nbse.nce of any 1' real '1 intervention at all?
The placebo effect is the redu ctio n o f pain when people think they're. taking
an analges ic drug but are no t actually d oing so. The subjective reduction of
Skin Spinaloord
lkh receptors
' "\ lnhibHo'J) '

lt<h
FIGURE 13. 12 Pain and itch stlrnula·

tlon are initlalty transmitted by separate Pain receptors ( ,/ ..' i
'
Mlabeled lines. n In the spinal ccrdt pain ,,. @' I
nQUrons conn9Ct to an itc h-inhlbiti ng Pain
neuron, with the result that painful stim-
ulation Inhibits itch. (After Ma, 20 10.)

TOUCH 405
pain by placebos is well d octune nted, a nd imaging of the spinal cord with
fMRI reveals that placebos actually inhibit nociceptive processi ng as ear ly
the d orsal horn (Eippert et al., 2009). Pbcebos can h ave positive effects as
;JS
well: imaging of the brain s of p eop le who self-adminis te red a n inert nasa l
spray that they \Vere told would heighten their responses to pleasant toud1,
sh o'"'"ed increased activation unde r pleasant touch in the sa me areas that were
attenuated by an..1.lgesic placebos under painful s timulation (Ellingsen e t al.,
2013). The location s of these effects e ncompassed em otion al areas as well as
sorna tosensory cortex.
Research has s hown tha t contact \Vi th those we love reduces brain activation
in a reas th at regulate our e motions and bodily arousal responses to the threat
of painful stimulation . In an fMRI s tud y by Coa n, Schaefer, and Davidso n
(2006), wom en experien ced the threa t of electric s hock under three conditions.
A '"'Om an could hold he r h usband's h and , the ha nd of a male wi th w hom she
was w1acquainted, or n o ha nd a t all. The ne ural pain resp on se was reduced
by tou ching a nother's hand, and thi s e ffect increased w ith the closen ess of
the relations hip . Broade r socfo.J influences on pnin have alSQ been fo und. For
example, reports o f pa inful s trains of the arm h-om tasks req uiring repeti -
tive m otion spread rapidly in Au stralia during the 1980s-like a contagio us
disease--but they were comrnunicate d by wo rke rs who did n othing m ore
tha n talk to one an other about their experiences. TI1e cross talk could have
h ad multiple effects in this exa mple, bo th heighte ning people's sens itivity to
their suffering an d providing a name for it.
PAIN SENSITIZATION Nociceptors provide a signal when there is impend- hyperalgesla An lrcreased or
ing or ongoing damage to the body's tissue. This is called " nociceptiveu pain. haghtened response to a
On ce damage has occurred, the si te can become m ore sensi tive, triggerin g the painful stimulus.
feeling of pain more read ily tha n before. This experience is hyperalgesla and
reflects an increased or heightened response to a no rmal ly painful stllnli.lus.
The resulting pain is called 11 inflamma tory,'1 and the heighten ed pain sensitiv-
ity usually goes away once the tiss ue heals.
Pain can also a rise in the absence of immediate tratumt, because of damage
to or d yshmction o f the ne rvous sys tem . The res ulting pain is called ' 1neuro-
p.:1thic." Some ne urop.:lthic pain reflects changes in the sensory fibe rs a t the
s kin that d o n ot n ormall y produce pain but n ow become pain inducers (a
phen omenon kn o\>\.'TI as allodynia); othe rneuropa thic pain arises from d-1anges
in the d orsal horn of the spinal cord. The n1ech.anisms by ""·hich neuropath ic
pain arises are increasingly understood at the rellular a nd molecular levels.
An important implication of this resea rch is that no single m edication will
alleviate all types of pain.
Tactile Sensitivity and Acuity

Now tha t \ve... ve covered hep lysfolo ca l substra te of the tou ch system, we
can turn to the psycho logical and psychophysical aspects. How sensi tive are
we to m_echanicaJ s timulation? Wha t are the limits on tactile acui ty in s pace
and time? Put a bit different! }'; w hat are thesmallest details th.at we ca n feel?
How Sensitive Are We to Mechanical Pressure?

To measu re the minim tun pressure that can be reli.'lbly sensed by a pa rticular
region of s kin, we need a way to present well-defined runmmts of pl'esstu e
over a nd over again. In the nineteenth cenhtry, rvfo.x von Frey (1852-1 932)
d eveloped an elegan t a nd simple way to d o thJs usi n g carefully calibrated
stimuli, including horse and human hairs. Modern researchers typically use

406 CHAPTER 13
nylon monofilaments {e.g ., fishin g lines) of va ryi ng diarne ters. TI1e s nl.:1.ller the
diameter, the less force the line applies to the s kin before it buckles.
To _replicate von Frey' s m ethod you rselt to uch different pa rts of your
s kin with a ha ir fro1n your head and a bristle from a hairbn1sh to reveal the
relati ve s kin sens itivity to these h vo different forces. \Vi th the th itm er ha i1·1
you \1,ill probably find tha t you can feel it on the more sens iti ve areas, s uch as
your lips and perhaps some par ts of your hand. You probably wi ll not feel it
pushing into your thigh or upper arm . With the bristle, however1 you should
d iscover tha t your s kin is sensitive to mechanical p ressu re all over, but n o t
unifo rmly so. For example, if you explore the skin on the back of your hand,
you sh ould be able to con vin ce yourself that there are s pots of g rea ter and
lesser sensitivity (Geldard, 1972).
Data from a rnore controlled pressure sensitivity s tudy reve al that fo r both
men and women, thresho lds va ry across d iffe rent sites o f the bod y (a high
thresh old means that tha t part of the body is less sens itive). In general, tactile
p ress ure sensitivity is highest on the face, foll owed by the tr unk uppe r
extremities (arms and fin gers) a nd then the lower ex tremities (thig h, calf,
and foot) (1Neinstell1, 1968). The for males a nd females is very simib r,
excep t that \Vomen tend to be m ore sens itive to p ress ure than m en. Sensitiv ity
to tem iceratu re chnn es as we Uns to pain, also va ries ma.rkedly as a function
1
o f bo dy site. A pa rticular sensitivity to pain in the fingertips w as discerned
by hea ting the skin with laser pulses, inducing a pinprick sensation from th e
A-delta nociceptors (Mancini et al, 2013).
Another approach to measu ri ng sensi tiv ity is to ask w hat the s ma llest
raised element is tl'l<."lt weca n feel asanotherv.isecomplete lysrnooth surfoce is
passed over the skin. Like the sto ried p rincess w ho detected a pea under a pile
o f ma ttresses, we a ppear to be very sensitive to the p ressure differen ce ca used
by a raised d ot on a s mooth surface. At a criteri on of 75% d etection, people
can detect a d o t only 1 microme ter high-that's a millionth of a meter, or 39
millfonths o f an ind iJ TI1e d ot seem s to tri gger d etec ti on by the FA I recepto rs,
w hich also help us de tect and correct for a n object slipping within our grasp.
Eve n m ore impressive, when a texture that is ma de of many raised d ots only
a very small fraction of a m icrom ete r high moves across the skin, the result-
in g vibrati ons trigger the FA I] receptors deep w ithin the s kin, en abling us to
d istinguish the d ot fro m a perfectly sm ooth surface (LaMotte and Srinivasan,
1991). How sensitive the skin is meas m ed to be d epends on the meas uring
sens itivity; it appears, as much as the skill. Recent tedmiques have allowed
\.vrinkles scaled in nano me ters (nm = billionth of a meter) to be manufactured,
and the s mallest wrinkle heights that can be d etected by rubbing the surfaces
approxima te 10 nm (Skedtmg et al., 2013)l
People a re also sensiti ve tod1anges in p ressure over time---that is, to trlctile
,,_; brn.tion. Figure 13.1 3 show s absolute v ibrato ry threshold (the minimum
amount that a v ibrating stimulus d isplaces the s kin in order to be detected ) as
a function of the frequency presented to the fingertip (LOfvenberg and Johans -
son , 1984). In thi s s tudy, people could de tect the presence of vi bra tions from
below about 5 hertz (Hz =cycles per second ) up to about 400 Hz, the highest
frequency that was tested. Othe r studies have confirmed tha t people can d etect
vibr01ti ons up to 700 Hz (Verrillo, 1963), the h ighes t &equency tes ted to date.
Although people can de tect v ibra tions over a wide frequen cy ra nge, they
are not equally sensitive to all frequencies, as Figure 13.13 d early shows. Since
the various mechanoreceptor populations are sensiti ve to different frequencies,
the overall psychophysica l functi on for the d etecti on of vibrati on reflects the
contributions of different mechanoreceptor populations at diffe rent regions
along it T.;ike a look at the corresponding v ibration sens itiv ities of SA I, FA C

8230336 AmMa Appa
TOUCH 407
FIG URE 13. 13 R98Ults of an experiment measuring

the minirnal {thrW"lolc;t amplitude o f vibration at the fin-
gertip that people can de;toct, as a function o f the \'ibra-
t ory frequ911cy. As labelo9d on the y-axis, the threshold "'
g
-10
Is mMSured on a decibel scale reolati'le to a refer9nce
\'ibration (see C hapter 9). The exptairlmentally obtaiood -"
-ll
··.
function . portrayed by the sold line , is to .... _____________ ___ ____?".!
reflo9Ct the contributio n of thrM different mochanore-
CE1ptor (SA I, FA I. and FA II), which are
shown as dlfferent-oolored seg rngnts. Each population
-
g
-30
'-. \
.,,
Is assumed to control the threshold In the limited fre-
qL1ency range where it is m ost s9flsitille. For compari-
son, the dashed lines in each color show the vibratory
-40
"" ".
threshold for the corresponding mechancreceptor pop- "':g -SJ FA ii
ulation across the i;intire range of frequencies t ested. o;l
These lines canno t be obtained from the experiment
portrayed h9re but, rathoa<, are based on the rQSUlts of
neurophys.lological studies of sing le-unit mechanorn-
j
,_.
-<O /
ceptor responses. Researchers have proposed that SA -?U
I units m 9Cliate our threshold for vibrations below about 10 20 SO 100 200 500 lOCO
5 Hz; FA I fi bers. for frequenc ies from about 5 to 50 Frequency {Hz)
Hz: and FA II units, for frequencies aboV9 about 60 Hz.
(After lbfvenbefg and Johansson, 1984 .)
and FA II mech an orerep tor populations from less than 5 to 400 Hz, also shovvn two-point touch threshold The
in Figure 13.13. TI1e SA[ tmi ts would seem to mediate o ur absolute vibratory minimum distance at wl1lcl1 t wo stimuli
thresh olds for frequencies below about 5 Hz; the FA I fibers, for frequen cies (e.g., two slmultaneous tou:::hes) are
Just perceptible as separate.
from about 5 to 50 Hz; and the FA [I tmi ts, for frequencies above about 50 Hz.
How Rnely Can We Resolve Spatial Details?

Pressure de tection is the t.1ctile equivalent of detecting a spot of light, where
the basic question is whether you can see o r feel anything a t all. For the tactile
eq ui valen t of visual acuity (can yo u m ake out the pattern of w hat you see or
feel?), try measuring your twoRpolnt touch threshold. As the na me s uggests,
this is the smallest separation a t which we can tell that we' re being touched
by two points and not just one. This experiment is best d one parb1e r,
although it w ill work to some d egree if you test yourself. A comp ass (the
kind that draws circ les) is a useful stimulator, but yo u can use anything that
enables you to vary th e sepa ration behveen two points, s uch as a bent paper
clip. Pick one of your own o r your partner's bo dy regions and see if you can
distinguish beh>\o"een a sin gle point and h'Vo points. Then repeat the proced ure
with different separations of the tvrn poin ts (e.g ., 0.5, 21 and 4 centimeters) and
flt different places on the skin (Fig ure 13.14).
Like sensitivity to pressure, s patial acuity varies across the body, but the
extremities (fingertips, face, and toes) s how the highest ac uity. Res ltl ts from
a syste1na tic s tudy of n.vo-point touch thresholds in fern.Bies as a htnction of
body site d emons tra te this finding quite d early (Fig ure 13 .15), and the pat-
tern for males is very similar. More-sensi tive psychophysica l methods sh ow
that on the fin gertips we are of resolving a sepa rati on of only about 1
millimeter (nun) (Loom.is, 1981). 1l1ese resul ts place tactile acui ty somewhere
FIG URE 13 .1 4 Two·pcint touc h thresholds are det91"mined primority by the con·
centraticn and receptive-1iek:i sizes o1 tactile receptors In an area o f the skin. The
triangles represent point stimulators. and the circles represent the areas of skin that
would respond to a sing le stimulation .

408 CHAPTER 13
FIGURE 13.15 Th9 minimal separa-

tion betwMn to.-vo points net!ded to
peroeive them as separatG (two-point
touc h threshold} wtiQn the pcints are
applied at different s ites on thEi body.
(After W einstein , 1008.)
m u m m B • a m
Mean threshold {mm)
behveen vision a nd audition: it is worse than vis u<ll acui ty, but bette r tha n
au d itory sp a ti1JJ resolution .
Note the corresponden ce bet\V'eeQ th,4'_ij.'.ltteril-o f tv.1o-point touch thresho lds
ac1·oss the body in Figure 13.15 and the Jlalive distortion b f differen t body pa rts
in the sensory homunculus of Figure 13.8. This m atch is n ot coi ncidental. The
d etermina tion that two closely spaced points ins tead of jus t o ne are touching
yom skin requires that your bra in receive two separate s ign als. Th is means
tha t a t the s kin there must be a s uffi cient concentra tion o f recep tors, each wi th
a sma ll e nou gh receptive field tha t the two contact points will e lici t d ifferen t
respon ses. An additional constraint is that as the s ignals are sent to the cortex1
they mus t not conve rge. A chunk o f co rtical real estate la rge en ough to recei ve
the m separately is necessary. In short, the two-point touch threshold is low
(tha t is1 the ability to discern two very close p oints as separa te is high) only
w hen the de nsity of receptors is relatively high, the receptive fields are s rna lt
and cortical con vergen ce d oes n ot occur. Tactile s pa tial acuity thresh olds a re
medi ated by the SA [ (and possibly FA I) tac tile receptors, which have relatively
s1na ll receptive fields and hi gh recep tor d ensities. (See Web Ac tivity 13.5:
Two-Point Touch Thre sholds.)
Although u sefut the tmditiona l h vo-p oint touch test h as so me drnw backs,
as you may see w hen you try your O \Vll experime nt. Even if the two s timula ted
p oints feel like one, it is n ot quite the same as s ti m u.la ting the skin w ith a s ing le
continuou s contact. Therefore, asking p eople w he the r they are really being
touched by one point or 1:\vo yields quite a different answer than d oes askin g

TOUCH 409
if it Juls like o ne p oint or hvo, especia lly in sensitive areas s uch as the
fingertip. Alternatives that are more objective have been su ggested, including
judging whe the r a n edge has :a gap or h ow a grating (a s urface v\.'ith
alternating grooves and ridges) that is applied to the skin is oriented-along
versus across the finger (Craig and Johnson, 2000).
How Rnely Can We Resolve Temporal Details?

It is somew h at m ore difficult to perform your O\<Vn rneasures of how well
people can resolve fine te mporal differences in tactile stimulation. Various
psychophysical methods have been used to address th is question . A com-
1non method requires subjects to decide whether t\110 tactile pulses delivered
to the skin appear to be ei ther simultaneous or successive in time. With this
method, s ubjects C.."'tn resolve a temporal differen ce of only 5 milliseconds (ms)
(Gesche ider, 1974 ). Tou ch proves to be better than vision (w hich is 25 ms) but
worse th.'10 1'.lllliitiun (0. 1 (Sherrick.a nd Ch olewiak, 1986). As with spatial
acuity, you V\rill notice that touch ttJ.15 somew here between vision and audition;
in this case, however, audition is the best and vision the vrnrst.
Do People Differ in Tactile Sensitivity?

If you a re a s ighted p erson reading th is book con ve ntionally, yo ur tn ctile
sensitivity in 10 years is likely n ot to be what it IS today; a decline with age
is to be expec ted . Studies of blind people wh o read Braille re\'eal an exci ting
excep tion to tllis trend, however: Legge and colle"gues (2008) tested the abil-
ity of blind and sighted people to identify raised three-dot patterns de1·ived
from Braille symbols, displayed a t different scales. They found that s ighted
adul ts los t about l o/u per year in their acuity levels between their teens and
their 80s, but blind Braille read ers showed essentially n o age-related d ecline
(Fig ure 13.16). The re is an unfortunate implica tion o f this trend for people
w ho lose their s ight la te in life la sizable proportion of the blind p opulation).
Braille symbols to be identified

j .: h :. d ·: f :·
5.73
455
'.i 0.2 3 .61

§
to
g 0.1 287
f"-
-"
Standard Braille line
----------------------------- -, ------.------ --- . 2.28"'
s.
g
FIGURE 13.16 T-acti9 acuity versus age:
Acuity is measured by th9 accuracy of
2, sym bol identification w hen using thB index
ii
-0.1 '!Bl fingar, across different scales that differ in
logarithmic units. The left v ertical axis uses
a log 1naasure. w he<e the spacing of dots
-D.2
.... ..· .·.... ... ---.
,• 1.44
1.14
in a standard Braille character is set to
{dashed line}. On the right vertical axis, the
measure is the distance betwiaen dot cen •
ters within the idlfrtitled characters in m ii·
limeters (mm). The bands arrund the lines
10 20 5() 60 so 90 that are fit to the data repr9S8nt Q6o/o confi-
Age(years) dence intEllWls. tA,fti;ir Legga et al., 2008.)

410 CHAPTER 13
11'1e zero point on the left vertical a xis in Figure 13.16 is set to the standMd
spaci ng behveen Braille dots; once the necess.-1ry font size passes above it
(arotmd 74 years of age), fingertip sensi tivity is not sufficient to read Braille.
Another factor that affects your tactile sensi tiv ity is, apparently, your
genes. Both tactile spatial acuity and vibratory sensi tivity are m ore correlated
in identical than in noniden tical twins, pointing to a ge ne tic contributi on
(Frenzel et al., 2012). The same study asked whether gen es that influen ce one
sense organ thilt respond s to pressure, the skin, might also influence ilnother,
the ear. TI1ey found that hearin g acuity was significantly co rrela ted \Vi.th both
tactile spatial acuity and the threshold fo r wannth; moreover, patients who
had genetic mutation s lea ding to early deafness s howed reduced s pa tial and
vibratory sensi tivity in toud1. Thus common genes appear to be expressed in
multiple sensory forms.
Haptic Perc eption

With the physiology a nd basic psychophysics of the touch system now tmde r
our belts, we can turn to questions about how vve use the i.nfomi.ation gath-
ered by our therrn oreceptors, muscle spindle fibers, Pacini an corpuscles, and
so on.. The term haptic perception refers to perceptual processing o f inputs
from multiple sensory subsystem s, including those in skin, muscles, tendons,
and joints. Haptic peT'Ception is usua ll y active a nd information-seeking: the
perceiver exp lores the world rather than passively receiving it.
Perception for Action

As mentioned earlier, touch relies on action to get i.n.fom1ation from the world .
Expanding on this p oin t a bit m ore, we can say that touch is active in ty...·o
mmplernentary ways . Using our hands to actively explore the world of s urfaces
and objects outside o ur bo dies i sactio11Jor percepfio11 . Using somatosensation
to contro l our impressive ability to grasp and manipulate objects in a stable
and highly coordina ted ma1uier a nd to maintain p roper posture and balance
involves perception for action .
In o ur discussion of kinesthetic receptors, we talked about how the l05s of
these internal tactile receptorsJeads to a devastating inability to know (without
looking) where our limbs are positioned. Westling and Johansson (1984) sh owed
that mechan ore::eptors in the skin also play critical roles when we're interacting
'\>vi th objects (Figure 13.17). After these investigators anes thetized the skin on
volunteers' hands, the voltmteers could no longer maintain a stable grasp of
objects that they could see perfectly well. Feedback. from the mechanoreceptor
populations in the skin appears to provide crucial in.formation about '\Vhen
art object is about to slip o n the skin . You may know this yo urself fr om trying
to unlock a door w hen yo ur fingers are very cold!
haplic perception Knowledge of

Action for Perception
the world that Is der1ved frcm sensory Let 's now consid er the action-for-pe.rception of hap tic p rocessi ng. Leder-
receptors In skin, muscles, tendons. man and Klatzky (1987) coined the term exploratory procedure for a partku1i.r
and joints, lnvdvlng active way of feeling an object in order to leam about one or m ore of its properties
exploration. (Figure 13.18). Each exploratory p=dure · optimal for obtaining precise
exploratory procedure A stereo· d e ta il s about one or two specific properties. For ex.ample, to find out how
typed hand movement pattern used to rough an object is, the best explora tory procedure is la teral mo tion-moving
tolJOh ctJjects In order to perceive their
properties; each procedure Is best the fingers back and forth across the surface. This is the proce-
for determining one (or more) object dure tha t people freely d1oose when they '\-vish to learn about roughness, and
properties. research indicates tha t it is also the one that works best.

TOUCH 411
b c d e f g FIGURE 13 .17 Foroo and p osltion during lifting , grasp·
==IN'
.1
ing , and a c ube. Region a= grasping; band c =
lifting: d = holding; Q = lowe<ing; f and g = reieasing. The
Loa d fotce II 1I ,Il 11 I load fo rCGi newtons [N]) ls in the gravitational dir9Ction ,
I I I I I I the grip foroo is imposed by th.:t fing ers pinching the cube,
(N)
----r--------1---- ---- ---------t -r---- --- -a nd the position is the height rielative t o the table. The ratio
of grip for09 to load fcrc9 is s9t so as to just prQV&nt the
cube fro m slipping. (After Westling and Johansson, 1004.)
Grip force
(NJ
l VCt !t\!
Positipn
(mm)
i/ ! i
2 !\ ! i 11
i
i
I
r t t
o 10 11
To exp lain why each e.xploratory p rocedure is linked to a specific object

property, we mus t consi der both the neural s truchues that transd uce informa-
tion and the processes tha t operate on that information. For example, it ha.s
been shown that we adopt two different processes \•;hen judging the roughness
of coarse as opposed to fine s urfaces.KO. Johnson and his colleagues (K. O.
Johnson, 2002) haveshm-\rn that the activity of s low ly adapting rnechanorecep-
tors (the SA I fibers) is a princi pal basis for the perception of s urfaces that are
at lea.s t m od era tely roug h. These receptors are ten times ::\S responsi ve wh en
there is relative mo tion between the skin and the s urface as w hen the fingers
statica lly rest against the surface ""ithout motion.
As y01.H fin ger sweeps across a s urface, the pa ttern of force across the
receptors varies "Vvith the hills and troughs on the object's s urface, providing
a kind of spatial map of the variations in skin defom1ation th at is sensed by
the SA I fibe rs (and for the coa rser s urfaces, possibly also by th e FA I fibers).
Lateral mot ion : Pre;sure: Static conL.lct

texture hardness lemper,:iture
FIGURE 13. 18 Exploratory p-ooodurQS

described by Lederman and l<latzky, and the
Unsupported Enclosu re: g lobal Conto ur following: glob.11 ob)Bct properties associatr9d each p roa:i·
h olding: weight shape, \•olume shape, exa=t sh ap? dure. (After Lederm,_'1fl and Klatzky, 1087.)

412 CHAPTER 13
Tl1is map is passed on to higher-level ne ural structures, which integrate the

lower- level infomrn.tion into an overall measure of the amount of variation .
The brain then uses this n e ural meas ure to estimate the ro ughness of the
s urface. However_, additional research has furthe r shown that when surfaces
are very fine, the mechanoreceptors responsive to high-frequency vibrations
(the FA If fibers) appear to encode surface roughness, although stroking with
the fingers is s till needed to set ttp the vibration (see Hollins, 2002). H ollins
s uggested that scientific results o btained from the study of both coarse and
fine s urfaces support a dual-coding theory (as opposed to a single type of
coding) for hrnnan roughness perception.
Hollins, Bensn1fila1 and \ Vashburn (2001) used a very clever psychophysical
method knovv"ll as selective adaptation to obtain scientific s upport for this dual-
rod.ing 1l1ey reasoned tlldt if the high-frequency (FA II) mechanoreceptors
are responsible for en coding the perceived roughness o f fine s urfaces, then if
the fingertip is first selectively ":::ldapted '' (fatigued ) to a very intense high-
&equency vibration (e.g., 250 Hz)1 fine s urfaces s hould subsequently not feel
as rough as they d o \.vi th the finger:. In contrast, selectively
the fingertip to a lov1t-frequency vibratio n (e.g., 10 Hz) should
not affect how rough these same fine s urfaces subsequently feel, because the
mechanorecepto rs that are most sensitive to su ch low-frequency vibration (SA
l fibers, and possibly FA I fibers) d o n ot encode the roughness of fine surfaces.
11-ie results of this experiment nicely confim1 the prediction s, th us offering
evidence in frwor o f dual-coding m echanisms for roughness. Selective adaptation
has been more generally described as the 1' psychophysicist's microelectTode,"
in that this method offe rs a noninvasive complement to recording directly from
n eural fibers when sensory sys te m-in this case, som atosensation.
FURTHER DISCUSSION of selective adaptation in spatial vision can be

found in Chapter 3 starting on page 77.
Role of Fingerprints in Perception and Action

8230336 Amm
lf you examine your finger pads, you will no tice an elliptically s haped three-
dimensional p.."l.ttern o f grooves and ridges that spirals inward toward the center
of each pad s urface (a dose-up is sho'"',1 in Figure 13.19). Those patterns are
FIG URE 13.19 Scanning electron

rnlcro;;,raph o f the skin ridges on a
human index finger pad (fi91d o f vi.ew is
5 mm). {From Qu illiam , 1Q7B.)

TOUCH 413
yours and yours alone {even iJ you have a n identical twin). So w h at are they
good for? Long before the hit show CSJ beca me p opuJ ar, the field of forensics
used two-dimensional inked versions of these ridged finge rprint patterns to
narrow dmvn. the field of suspects in a cri m e.
More recently, however, scientists have prop osed that the finger pad ridges
1nay pby at least tv;o other va luable fun ctions in daily life as we manually
explore, perceive, lift, transport,. and manipulate objects. One is to enhance
the perception of fine s urface texh.u-e.s. Research .suggests that when finger
pad-like ridges are used to actively explore very fine surfa ce textu res, they
selecti vely a rnplify frequencies ranging from about 200 to 300 H z by a factor of
about 100, while filtering out other frequencies (Sch eibert et a l., 2009). However,
ne ither of these effects h as been observed in conjunction with coarse surface
textures. If you recall 01u earlier disclLssion of the two processes for the manual
perception of s urface roughn ess (fine versLLS C'08.rseL you w ill realize that the
hig h-freq uen cy selection and a mplification imple mented by the finger pad
ridges dovetails with the m axinum1 sen sitiv ity of the FA II m echanorecep tors,
whidi.have been implicated in the ne mal processin g tha t tmderlies the manual
perception of fine s urface texh ues. In short, the finger pads are ttmlng the signals
from exp lora tion o f finely textured s urfaces to the relevant n eural p rocesses.
Scier1tists ha ve long speculated that fingerpri nt ridges play a second role,
helping our fingers maintain a s tab le g rasp on objects by increasin g the amount
of friction behveen our ridged s kin and the object. In fact, these skin protru-
sions are often called " friction ridges .'' H O\·veve-r, Peter \ Var man and Roland
Ennes (2009} have challenged the validity o f this hmction by experimentally
shov..ring tha t ridged fingertip skin prod uces less, not more, friction during
contact b y reducin g the ski n contact area. But take a look a t Figure 13 .19,
shovi.ring a scan ning electron m icrograph o f the skin ridges on a hwnan finger
pad, w hich s uggests a different \ ..·ay tha t fin gertip ridges amid <lffect grasping.
Notice th e drople ts of swe.at that are prod uced along the tops of the rid ges
at short, regula r interva ls. Co uld m oisture (which was n ot addressed in the
previous: s tudy) help stabilize grasping by increasing the fr icti on be tween the
finger and the g ripped objed? Measuremen ts o f the s lip threshold (see Pigure
13.1 7) for different levels o f grip force and skin m oisture content s uggest tha t
moderate a mOlm ts o f swea t could fuc ili tate stabl e manipulati on of objects by
increasing fri ction (Andre, Lefevre, a nd Thonnard, 2()0;)}.
The What System of Touch:

Perceiving Objects and Their Properties
Ch apte r 4 described the p rocesses tha t unde rlie visual object recogni tion. \ \le
1
need somatosensa tion to control s imple actions such as s tanding or grasping

and to wan-i. of danger th rnu gh pain, but h ow rnuch v alue d oes the sense of
touch h ave as an object recognition system ? You know tha t touch a lone can
function quite well to identify objects if you have ever gotten out of bed in the
d ark to use the ba throom . An d the n ext time you get d ressed, try to keep your
gaze constantly focused on your hands as you button your sh irt. This si mple
exercise sh ould con\o;n ce you that even \vhen you ca n u se vision , you some-
times rely on toud1 to recogni ze objects an d their parts. On tli.e other ha nd ,
object recogni tion by toud1 has its limits: the designers of coins know that it
is imperative to be able to tell one fro m another in yom pocke:t, but the Susan
B. Anthon y dollar none tl1eless failed, because it fe lt too s imilar to the quarte r.
PERCEIVING MATERIAL VERSUS GEOMETRIC PROPERTIES People can

perform haptic object recognition very welL Klatzky, Lederman, a nd Metzger
(1985) asked people tq iden tif):'.eai;l1 P.flCX/ objects (e.g., a fork, a brush ,

414 CHAPTER 13
FIG URE 13.20 Examples of compa

rmn objticts that are easy to recognize
\'isually in two-dimensional form but ,
when raised for haptlc are
not easily recognized by t ouch. {After
Ledtfman et al .. 1000,)
and a paper clip) placed in their hands. Not only did people perfom1 a lmost
perfectly wi thout looking at the objects, but they also gen erally responded
in less than about 2 seconds. Ho wever, the information_used in haptic object
recogniti on is quite diffe rent fro m that used in vis ual object recognition. Con-
.sider the difference beh veen material properties-those that d o not depend on
the s tructure of a particula r object, like its s urface roughness-and geome tric
prope rties like size and s hape . In haptic perception, the observer is in contact
with the object being observed, so ma terW p rop e rties of the object {is it soft?
cold ? fu zzy?) ar e easy to perceive a nd pl ay a crucia l role in the recogni tion
prcx:ess. In vision there is n o physical contact, so thermal and tex hual proper-
ties of objects are much mo re difficult to perceive.
11-ierefore, the geometric prope rties of objects al'e the m ost imp ortant fo r
visual recognition. Indeed, sparse line dra\vings may be quite e.asy to recogn ize
visu ally, but they a re hard to recognize w hen presented ha ptically as raised
contours (Figure 13 .20). To d eterm ine the overall s hap e o f an obj ect h apti-
c.olly, we usually must explore the object by tracing along its contou rs \Vi th
our fin gers. Integra ting tactile info rma tion over time is possible but not very
efficient, w hich is w hy the ins tantly recognizable material p roperties tend to
be much more important in hapti.c recogniti on . (For more on th.is topic, see
Web Activity 13.G: Haptic Object Recognition.)
HAPTIC SEARCH As \'\'"e saw in 01apter 7, a ntm1ber of so-called preattentive

features in the visual d omain are p re.sum ed to be cri tical in the visual object
recognition process. These features can be identified by the extent to wh ich
they " p op out'' in a visual sea rch ta sk. For example, if you are sea rching for a
red object, you will be equa lly fast at finding it regardless of how rn.any g reen
objects are p resented along wi th it. TI1 is res ult imp lies that the /1 redness'' of an
object is available to recognition processes befo re a ttentional mechanisms ex-
amine the objects in the display a nd integrate the v ario us features of each one.
Does the sense of touch also support preattentive feah ue d etection ? To find
out, Led erman and Klatzky (1997) cons tructed a set of s urfaces somewhat like a
tactile s lot machine with one key for ead1 of six fingers. The s ubject's right h.,1.nd
is s hO\vn in Figure 13 .21 a . Stimulus patches were m ounted at'ound th e plana r
edges of ead1 of six stimulus wheels. On e..1ch trial, the '"·heels were rotated until
the d esired stimulus patches w ere facing upw ard to fo rm the h aptic d isp lay.
1l1e entire display was th en m oved up to contact different combinations of
the middle three fin ger tips of e..1ch hand . Us ing this a pparahLS, Ledennan and
Klatzky found that a number of haptic feahu-es d o indeed pop out. As Figure
13.21b sh ows, p articipa nts in these expe riments were just as fast at de tectin g
a rough s urface when there was n o sn.10o th s urface in the tactile " display" as
when there were a s many a s five s mooth s urfaces. Sim.ila rl}'i a hard s urface
p opped out of a grm.ip of soft surfa ces, a cool surface p opped out of warm
s urfaces, and a s urface wi th an ed ge popped out of perfectly flat s urfaces.

TOUCH 415
(b)
90J
OSnV>O&..timub
ORou/#>•timuli
700
'forget present Target absent

&. 500 (slope =-1) (>lop• =9)
I
Number of fingers stimulated
FIGURE 13.21 An experiment investigating whett1er touch supports preatt€<'rt1ve

(c) feature detQCf.lon. apparatus used to display targets to th& fingoartlps. The
rotating drums bring either a stimulus patch or a cut out (no stimulus) t o the upper
surface, and then they rlse as a whole to contact ttw mldd!Q throo fn1gQrS of both
hands {only 0119 hand is sho wn). (b) The ftrne rgqulrQC:l to a rough
target among smooth distractors as a function of the number of fingers stlmu·
lated. (c) A three-dimensional version. wh9'9 a target obj9Ct (sphgre among
in the o9Xamptc;i shown) is detQCt:Qd among objects graspi;Kj In thQ hand Qnsat).
(Parts a b after Lederman and t<Jatzky, 1007: c after Plalsier, Bergmann Tlest. and
Kapp€1rs, 2000.)
Plaisier, Bergmann Tiest, and Kappers (2009) d eveloped a three-dimen.sional

version o f the h a ptic sea rc h p aradig m in w hich dus te rs of objec ts were
grasped in the ha nd, :as sh m . .n in Agure 13.21c. Objects w ith different shapes
had corresp ondin g differences in the loca ti ons of ed ges, s urfa ce curvature,
surface a rea, and height-to-wi dth ratio. By means of a single grasp, subjects
exh ibite d highly efficient search w he n ta rgets and dist ractors diffe red in
the type of sh apei for exam ple, a cube grasped within a handful of spheres
ten ded to p op out.
H mvever, not every ha.ptic differen ce s upports efficient search . For exa mple,
response times increased \o\ith the number of d istractors w he n the tas k was
to find a target with a horizontally oriented edge among d istractors having
verti cal edges. Note that horizontal targets do p op out of vertical dis tractors
in visual sea rch tasks. This d istinction fits nicely wi th the previous observation
that h aptic recognition relies extensively on m aterial properties, but that the
tac tile sys te m d oes not appear to be set up to efficien tly differentiate object
contours by their s patial layout.
PERCEIVING PATTERNS WITH TH E SKIN Even if pa ttern percep tion by

touch is not terribly e fficient, it can be d one, especially if the p..'lttems are sma ll
e nough to be perceived by a s ingle finge rtip . Loomis (1990) sug.gested that, to
some extent, touch acts like blurred vision w hen the fin gertip explores a raised
pattent. He tes ted people's ability to identify a set of pa tterns induding Braille
symbols, English and Japanese le tters, and geometric for ms--a few of which are
shown iJ1Figure 1 3.22. Sornetimes the pa tterns '"'-ere presen ted to the finge rtips
as raised elem en ts. Oth er times they were presented visually behind a blurring
screen that rnatched the resolution of th.e eye \·vith the more lim ited acuity of
fingertip s kin . Interestingly, Loomis fo und very s imilar patte rns of \oisual and
tactile confusion errors-that is, responses in w hich one pattern -....·as confused

416 CHAPTER 13
FIGURE 13.22 Sel9cted charac- : .. •: ·. : • :: :.

tEf recognition sets used by Loomis.
induding Braill9 (top). Loomis,
1900.) ABCDEFGH KLM N OPQRSTUVWXYZ
DDDDDDDDOOODDDDDDDDDDDDDDD
for an other. This findin g s uggests that a conunon 1 d ecision process operates
on both haptically l'\nd visi.rnlly perceived patterns.
In the Braille alphabet, shown in the first row of Fig\tre 13.22, each letter
is formed by raising some of the dots in a 2 x 3 array. For the letter A, for
example, a single do t is in the top le ft posi ti on; and for the letter Q,
all dots except for the one in the lower right posi tio n are raised. This design
reflects a compromise behveen the s kin 's acuity and its "field of view," the
o f skin that ""-e can take in all a t o n ce. It would be nice t o include more
than six d ots in the array, but because of the s patial blurring im posed by the
s kin, denser patterns would be difficult to resolve and discriminate {remember
tha t two-point toud1 thresho lds on the fingertips are about 1 mm ). Sp reading
a grea ter ots a larger area would n ot work ei ther,
because then the pattern would extend beyond the fingertip. Unforh.mately;
people are unable to "read" more than one finger at a time, s uggesting that
our tactile fieJd of view is very narrow.
tactile agnosia The lnabllltyto Iden- TACTILE AGNOSIA Just as lesions in the temporal lobe can p roduce visual
tlfy objects by touoh. agnosia (see Chapter 5), lesions of u,. parietal lobe can produce tactile agno-
sla, an inability to identify objects by touch. In making a diagnosis of t.'lctile
agnosia, the ne urologis t n eed s to be able to eHmh1ate o ther possibilities. Is
the p roblem impaired m otor control, w hich would pre\•en t the exploratory
procedures need ed to effectively lea rn about an object's properties? Or mi ght
the problem be a higher-level cognitive dysfw1ction , s uch as a loss of access
to object nanles?
\Ve already described a patient who could n ot recognize objects by to uch
but could locate them. She had tactile agnosia with her right due to a
lesion in the left inferior parietal region of her brain, but the d efici t did not
extend to h er left hand. Reed and Caselli (1994) d ocumented that, altho ugh
the patient could not recognize objects s uch as a key chain o r a combina tion
lock w ith he r right hand, she co uld easily recognize these objects visuall y or
with her left hand, ruling out a genera l loss of knowledge about objects. O ther
capabiliti es were no rmal in both hands, including sensory threshold levels
and the movements wi th which objects were explored . The patien t could also
d iscrirninate between objects wi th different levels of weight and roughness
us ing either hand . And she could answer questions about the haptic p roper-
ties of named sud1 as whether an orange was harder than a n apple,
indicating that s he had the ability to remember and imagine how objects felt.
11-1.us, the patient could acqti.ire information with her impaired hand about
an object's properties (e.g., its weight and roughness), and she ha d intact haptic
knowled ge about objects she had encountered in the past. \Nhat s he lacked
was a connection between these tv>.·o components of object identification. That
is, ei ther she was unable to integra te the perceived prope rties into a cohe rent
object representation, or she was tmable to m atch perceived representati ons
to stored representations in memory.

TOUCH 417
The Where Sys tem of Touch: Locating Objects

As vdth othe r sen sory m o dalities, knowing what a h a ptk stimulus n1 ight frame of reference The ccadlnate
be is only p art o f the perceptual problem. We a lso nee d to know where that system used to del ne locations In
stim ulus is located because we often w antto d o som ethin g '"'ith it. If you are space.
al re:1 dy to uching an object, you obvious ly know w he re it is (a tree limb tha t egocenter The oent€f of a refe ren::::e
you bump your head on is in the air; on e th at you sturnble over has fallen to frame used to represent locatlons rela-
tive to the body.
the ground). [f you a re n ot yet touch ing the object but can see it, your sense of
vision can work out where the object is a nd guide your reaching beh avior. But
w hat about groping fo r the s nooze button on your alann clock \·vh en your eyes
have n ot yet opened for the day? As mentioned alrea dy, there is evidence that
tou ch, like vision, has a specialized n eural p athway for dea ling w ith questi ons
of w here objects are located, as compared with knowin g w hat they are like .
HAPTIC OBJECT LOCALIZATION Like visual and auditory local iza tion , hap-
tic object loca!iZ.'l tion first requi res tha t we establish a frame of reference. For
vision, the center of the refere nce framt>-the egocenter-is 101."'ated nea r the
brid ge o f the nose, betv,.-een the two eyes; the auditory egocenter is a t a point
smack in the middle of the head (ben veen the hvo &'lfs). On e way to pinpoint
your haptic egocente r is to place your left index fin ger on top of the edge of a
d esk o r t.tble in ;:i nah.rr,11 p ositi on on the left side of your bod y, d ose your eyes,
and try to ma tch thi s location by placin g your right index finger on the bottom
of the desk. (Figure 13 .23 s hows a vers ion of this task u sing a stylus in eL\ch
ha nd instead of the index finger.) II you d o this ma ny times, you may find that
you consistently go too far to the left. Comrerse.ly, if you try to ma tch the loca-
tion of your ri ght index finger wi th your left, you ,. ,.m be more likely to e rr too
far to the right. A careful analysis o f errors in a task of this type led Haggard
(11) Topview
• • • •
Amma Appa
FIG URE 13.23 Loca.ting the haptic egoCtilnf.er. Q OQ

hand places a stylus on the target on the upper table
surfac e, and the other hand atto9mpts to m atch up under-
n eath the tabl9 in the oorresponding location. i;A.fter Hag-
gard et al ., 2000.)

418 CHAPTER 13
FIG URE 13.24 Haptic perception of

tabletop space. The two bars in thS
figure are arrangQd to f'1pr9Sent how a
typical experimental sub;=.ct wruld ori-
9nt tvvo b ars t o appear dose to •par.:il-
lol.' (Afte< Kappers, 2007.)
I
et al. (2000) to conclude tha t there is, in fact, no single, fixed &ame of reference
for the haptk perception of loca tions. In the case of the index fin ger reaching
task, the egocenter appears to be loca ted at the sho ulder of the am1 doing the
reaching. In other tasks, the egocenter may move to othe r positio ns on the body.
A lthough you mig ht think you 're pretty acc urate a t de termini ng h ow
external objects are oriented in hapti c space, research h..LS show n that peop le
make surprisin g ly large and systematic errors. Try the followin g task, which
is an f_n fo rmal demonstration of the experiment illus trated in Figure 13.24
{Kappers, 20C17). Haven friend place rn.·o pencils in front o t you wi thin reac h,
on e far off to yo ur le ft and the othe r far off to your right. Position your hands
in a natural way on both pen cils. The orientation of the lef t pen cil should be
fixed so that its len gth lies crosswise to the fingers of your left ha nd. 1lle right
pencil sh ould be set in a random posi tion benea th your rig ht hand. Ro tate
the right-hand pencil tmti l it feels paraUel to the left-hand pencil. Now have
a look a t h ow the hvo pencils are aligned with respec t to each other. Are they
physically parallel?
In the ach1al experim ent 2007), the; ro d at the RDSitions;
illustrated in Figure 13.24 were rotated to orienta tions differing by40 degrees
on average! Althou gh individ uals varied in their susceptibility tospatia1 errors
in this haptic paralle li sm task (errors for the positions shown ranged from 8
to as much as 91 degrees}, the average trends were quite consistent. Subjects
tended to judge w ha t was para llel by weigh ting their orientation judgments
""ith respect to two different spatial frames of reference: one was egocentric
(with reference to the body, and commonly centered on the hand); the other
was allocentric (with reference to extem a l space).
Tactile Spatial Attention

Although the last 50 years or so have produced n considerable amount o f re--
search on visual and a uditory D.ttentio n$only in the last decad e have scientists
begun to investigate the nahue of the p roc:.es.ses that tmde rlie tactile.a ttention.
ill this section we focus on tacti le s patia l (as opposed to nonspati..'11} atten tion.
FURTHER DISCUSSION of attention as it relates to visual perception can

be found in Chapter 7.

TOUCH 419
\Vhen people anticipa te being toud1ed in a pa rticular location, they can endogenous spatial attention
volu::ntarilv d irect their a ttenti on to that loca ti on. Attention th at is d irected A form of top -down Q<nowledge-
to the tactile mo dality in th is way is knov1.'n as endogenous (or top-down ) d riven) o:ntrol o f spatial attentlcn in
which attention Is voluntarily directed
con trol. In o ne s tudy (Spence, Pavani, a nd Driver, 2000), par ticipants were tO\ovard the site whae ttie cbserver
'1sked to indicate w hether a sus tained force or a series of pulses was de livered anticipates a stlmulus will occur.
to a fingertip (Figure 1325). The stimulated fingertip could be on the left or
exogenous spatial attention
the right hand. A visu a l p recue, in the for m of an arrow, predicted w h id1 A form of bottorn-up (stimulus-driven)
hand would receive the s timulation. If peop le could make use of this p recue spatial attention In which attention
to d irect a ttentio n to the predicted hand, it \.\.'as expected that they would be Is ral exwely (Involuntarily) directed
faster at d ecid ing w heth er the s timulus v,ras sustained o r pu lsed. And ind eed , toward the site at which a stimulus l1as
abruptly appeared.
this \Vas the case: the precu e sped up responses rela ti ve to a no-cue con trol.
Occasio nall)'i h owever, the p recue directed the participant's a ttention to the
w rong h and , and on s uch trials people resp onded m ore s low ly than in the
no-c ue cond itio n.
1l1ese effects are exactly an alogous to atten tiona l cuein g effects in \·i sion
and aud ition. Thus, we see th at tactile attenti on.1 li ke vis ua l a nd a udito ry
attention, is a limited resource that mus t be alloca ted in one way o r
You w W p robabl y h3ve rea lized tha t directin g one's a ttention endogenous ly
(vohm tarily) to a stimul us event is not the only way in w hid1 people con trol
spatial a ttention. It is also p ossib le to reflexively direct one's sp ati.a l attention
to an abrup tly occu rri ng tac.tile stinu1lus that takes pbce at a partic ular locn-
tion or part of the body. Such reactive control of s p3tial attention is lmmvn as
exogenous (or bottom-up) processlng
The sense of touc h also d emonstrates the attentiona l p henomenon of change
blindness, described in Chapter 7. ln a study mimicking the !licker paradigm U5ed
in visual change blindness, Gallace, Tan, and Spence (2007) stim ulated subjects
with vibra tions at hovoor th.ree sites on the body and asked them to detect when
FIGURE 13.25 S tudyi ng competition between siE:>nsory modal-

iti9s. In 9ach hand the subj9Ct hcids a cu be that has vibrotactile
stirnulators and tights, either of w hich can s4gnal the r€oQuired
respcnSEll. Th@arrows miar the fE1edback lig hts are usecl to direct
attenUon. (Aft.er SP€'JC9, Pava:n l, and Drlw r, 2000.)

420 CHAPTER 13
the nurnber of vibrations changed fro m one presentation to the next. \ Vhen the
second presentation \Vas preceded by a brief period in which multiple sites were
stimulated , the ability to detect a change decre<lSed dramati cally. Even a sin gle
irrelevant v ibratory s tirnulus, called a "mudsplash," ben.veen the t>vo vibratory
pattem.s substantia lly decreased the ability to d etect differences betvveen them .
Social Touch
Influen ces o f touch extend beyond perception and ac tion even to encompass
social interaction. Many words describing interpersonal responses refer to haptic
properties, as when ,...,.e describe son\eone as ''warm,'' "cold,'1 "hard/' or even
''a soft touch.'' Work of Inagaki and Eisenberger (2013) suggests that these ver-
bal connections are not ju.st metaphoric. After reading loving messages from
o thers \.'fi th whom they had dose relationships, people felt physical ly warmer,
and conversely, holding a heated pack led to heightened feelings of social con-
nection. Brain im..'1.ging further t'evealed that activation by both physical wannth
and loving messages was found in common brain areas, particularly the insuk\,
an area that we have seen is associated \1ri th positive touch-related experien ces.
The connection between physica l and social wa m1th m.Jy be the res ult
o f past experien ce \vith the physical of a caregiver, but it may also
have a gene tic component. It has been fo und, in fact, that touch affects social
interactions through indh;dual characteristics tha t we think of as geneti-
cally determined 1 like personality. Work in the field o f epismetics has led to
a mode l describing how the expression of genes related to social intemction
is controlled by to uch in the Mother rats who lick and groom thei r pups
produce offspring wh o tend to lick.and groom their ow n pups as well. ;'Ahat"
we m.ight say, there ll"tus t be a licking-and-grooming gene that is passed from
one generation to the next. But to complicate matters, if pups fro m attentive
and remote rat moms a re switched at birth1 as in the old fable, they 1'inh erit "
the behavims of their ad op ti ve, rather tha n biological, m others (D. Francis et
aL, 1999)_ 1l1eir maternal behavior is governed not by their genetic makeup,
but by the grooming behador they have ex-perienced .
Have the pups lea rn ed from their experience, then, to lick and g room ?
The m echanism foy pas.sing on the behavior is not wh at we normally think of
as learning by association. There is in.stead an epigenetic process that turns
licking-and-grooming genes off or on, according to the pup's experience with
the same behaviors 1 by tegulot_m g neJ.uoendocrin aystem.s (Champagne,
2008). Complex beha viors are transmitted from one rat generation to the next
by regulation of the genes, no t control of their component DNA. Evidence for
a similar m echanism in humans has been obtained (McGowa n et a l., 2009).
TI1e rat mother 's con tact has col'\seq uenre.s well beyond the ma temal care
exhibited by the offspring when they in turn have pups. Off.spring of licking-
and-grooming mothers turn out to be less timid than those of rem ote m oth ers.
Interactions between Touch and Other Modalities

Touch does not occur only in the absence of other sensory input, of co urse.
We comrnonly touch objects that we see, a nd we hear the consequ ences of
cont.act. How d oes the perceptual system as a w h ole deal with s ignals from
multiple modalities? Sometimes they compete, and sometimes the wh ole is
an integrated. combination of th e different inputs.
Competition e::1n arise w hen resources are llmited-thatis, whe n attention
comes into play in a particular tas k. Do the different modalities compete \·\rith
each othe r for a ttenti.o nal resom ces as well? Consi de r the fuct that the pressure
stimulus of your posterior on your seat seems to have lost out to this visual
text in the current competition fo r you r (until n ow, th:1t is). ln the

TOUCH 421
lab, Spence, Nicholls, a nd Drive r (2001) did a cross-modal version of the same
sort of endogenous cueing experiment that was described previously-: nley
led particip<mts to expect a s timulus to be presented via one modality and
then so metimes presented it in a different modality. The participants were
instructed to indicate vvith a foot pedal v.1hether a target sti..rnulus appeared
on their left 01· right side. The s timulus could be n oise frorn a loudspeaker
(auditi on), a red cirde at the location of the loud speake r (visiont or a rod
pressing the finger lvhile it to uched the loud speaker (touch ); a nd a n1e could
direct a ttenti on to\-.·ard any of the three rn od.a lities.
Again, respon ses were fa ster w hen the cue was valid and s lo,,..... er w hen it
\\.'as invalid. In terestin gly; the g rea test cost for an invalid cue occurred when
observers expected a tactile stimulus but a visual o r auditory stirnulus was
presented instead . This result may imply that the sense of toud1 has a par-
ticularly restricted atte ntiona l duumel-that once a ttention is focused. on the
touch m odality, it is relatively di.Hicult to realloca te it. O r it be lAppa
attention and a uditory a ttention are shared to sonle exte nt, because expectan-
cies in those m odaliti es could be di recte d. to a common loca tion in external
space, w hereas the expectancy for toud1 is directed to a location on the body.
In contra.st to attention['!! competition, intersensory can occur
w hen different modalities receive in.formation about the same object.Suppose,
for exa rnple, tha t you 're to uching sandpaper. The roughness you feel ::i.lso
d epends on the roughness you see. Participants in an experiment by Led er-
man, Thorne, and Jone5 (1 9&6) snw one sandpaper surface and felt another
but were told the tvvo s ur faces were one and the sa me. \ Vhen they were asked
about how closely packed the elements in the surface were, they were more
strongly influenced by vision . \'\'hen asked abo ut d1e roughness of the surface,
however, to uch became more imp ortant and vision less important.
In some circumsta.nces, one modality may appear to d ominate. In a classic
study pitting vision against touch, Rock and Victor (1964) had people grasp a
sq uare while looking at it through a distorting lens. \Vhat these participants
felt '"'as p retty much w hat they saw: a rectangle. That is, feeling that the sides
of the sh.ape weceeq ua l in length had essentially no influence. But dominance
by one modality over the oth er is not the rule. A more gene ral model is tl1['1t
people sign..1.ls from hvo m0t-l alities, producing a weighted average.
Tha t is, tJ,ey u se x percent of the information from one modality and {100- x)
percent from tJ,e other. The relative weighting reflects the quality of the signal
from each modali ty.
Ernst and Banks (2002) demonstrated such integration with an appara tus
tlu'l.t si multaneously created touch and sight of the same ,,; rtual display.
The display consisted of a plane with a raised bar across the middle (F1gure
13.26). The virtual visual display was produced with stereo g lasses and a n
app ropriate pair o f random dot s tereogrn ms, one presented to each eye (see
Figu re 6.33), cre.:1 ting the visua l il lusion of a bar stepping up frorn a plane. The
corresp onding vi rtual tou ch display was created wi th a device that gen era ted
forces that pushed back on the hand whenever contact was made wi th the
s.imulated s urface. When the bar heights presented to the two mo dalities did
not matd1 (the to uch ed bar was higher than the viewed ba r, or vice versa),
the perceived height of the bar was a 'veighte d compro mise betvveen them ,
wi th vision more strongl y weighted tha n touc h. Wh e n the investigators made
the information from vision less reliable by changi ng the apparent
height of some of the surfa ce d ots, the weight assigned to tou ch increased,
and it played a g reater r ole in de tennining the perceived height of the bar.
Multiple modalities ffkl}' collabora te by signaling different, but complemen-
tary, sources of information about an object. O ur discussion of object percep-

422 CHAPTER 13
l=IGURE 13.26 Testing the intggra-

tion of sensory modalities. The obsetv-
er could SQQ a virtual surface of dots
through stereo goggles, which gave
the appearance of a raised bar a;ross CRT
the surface, and could touch the 'w'irtuaJ
Sl.lrfaoo and receive resisting forces
consistent with th€I surfaca h91ght.
'"-ft"' Ernst and Banks, 2002.)
Opaque mirror
Visual and
hapticBCElE'
tion emphasized that vision and toudl are intrinsically complementary: one
8230336 A is well Suited to convey an object's geometry; the other, its material. Perhaps
the phil050pher Molyneaux had this complementarity in mind when he asked
his famous question about whether a blind person who gained sight would
recognize familiar objects with this new sense.
For a quite different example of intennodal cooperation, consider a study
about the perooption of object shape by Newell et al. (2001). In the learning
phase of the experiment, observers studied a set of Lego shapes, each fixed in
a partiaJar orientation, using vision or touch (Flgure 13.27). In the recognition
phase, previously studied shapes were presented, oriented either as they had been
during the study phase or reversed 180 degrees, along with unstudied shapes.
Again using vision or touch, the subjects were required to indicate whether
each tested shape was old (previously sh1died) or new. Not surprisingly, people
who studied and were tested in thes...'IDl.e modality found iteasier to recognize
old shapes when the orientation did not change. But now for the surprise: for
people who studied and were tested in different modalities-vision changed to
toud1, or vice versa-recognition was actually better when the shtdied shapes
were reversed in orientation during the recognition phaset Why should this be?
lt tum.s out that the natural way to hold an object is thtunbs in, fingers out, so
that we learn more haptically about tl1e back than the front. When an object that
we viewed is then explored by touch, reversing it means the more effectively
FIGURE 13.27 An 0)(8.mple of the explored back surface matd1es what was accessible to vision. Thus, it would
object shapes used In Ngwg1·s experi- appear that two modalities might most efficiently process an object's shape by
ment. (After NQll.l\IQll et al., 2001.) a nah1ral collaboration: vision for the front, toud1 for the back.

TOUCH 423
Haptic Virtual Environments

Anyone w ho has played a \'ideo game has been in a haptic virtual haptic virtual environment A syn-
environment. The actions of the player (e.g .. button presses or joystick thetic world that may be experienced
movements) are sensed by the m achire and filtered thro ugh a program haptlcally by operation of an electro-
that creates a simulated wor1d , causing new events and rutcomes that mechanlcal device that forces
are fed back to the user thrcugh vision and audition. Although some to the hard of the user.
joysticks ernpoy crud e vibration. vvhat is missing from most of these
environments at present is haptic fee::ib:lck. However. Interfaces have
been developed that provide such feed back in the fonn of vibratio n o r
sustained foroes to the hand. diverse objects varying in shape,
size. surface texture, and softness to be rendered .
Haptic virtual environments are useful in many applications other
than video games. They enable an operator to manipulate oqects \Vith
a remote robot and feel the forces t11e robot hand encounters. Tuey are
used to train novice surgea1s for minimally Invasive surgery, where the
operator manipulates an implement inserted th rough a small Incision
while viewing the surgi cal site on a video display. In the haptic virtual
training environment. the patient's body Is replaced by a dummy, and
the surgical tool conne:::ts t o a ocm pLrter that tracks the tralnoo's move-
ments. The computer contains a simulation of the pati ent that describes
body structures ard their properties, such as slipperiness and softness.
As the ccmputer tracks the surgeon's actions with the tool, It deter-
mines the effect those actions would have on the simulated patient ,
and It generates high-precision grapl"jcs and forces to feed back to the
surgeon (Rgure 13.28). Commerce on the Internet Is a potential domain
for force-feedback devices that could allow products to be felt as well
as seen.
FIGURE 13.28 A virtual surgical trakl&r. A navi09 SLl"QQOO rQCQ!v9S high-

p<ecision graphics and force fe&dback about a bkx:>d vessel that Is being
repaired. (AftEf photographs from Boston Dynarrics.)

424 CHAPTER 13
Summary
1. TI1e sense o f touch produces a number of distinct sensory experiences. Each
Refer to the type of experien ce is media ted by its own sensory receptor system(s). Touch
Sensation and Perception receptors are respons ive not only to pressure, but also to vibration, changes
Ccmpanbn Website in temperature, and noxiou s stirnulation. T11e kines thetic system, which also
contributes to o ur .sense o f touch, is further involved in sensing limb posi-
slles.slnauer.com/wolfe4e tion and the movem ent of our limbs in space. Pleasant or emotiona l tou ch is
for actMties, essays, study anothe r form of sensory specialization .
questioos, am other study ads. 2. Four d asses of pressure-sensitive (me.cha.no-) receptors have been found
w ithin hairless skin, and an o ther fi ve classes ' ""ithin hairy skin . The o rgans
u sed to sense limb position and movement (namely, o ur muscles, tendons,
and joints) are more deeply situa ted l 1rithin the body. TI1ermoreceptors
1
respond to changes in skin tempera t ure that occur1 for example1 w hen we
contact objects that are w armer or cooler than our bodies. N ociceptors sig-
n al tissue d amage (or its potential) and give rise to sensations of pa in.
3. The p a thw ays from touch receptors to the brain are complex. Two major
pathways have been identified: a fas t one (the d orsal column-medial le m-
n iscal p athway) tha.t carries information fro m mechano recep tors 1 and a
8230336 Anma App s lower one (the spinothalamic pa th way) tha t carries thermal and nocirep-
tive information. Both en ter the dorsal hom of the spinal cord, whi ch itself
has dense ne ural connectivity. 111e pathways project to the tha lamus and
from th ere to the primary somatosensory a rea, loca ted in the pa rietal lobe
just behi nd the central sukus . TIUs a rea contains several somatotop ically
organ ized subregions, in w hich a djacent areas o f the body p rojec t to adja-
cent areas of the brain. TI1e neural o rgan ization of the brain for touch has
been s ho\\11\ to be remarkably plastic, even in adul ts.
4. Do\¥nward pa thways from the bra in play an important role in the percep-
tion of pain . According to the gate control theory , signals along these pa th-
ways interact at the spinal cord with those from the periphery of the body.
Such interac tions can block the pain signals tha t would otherwise be sent
fonvard to th e brain. The .sensation of pain is further moderated by areas in
the cortex.
5. In vestigators have measured sens iti vity to mechanical for ce by app ly ing
nylon hairs of different diameters to the skin. They d etermine s patial acuity
of the skin by the n ..•o-point touch th reshold, and more precisely
by d iscriminating the orientation of gratings applied to the skin. Tactile
pressure sensi tivity and spatial acuity vary with body s ite because of vary-
ing con centrations of different types of mechan orecep tors. The minimum
d epression of the skin needed to feel a stimulus vibrating a t a p articular rate
(frequency) provides a measure of v ibra tion sensitivity.

TOUCH 425
6. The sense of tou ch is intimately related to our ability to perform actions .

Signals fro m the mech...m oreceptors are necessary fo r simple actio ns such as
g rasping and lifting an o bjec:.t. Conversely, o u r own movements determine
how to uch receptors respond and.1 hence1 which pro perties o f the concrete
world we can feel. To uch is better adapted to feeling the material properties
of objects than it is to feeling their geometric features (e.g., shape), particu-
larly wh en an object is large enoug h to extend beyond the fi ngertip.
7. Like o ther sensoiy modalities1 tou ch g ives rise to internal representations of
the world, w hich convey the positions of objects using the bod y as a spatial
reference sys tem . Touch-derived representations.are inputs to hi gher-level
functions like a lloea tio n of a ttentio n and integra tion w ith in.forma tio n from
o ther modalities.
8. The psychological study of to uch is useful for a number of applications .
Virtual touch environm ents that trans mit forces to the to uch recepto rs can
provide a basis for training people to perform re mote operations like s u r-
geiy and perhaps, in the fuh1re1 will convey the illus ion of to uched obj ects
over the Internet.
8230336 Amma Appa

CHAPTER 14
8230336 AMma Appa

Olfaction
THE STORY OF Tl-IE NEXT TWO CHAPTERS begins at the d awn of life itself.
When sing le-celled orga n isms first appeared on Earth, their basic p urpose in
life \.vas to take in som e s ubstan ces (food) and avoid others (toxins) . As these
organisms evolved into multice llular crea tures, de tecting ch emicals in the en-
vironment continued to be crucial for surviva l. Sys te ms to de tect and analyze
e n\oironrnenta.l molec ules were thus the first senses to evolve.
Humans have hvo main chemical de tection syste ms: one for m olecules
fl oa ting in the air, and a nothe r for m olecules that we put in o ur m ouths. The
technical names for these h vo systen1S are olfacUon and respectively.
The form er, m ore commo nly known as "smell,'' is the s ubject of this d lapter.
G usta tion, which you p robably know as "taste/' "'ill be explored in Chapter
15. Another ch e mical-sensin g syste m that is important fo r our exp eriences of
bo th smells and tas tes is the trige minal system, llmervated by the trigeminal
ne rve. The trigemina l system enables us to feel gustatory .:ind olfactory experi-
ences, like burning a nd cooling, and you w ill lea rn rnore about this system
in Chapter 15. Our sense of s mell is cri tically involved in otu exp-erience of
food. The reason is because there are t\vo routes throu gh which we perceive
odors. First we have orthonasal olfactlon, wh ich occ urs w hen we sniff odor-
ant molecu.les through our nostrils and they tra vel up o m n ose and onto the olfactlon The sense of smeU.
olfactory epithelium---as happens when we smel l a rose. Orthonasal olfaction gustatlon The sense of taste.
is the primary topic of this The second ro ute is ca lled retronasal orthonasal olfactlon Snlffl ng in and
olfactlon, w hid1 occurs when we breathe in odorant m olecules throu gh our perceiving odors throu:;ih our nostrils.
m outh and they travel up from the back of our mouth into o ur upper n asal which occurs when we are smelli ng
cavi ty and onto the olfactory epi the lium. Retronr.sa.1 olfoction occurs w hen we somethirg that Is outside of us.
areeatin ganddrinkingand produces the sensation of "flavo r. "You w ill lea rn retronasal olfactlon Perceiv-
m o re a bout retronasal olfaction a nd our experience of flavor in C hapter 15 . ing odors through your mouth while
breathing and chev.1ng. This occurs
when we are smelling something that
Is Inside our mouth ard Is vmat gives
Olfactory us the experlence of flavor.
odor The translation of a c hemlcal
Odors and Odorants stimulus Into a smell sensatlc.n. For
O lfac tory sens<i ti ons are ca ll ed odors. 111e s timuli for odors are che mical example, ' The cake has a chocolate
compounds ca lled odorants. An analogy c..111 be made '"·i th vis io n , w here OOor."
wavelengths of light are stimul i and colors are the vis ual sensa tions . However, odorant A molecule t11at Is denned
n ot every che mical is an odoranl To be sm elled, odorant molecules must be by Its ph'fsloochemlcal characteristics,
volatile (able to fl oat through the air},small (less than abou t 5.8 x 10-22 grams}, which is capable of being translated by
and hydrophobic (repellent to wa ter). Figure 14.1a shows the che mical s truc- the nervcus system Into the perception
of a smell. For ex.ample, "You were
h.tres of two odorant molecules. H owever, many m olecules that would seem to given the odorant methyl to
meet the basic requirem ents sti ll d on' t sm ell to us. Two examples are natural smell, vltilch has the oda of winter-
gas (me thane) and a by-pro duct of meth..1.ne, carbon m onoxid e {Figure 14.1b). green mint."

428 CHAPTER 14
FIG URE 14.1 Odorants.. (a) Most

small, volatile. and hydrophobic m ol-
8CUl9s activati9 the senS9 of smell, '1J)
but are notable exceptions to
th9 rul9, suc h as IT!Qthane and carbon
rnonoxide.
(b) 1'.'fethane Carbon monoxide
O ur evolutionary ancestors would have ha d n o reason to d e tec t these sub-

s ta nres, w hich no t d an.gerou s in the con centra tions found in nature. But
82303 th buildup of carbon monoxid e in enclosed spaces s uch as homes
wi th furn.aces can be- fatal, gas companies add a comp ound (tertiary-butyl
mercn. pran) that we smell as rotten eggs, to act as a wa rning signal when a
s tove 's pilot light goes o ut. We a lso sm ell the m olecules tha t make up
the a ir we breathe, such as oxygen, helium, and nitrogen .
The Human Olfactory Apparatus

Unlike the visual a nd auditory sys te ms, but like the syste ms of to uch a nd
taste, the human olfactory system is tacked onto an organ that serves an othe r
FIG URE 14.2 The nose. AJthough the primary func-

Au and tion o f the nosQ is to warm and humidify th9 air that w e
odorants bfoo.the, the no se also dirnct s odorants o nto the olfactory
epithelium.

O LFACTlON 429
Mitralall Tufte::l.cell Olfadorrbulb
olfactory clett A namw space at

the b ack of tha nooe Into which air
fiows and where tha olfactory epltha-
llum Is located.
olfactory eplthell um A secretory
muocus membrane In the hUman nose
Olfactory whose prmary function Is to detect
odorants In Inhaled air. Located on
Olfactory Basal all both sides of the upper portlm of the
&eNO'J' nasal ca\Aty and the ollactory clefts ,
the olfactory eplthallum contaJrs three
types of cells: olfactory sensory neu-
FIGURE 14.3 The r&tlna of the nos&. The olfactory eptheHum contains three types rors, basal eel.,. and supporting eel.,.
of cells: olfactory sensory neurons (OSNs), basal cglfs. and supporting CQ!ls.. Thg nasal dominance The asymmetry
OSNs are located beneath a wat&ry mucous layer on the eplth911um; the hairlike cllaraclerlzlng the Intake of air by tile
olfactory cilia of the OSN dendrit9s project through the mucus and are the recgptor two nootrlO;, which correspmds to dW-
sites for odorant moli9Cules. The d lfferent colors of the OSNs in this Illustration repre- fenng sensitivity to odorants between
sent the glomerull onto which they will cor1V9rge. .AJI blue OSNs go to blue glornerull, the two nostrils. Nasal dominance
all purple to purple, and so on. This schematic Mlustrates the fact that different OSNs alternates nostrHs thrrughout the day,
expressing the same receptors converge on the same glomgruH, no matter whierra but there Is no predlctablllty about
thsy are In th8 olfactory eptth911um. when the nostrils alternate.
support! ng cell One of the three
types of cells In the olfactcry epi-
thelium. Supporting eels provide
purpose. The primary function of the nose (Figure 14.2) is to filter, wann, and metabolic and physical support for the
humidify the air that we breathe. But the in.side of the n ose h as small ridges dfactory sensory neurrns.
called turbina tes that add turbulence to incoming air, causing a small puff of basal cell One of the three types of
each breath to rise upward, pass through a narrow s pace called the olfactory cells In the olfactory epithelium. Basal
cells are the precursor cells to dfactory
cleft, and settle on a yellowish patch of mucous membrane called the olfactory
sensory neurons.
epithelium (Figure 14.3). Notably, our two nostrils take in different amounts
olfactory sensory neuron (OSN)
of air, and this nasal dominance alternates between our nostrils throughout
One of three cell twas-the main
the d ay. This means that the two nostrils continually vary in their sensiti\o;ties me-In the olfactory eplth>lurn. OSNs
to odorants as a ftmction of the amount of air inhaled. are small neurcns located beneath a
The o lfactory epithelium is the "retina of the nose." \Ve have an olfactory muorus layer In the epithelium. The
epithelium at the back of each nasal passage, about 2% inches up from the c ilia on the OS N dendntes contain tha
nostril. Each epithelium measures about 1-2 square inches (depending on the receptor sites for odcrant molecules.
size of the n ose) and contains three types of cells: supporting cells, basal cells, cilium M'f of tile halnlke protrusions
and olfactory sensory neurons (OSNs). (See Web Activity 14 .1: Olfactory m the dendrites of olfactory oorsory
neurons. The receptor sites for cdorant
Anatomy.)
molecules are on the cllla, which are
OSNs (Figure 14.4} are small neurons that have cilia {singular cilium} pro- the nret structures Involved In olfactory
tnid.ing into the mucus covering the o lfactory epithelium. These cilia, which signal transduction.
are actually tl1e OSN's dendrites, have odorant receptors (ORs) on their tips. odorant receptor (OR) The reglcn
The basic rule of o lfactory sensory physiology is "one to one to one"; each OSN actory sensory nw-
m the cllla of o W
expresSES only one type of odorant receptor (OR), and all OSNs expre;sing the rors where odcrant molecules bind.

430 CHAPTER 1 4
FIGURE 14.4 A flucr9Scence imag9

o f an olfactory senSCf'/ neuron {lower
right), with a schematic graph (upper
right) of an action potentlaJ sequence
fol bwing odorant appHcation. The
OSN collects odorant rno loc:uloo \lia
receptors on its dendrites and sends
action potentials to th9 brain through
its axons. Note that pA = picoampgoo. 0 12 3 4 56
(FluoresC9na;;. image court9sy o f C. Time(s)
Balmer and A LaMantia.)
same type of recep tor project to the sam e g lom erulu s (see Figure 14.3). The
interaction between an odorant and the OR stimulates a cascad e of biochemical
events, ultimately producing a n action potential that is tran smitte d a long the
axon of the OSN to the olfactory bulb {Schild and Restrepo, 1998). To initiate
an action p ote ntial, about seven o r eigh t odor.mt m olecules must bind to .1
receptor, a nd it hikes about 40 of these nerve impulses fo r a sm ell sensa tion
to be reporte& You 1night be wondering how many od ors can we d etect? Th e
latest findings suggest that the ans\ver is over one trillion (Bushdid et al., 2014)!
Jn other words, the quantity is infinite. \.Ve can de tect any and all "smellable"
rnolecule mixtures. TI1is is far grea te r tha n the munber of colors we can see
(up to about 75 million) and the lllU11ber o f tones \ Ve can hear (approximately
340,000). Given that the universe of sm e ll is rig ht under our n ose, can we
actu ally perceive the scent of a trillion odors? The an swer is "no"-for various
reasons. Beyond the fact tha t we couldn't possibly li ve lon g enough--one tril-
lion seconds is about 32,000 years--there are other m ore mundane con s traints
as you \vill discover throughout th is chapter.
Tilere h..'1.S been controversy over how many 0'3Ns hm:nan.s possess, because
the ntunber of OSNs in the human olfactory epithelfa h as never actually been
cow1ted. 111erefore, the nwnber arrived a t hils been extrapolated from research
in other m ammals. Until recen tly it was believed th at we had about 20 million
C>SNs, s plit the epitheHa of our right and left nostri ls. H owever, cur-
rent°'researm in rod (L<am and Mombaerts, 2013) and mic.rographs of the
hum an epithe lia (Holbrook et a l., 2011) now suggest that the number o f OSNs
is 5-10 million, and that there are m ore OSNs expressing the sam e OR than
previous ly thought (personal com nnmication Charles Greer, June 4, 2013). Mo!'e
research may soon clarify this issue. Whatever the exact ntunber of OSNs, we
know that vision is the only sensory system tha t has more sensory neurons
than olfaction. Despite this la rge number we are n ot the extreme sniffers of

OLFACTION 431
(II) (b) FIGURE 14.S Tracking scents.

(a) The path (red) o f a do;J following
the scent trail (yellow) of a pheasant
dragged thro ugh a fi91d. path
l)'ed) of a t1uman fcilowing a scent
trail of chcx::dat.e essential
oil through a field. (After Porte:ir et at,
2007 .)
crfbrtform plate A bony structure

riddled with tiny holes, at the level of
the eyebrows, which separates the
nose from the brain. The axons from
animal king dom. Dogs have a t least 100 times moreOSNs th.an humans, and a the olfactory sensory neurons pass
mud1 hi gher proporti on of the dog's brain is dedicated to olfoction: abou t 5%, thrrug h the tiny holes of the cllbnform
compitred wi th 0.1% for humans. Resear-chers s uspect that human ::i. a m s me ll plate to enter the brain.
about the same nmnbler o f Scents ftS dci"gs-(the bloodhounds aren' t t..'tlklng,so anosmla Tile totat lnablllty to smal,
we can 't be sure), but d ogs can sense odors at concentrations nearly 100 million most often resulting from sinus Illness
times lower than the concentrations that humans can de tect {KresteJ et a l., 1984; cr head trauma.
Willis e t aL, 2004). Nevertheless, in a recent s tudy humans were able to fo llow
a 10-mete r-l on g scent track of chocolate aroma while on all fo urs in an open
grass field (Porter e t al., 2007), and the tracking patten\ th.at the participants
used was s trikingly .sirnibr to that of a d og (Figure 14.5 ). O ther amazing
srnellers indude pigs, whic h ca n s mell the scent of truffles (the mus hroom,
n ot the chocolate) under 6 inches of soil, a nd salmon, ""· hkh use s mell to find
the wa ters of their birth from hun d reds o f rniles away (Dittman and Quinn,
1996). But the m ost extreme s niffer of all appears to be the African elephant
(Figu re 14.6), i.-.rith approxirnately 2000 ftmctional ORs (rnore than fi ve times as
1nany fun ctional ORs as humans) an.d the longest nose in the animal king dom
(Niimura et aL, 2014). Their incredible proboscis a llmvs the m to distinguish
w ith jus t a whiff two ethnic groups wi th whom they share their habitat: the
Maasai wh om they are afraid of because to sh ow their virility young Maasai
men spear them, and the Kamba whom they ignore-the Kamba are farmers
and do n ot bother the e lephants (Ba tes et al., 2007).
The axons o n the ends of OSNs o pp osite the cilia (<lend.rites) pass throug h
the tiny s ievelike holes of the cribriform plate, a bon y s tnictme at the level of
the eyebrows tha t separates the nose from the brain. A hard blow to the front
of the head can cause the cribriform plate to be jarred back or frad:ured, slicin g
off the frag ile olfactory axo ns , and consequently inducin g anosmla e·sm e ll
blindness"), the to tal absence of a sense of s mell. Stern cells in the olfacto ry FIGURE 14.6 The impressive
e pithelium can fonn new OSNs; indeed, all of o ur OSNs die and regenerate proboscis. of an African e4ephant .

432 CHAPTER 1 4
about once every 28 days. However, frach.ired cribriforrn plates typically scar
over, preventing the new OSN axons from passing through to the brain, and
the sense of smell for life.
Anosmia
Anosmia rsmell blindness) Is the total absence of a sense of smell.
Hypcsmia is a signlficantty redu::::ed sense of smell. Smell loss is much
more prevalent than most people think. From a suivey by the National
Institutes of Health (NIH) In 1994, it was conservatively estimated that
one in every 100 people suffers from anosmia. However, mere recent
estimates indicate that as many as 14 million Americans over the age of
55 have a severely cornpromise:::1 sense of smell. When younger adults
are included. it appears that about one in 20 Americans may have some
olfactory dysfurr:::tion. However. compared v•th loss of vision or hearing,
olfactory loss Is paid little attention by both the medlca and the gen-
eral community. The American Medical Association val ues the loss of
smell as 3-5% of a perscn's life worth. \Nhereas loss of vision is valued
at 85% , In a questionnaire administered to students at the University
of Pennsylvania. loss of the sense of smell was ranked equal to loss of
one's big toe ('Nrzesniewski. McCauley. and Rozin. 1999).
Despite general indifference to sm01 loss, anosmlacan cause
great sUffenng. Indeed. It has been reported that atthough the trauma
of being blinded is Initially worse than losing one's sense of smell, after
one year blind people have begun to cope 'Nell, 'IJJhereas those with
anosmla are faring worse and detertoration in their quality of life typl-
cally becomes progressively more extensive (Herz. 2007). The reason Is
that loslng our sense of smell affects almost every aspect of our lives.
The consequences at smell loss are most obvious in our enjoyment of
food and dr1nK and the experience of flavor. But anosmla can also lead
to ser1ous health oonsequerces , as people cannot detect food poison-
ing that doesn't have visual cues, and they cannot perceive smoke or
the scent that signals an accumulation of carton mcnoxide. Anosmia
also affects peoples ability to function socially and to be intimate in
sexual relationships. It impairs oogritive and spatial ability, eliminates
some of our most significant memories , alters our sense of self. and
can profoundly disturb emotional health. People who lose their sense
of smell often fall into clinic.ally depressive states, and because the lack
of olfactory stimulation leads to progressive emotional deterioration, the
depression that results can become dangerous.
Being born arosmic Is quite rare and affects less than 0.06% of
the pcpulation. The most common cause of anosmia is sinus infection,
followed by nasal polyps. Chronic sinusitis and allergies can also lead
to anosmia. In these cases anosmla Is due to blocked nasal passages,
which prevent odorants from interacting vlith tile olfactory receptors.
Usually v.nen these conditions are c ured, the person regains notr11al
smell function. HO\i'1ever, about 30% of anosmia is caused by head
injury. which can oo::::ur In sp:Jrts like football or boxing or through car or
bicycle accidents. In these cases smell loss is usually pennanent. Inju-
ries tt1at involve a blow to the eyebrow-forehead region or the back of
the head cause the cr1brifonn plate. which is riddled with tiny ho les. to
be jarred. Dislodging the a-ibr1forrn plate sll9ars off the o lfactory axons
through which the olfactory axons pass. and although olfac tory neurcns

OLFACTI ON 433
c an regenerate, Injury to the crib rifo rm plate usually results in scarring

OJer of the tiny holes; t.hus the o lfactOfy axons c an never enter
the brain.
Many everyday toxi ns. including gasoline and halrdresSng chemi-
cals can c ause pennanent smell dysfunction (Smith, Da\.1dson, and
Murphy, 2009). Certain m edic ations c an also c ause smell loss or
distu1bance -particular1y those used in chemotherapy. In fact. many
common drugs, Inc luding antibioti cs, antidep-essants. and even sane
antihist amines, c an impair olfactory funcuon (Doty and Bromley, 2004).
Lo ng-tenri excessive alcohol consumptio n also red uces olfac tory func -
tion (Rupp et al., 2003). By contrast , marijuana m ay have amelioraUng
effects. as we' ll discu ss shortly.
Smell function also declines w ith age: however, the rate is gradual
and there is w ide lndMdual variability in severity. NotatJy, loss of smell
am ong the elderty who live alone c an lead to missed meals and i:oor
nutriti on . The symptom s of malrourtshment c an b e misinterpreted as
d ementia, and people m ay b e presumed to be cognitwely Impaired
when they are "only" experiencing the normal olfac tory dysfunctio n of
aging. Fortunately, when a healthy diet is implemented in these In-
stances, sig ns of oognitive impairment go &Nay. However. it Is also
the case that smell loss Is the first Viarnlng sig n of several neurolo gi-
cal disorders -in partirul ar, Alzheimer's and Parkinson's
disease (PD)-and sympto ms often appear years before any
other signs of the illness are present . If you know som eone over 4 0 1NhO
has sudd enly started having trouble identifying familiar smells. you may 36 Anma Appa
w ant to recommend a visit to a neurologist. The ear1ier AD and PD c an
b e d iagnosed, the better treaunent oLrt look and p rcgnosls are. Several
tests can be administered to assess smell loss. The m ost common are
the University of Pennsylvania Smell Id entific ation Test (UPS ll) and Snif-
fin ' Stick s. The UPSIT Is a 40-ltem scratch-and-sniff multiple-choice test
that m easures detection and Id entification . S niffin ' Sticks involve three
oifactory tests t o evaluate threshold d etect ion , cdor discrimination . and
odor Id entification .
Many smell and taste clinics throLQhout the c oontry administer
simple tests to deterrnh'le the causes and treatment possibilities for
o lfactory loss and dysfunctio n . However. currently there is no equivalent
of a hear1rg aid o r g lasses for the nose: therefore, if sm ell loss cannot
b e treated b y addressing the undertying c ondltlrn (e .g ., sinus infection ,
m edi catio n use). it is likely to t:e p ermanent.
olfactory (Q nerves The first pair of

cranial nerves. The axons of the olfac-
Neurophysiology of Olfaction tory sensory neurons bundle together
after passing through the crlbrWorrn
In som eon e \...·ith a healthil y functionin g sense of s mell, the OSN axons pass plate to form the olfactory nerve. whk::h
throu gh the cribriform pla te1 bundle together to fo rm the olfactory nerve c onducts Impulses trom the olfactory
(cran ial nerve 0, a nd enter a blueberry-sized ex tension of the brain just above eplthella In the nose to tho dfactory
the nose called the olfactory bulb (Figure 14.7 }. We h ave hvo olfuctory b ulbs, bulb .
one in each b ra in hem isphere. Unlike the other senses we've stud ied so far, olfactory bulb A bluoberry-slzad
olfaction is lpsilateral,. m eanin g tha t the right olfactory b ulb gets in for m a- extension of the brain Just above tho
tion from the righ t nos tril, and the left olfactory bulb gets in forma tion from nose. where olfactory infamatlai Is
first processed. There are two olfactory
the left n ostril. bulb s. one In each brain hemisphere,
a lso kno \.VT\ as the synthetic chemical methyl sa.licybte, has an odo r c orresponding to the right and left
that we id en tify as \Vintergreen mint. Like o ther o dor an ts, m ethyl salk y late nostrils.
0 111 activate severa l different types of ORs, a nd it d oes so \vith different de- lpsilate ral Referring to the sam e
grees of "weightin g" depend ing on the recep tor 's affinity to the ,,,-in tergreen side of the body (or bralrv.

434 CHAPTER 14
FIGURE 14.7 How srrells are perceived. '8) The pathway of olfac-
tory perc.6ptlon, from odorant molecule to the olfactory b.Jlb. {b) A
cross-sectlonat view of the nsural organization of olfactlon. Olfactory
Information Is transmitted from the dfactory bulb to the primary olfac-
tory c atax comprising the amygdala-hlppocarnpal complQX of the
Umblc system In tM first stages of olfactory processing. The orbito-
cortex ls cxmsldQ<ed to be the s€1condary olfactory cort9X and Is
rasponsit>IQ for conscious odor peroeption.
molecule. Humans have between 350 and 400 different types of hmctioning
ORs. Most of these receptors won' t be activated by methyl salicylate; a few
will be \'lleak1y activated, and one or two wW be strongly activated (this will
become dearer as you read the following sections).
TI1e first relay for the OSNs in the brain is in the olfactory bulb, w h ere the
sensory nerve endings gather together to form tiny spheres called glomerull
(singular glomentlus). Molecular genetic studies in mic:e have shov.m that all
neurons expressing a particular OR type, no matter where they are on the nasal
epithelium., send their axons to the same glomerulus pair (consisting of one
medial and one lateral glomemlus) in the olfactory bulb (Mombaertset al., 1996).
The d..istinctpattemofORactivation for a specificod.orant is then translated to
a specific pattern of spatial activity across the glomeruli. Methyl salicylate, for
example, will activate a set of specificORs and consequently produce a pattern
of glomemlar activity that is unique to it The compounds of rose aroma will
initiate a different activity patten11 and so on. The specificpattem of glomeruli
acti\.;ty is then interpreted by the brain as indicating a specific odor.
However, this relatively simple picture is complicated by the fact that each
glomerulus may receive axons from several different receptor types. Moreover,
our personal experience with an odor can actually dvmge the pattern of activity
that is produced by the glomen1li in the o lfactory bulb. For example, if you
liked methyl salicylate (winteigreen) mints but one day became ill after eating
tl1em, causing you to thereafter find the scent very unpleasant (illustratin g
glomerulus Alff of the spherical what is known as a "learned taste aversion," whid1 will be discussed further
conglomerates the Incom-
ing axms or the olfactory sensory neu- tOW'ard the end of this chapter), the pattern of activi ty produced by the scent
rons. Each OSN converges onto two of methyl salicylate in the olfactory bulb would be different from what it was
glomerul (ore medial, me lateral). when you liked wintergreen mints. Astonishingly, what this means is that

OLFACTION 435
there is no fi xed code for odor perception; rather, our personal experience \.vi th juxtaglomerulcr netrons The first
an o dor d etermines how it will be processed by the olfactory system, even at layer of cells rurrrundlng the glomerull.
ve ry early le vels (\'\'ilson, Best, and Sullivan, 2004). They are a mixture of excltatay and
lnl1lbltory cells and respond to a wide
Lrnnediately s urrounding ead1 g lornentlus is a rn.ixed p opulation of excit- range of odcranlB. The selectMty of
atory and inhibitory cells called juxtaglomerularnet.rons. These cells respond neurons to specific cdorants increases
to a mu ch wider range of od orants than the next layer of neurons, w hich is In a gradient the surface of the
composed of tufted cells, which them.selves respond to more odorants than dfactOI)' bulb to the deeper layers.
neurons in th e deepest level of neurons in the olfactory bulb, called mitral tutted cell The next layer of cells
cells, which resp ond to just a few s pecific odorants. Tha t is, the selectivi ty after the Juxtaglomerular neurcns. They
of ne urons to a specific od orant is increasingly s harpened in a gradient from rep;:ind to fW/er odcrants than the
Juxtaglornerular ools. but more than
the s urface of the o lfactory bulb to th e d eeper layers. 1l-1e mechani sms that neurons at the deepest layer of cells.
determine the response profiles to specific odorants are s till unknown, but its
mltral cell Tue deepoot layer of neu-
seems that this increasing fine tuning through neuronal layers is important rons In the dfactorybulb. Each mltral
for helping the brain distinguish behveen s imilar od ors (Kikuta et al., 2013). cell responds only to a few specific
Furthe rmore, at the d eepest leve l of the olfactory bulb, granule cells, an OOorants.
extensive network of inhibitory neurons, integrate input from a ll the earlier grarule cells Like mltral cells, gran-
projections and are thoug ht to function as higher-order hire detectors--c.J.- ule ceMs are at the deepest level of
pable of d etecting and learning specific combinatorial patterns of mitral and th3 olfactay bulb. Tuey comprise an
rufted cell activation and thus resp ondin g specifica lly to different odornnts extens.,.e network of Inhibitory neu-
rons, Integrate Input from an tl1e ear11er
(Koulakov and Rinberg, 2011). Axons o f the mitral nnd h1fted cells of each bulb
projections, and are thrught to be the
crunbine and fo rm the olfactory tract, one in ead1 of the brain, basis or specllc odcrant ldentmcatlon.
that conveys odor infom1ation ipsilateraliy to the primary olfactory cortex,
olfactory tract The bundle of axons
also knovvn as the plrlform cortex. The primary olfactory cortex compri ses
of the mttral and tutted cells within the
the nmygdala, parahippocampal gyrus, and interconnected areas (also known dfactory bulb that sends c:dor informa·
as the complex), and it intima tely i.nterncts with the tion to the primary dfactory oortax.
entorhlnal cortex (see Figure 14.7b). primary oWactory cortex or pir1-
Thou gh they numbe r in the milli ons, OSNs converge onto a relatively form cortex The neural area where
s mall number of gl omeru.l.L The m ouse, whose brain and OR physiology are dfactOIY Information is l rst processed.
mud 1 m ore devoted to offaction tha n a re those of htmtans, has360J glomeruli It comprises the amygdala. parahlp-
(Richard, Taylor, and Greer, 2010). lt therefore speculated that hum.J.ns, pxampal wus, and Interconnected
areas; an:J It Interacts dosely with the
w ith far fewer functi oning ORs, would ha ve about 700 glom eruli . However,
entorhlnal cortex..
immunohistochemis try research fr om C harl es G reer's laboratory at Ya le
University has reveal ed that humans ha ve as many as 60)()glomeruli - n early ppocampal com-
plex Tlie conjolnad reglrns of the
twice as many as the mouse for o nly a third the number of receptors (Maresh arnygdaJa and hlppocampus, wt>ch
et al.r 2008). This discovery was quite a s urprise and s uggests that using the are key structures In the limbic system.
mouse o lfactory system as a rnodel for h.tmUl.ns, which has been the traditional ltlls ccmplax is crltically lnvo.,.ed In
approad't, rnay not be as useful as once thought. the unique emotional and associative
The central brain structu res that process olfactory information are a ll pa rt properties of dfactory cognition.
of a netwo rk of structures known as the llmblc system, which is involved in entorhlnal cortex A ph',1ogenetl-
many aspects of erno tion and memory. As ,..,,e wil l see later, these connections cally dd cortic al regrn that provides
are key to the tmi'}ue emotional properties of olfaction. the major sensay association Input
Into the hlppocampus. The ootorhinal
It may be sttrprislng, but paleontologic.:1.l research h as sh o\VJ"\ that mammalian cortex also receives direct projectla1s
brain evolution was precipitated by an incre;tse in olfactory abil ity (Rowe, frcm oWactory reglrns.
Macrinin, and Luo, 2011 ). That is, the first s tructures in mammalian brains to
limbic system The enccrnpa.sslng
increase in size and complexity were tho.seas.sod.rited with the sense of sm ell. It group of neural strlciures that indudes
is theorized that increased olfactory c..1 paci ty enabled om 200-million-year-old the olfactay cortex, the arnygdala,
predecessors to hunt at night, giving. them the evolutionary ed ge they needed the hlppocampus, the plrtfcrm cortex.
to d evelop further and eventually evolve otrr current brains, which, ironically, and the entorhlnal conex. The limbic
system Is invotved In many aepect:s
along the way traded off olfactory advantages for vision (see disc ussion of of emotion and memory. Olfaction ls
pseud ogenes in ''The Genetic Basis of Olfactory Receptors" below). unique amcng the senses for Its direct
OUactory sensory neurons are diffe rent from all other sensory receptor connection to the limbic system.
cells in that they are not mediated by a protective b.urieT and instead make
direct contact with the brain. By contrast, visual receptors are protected by
the con1ea, 1·eceptors for hea ring are protected by the eardrnrn, and taste buds

436 CHAPTER 1 4
are buried in papillae. One consequence o f the fact that the olfactory sensory
n euron s are d irect conduits into the brain is tha t man y drugs can be inhaled.
In spire o f their di.red linkage into the brai.J\, OSN axons are a m ong the
thinnest a nd s lowest in the body. Therefore, eve n tho ugh the n ose connects
di rectly to the bra in, the time it ta kes to process a sensation (in this an odor)
is long, compared wi th mu o ther sen sory experie nces. The lag time between
sniffing and the brain's registerin g a scent varies, averaging approximately 400
milliseconds (m s), a lmost ha lf a second; compare this to the 45 m s it takes fo r
the visual cortex to register an image presented to the retina . This half-second
duratio n for odor d oes n o t take into account the time it takes to
react to a scen t, w hich effectively d oubles the pe rceptual time, making olfac-
tion a particularly s low sense. You ha ve probably observed tha t smells seem
8230336 Arr to mt'.fge gradually, rather than flashing into your awareness.
TI1ese d istinctions b ring up the s ubtle diffe ren ces between sensation and
perreption in o lfaction . Sensution occurs w hen an odor is ne urally registered;
percep tion occurs when we become aware o f d etectin g a scent, Odor clearance
is also slow, and you may have noticed tha t od ors tend to linger. 1his is due
both to a mbient air C'l.trrents (e.g.• a breeze vs. still air) a nd the time It takes
ORs to dear an odorant. The relatively s lo1N speed and lingering fea tures of
o lfa ctio n have been the central obstades fo r d eve lopin g effec ti ve 11sm ell-o-
visionu and olfa ctory virtual-reality technologies.
The Genetic Basis of Olfactory Receptors

ln 1991, molecular biologists Linda Buck a nd Rid1ard Axel {who were rewarded
\\lith a Nobel Prize for their efforts) showed that the m ammalian genome con tains
betvveen 1000 and 2000 different odorant receptor genes, depending on the spe-
cies (nxl.ents have the most), each of w hid1 codes for a single type of OR. The OR
genome is the largest gene fumily known in mammals. A_ll mammals appear to
have pretty much this sa me set of genes, but some OR genes in each species are
n onfunctional "pseudogen es'': the genes are on the chromosomes, but
the proteins coded fo r by the genes never get produced.. Tile latest figures sug-
gesttha t d ogs hove 811 func tion.<! OR genes 2011); therefore, about
19o/o of their OR genes are pseud ogenes. [n humans, the OR genome contains
about ] COJ genes, and the proportion of pseudogenes is between 60% and 7QO/o.
Some researchers have recently s uggested tha t the high proportion of OR
pseudogenes in humans is the of an evolutionary trade-off beh·veen vision
and olfaction . Yoav Gila d et al. (2004) observed tl»tOld World primate species
s ud1 as gorillas and rhesus monkeys have about 30o/oOR pseudogenes, but most
New\ Vorld species (e.g., squi rrel monkeys) have a lower proportion (abo ut 18%).
The one New \'\J'orld exception is the howler monkey, which also has about .30o/o
OR pseudogenes. It tunlS ou t that the howler monkey has something in common
wi th O ld \iVorld primates tha t other New World m onkeys d o no t: tridlI'omatic
color vision . Brains can get only so big, so to &-ee up the brain spa re necessary to
house human ancestors' evolvin g visual analysis tools (including trichromatic
vision), they may have d ropped the ability to analyze the od orants de tec ted by
1.."'ertain OR genes, and those then became pseudogenes. Fm our evolutionary
predecessors, the payoff from s upe1ior visual detection mus t have o utweigh ed
the disad vant.'lges from diminished olfacto1y acuity.
\ Ve don't have a p recise percentage of p seud ogen es for huma ns, because
researchers have found individual va riability in the number of fun ctional
recep tors expressed; the r of hmctiona.I ORs seems to be ben.veen 350 and
400. lndeed, there is enom1ous diversity in the repertoire of fon ctional od orant
receptor genes among differen t people. ln olfaction everybody actually has a
unique nose (McRae e t al., 2013; Menashe e t al., 2003). Thus, one person may

OLFACTION 437
express 358 ORs w hile another expresses 388, but both have a ''normal" sense of
smell. This mun her l'e flects both w hich genes are expressed as functional recep-
tors and how m any copies o f a specific recepto r a person may have. TI1e m ore
copies of a sp ecific receptor you have, the m ore sensitive yo u \vi ll be to certain
odmants (tha t is, certain specific di.enUcals will smell stron ger to one person
than they d o to an other because of the munber of receptor copies they have); and
whether you have a pseud ogene or a ftmctional gene for a given recep tor also
alters od or perception {Keller et aL,2007). Forexarnple, people w ho " hate" the
aroma of cilantro (also knm'ln as coriander) are rn issing the gene for detecting
the herbal flora l component of this aroma and therefore only detect the soapy
note(Kurz, 2008). Most recently the genes associated with the O R expression that
d etermine our sensitivity to fo ur more food -re levant beer, blue
cheese, and identified (McRae et al., 2013). However, as of nm.o\'1
the specific gen etic basis for our perception of m ost other arom as is tmknown.
The number of particular re ...""eptors people express influences their liking
for a n odor, because ha ving m ore receptors lead s to a m ore intense smell, and
strong scen ts in genera l tend to be perceived as less pleasant th._"l n the same
scents a t a lmver intensity (see the "Olfactory Hedonks" section of this cha pter}.
We see this relationship with intensity in all our senses, Very bright lights and
loud sounds are inherently m ore unpleasant than moderate-intensity lights
and sounds, and our relative sensitivity to light and sound w ill affect how
much each of us can tole rate (see C hapters 5 and 9 for more d eta il). Having
fevv receptors of a given type can lead LIS to perceive a scent weakly, and if we
like an a roma, we m ay expose ourse lves to more of i t to get the same punch as
someone 'vi th more ORs th at are sensitive to that od orant.
'When it comes to food, the m ore intense our perception of the retronasal
ar oma of the food1 the we te nd to consume of it (R uijschop e t al , 2008). For
example, if our OR expression fo r the odorant tha t contributes to the banann
aroma in bnnana cream pie is vveak, we may eat m ore pie than someone w ho
can perceive the banana aromastrongly, and this may lead us to cons ume m ore
d essert than we might othe rwise prefer to do. l11ese finding.s de mons trate the
subtle but irn portant ways by v.lhich OR variation o m influence our food choices
as well as our food intake. lt should, however, be noted that many psychologi-
cal factors inedfo te our food consumption as \vell as our odor preferences (see
"Olfac tory Hedonics"). The refore, our genetic makeup cannot predetermine
our response to a ny given odorant or food .
O ther factors rn n also increase our sensi tivity to odors te mporari ly. For
example, even wupt)on (e-,S,., three drinks per hour)
impairs olfactory sensi 1"1ty, h aving ln y one dri}lk\b)ood alcohol level less
than 0.06%) improves o lfactory acuity (Endeve1t-Shapira e t a l., 2014). This may
be a good reason to drink moderately whi le dining- with one glass of w ine
the fl avor aroma tics of your meal will be enhanced, b ut th e more you pmu
the less fl avor you' U perceive. Ano ther popu lar recreational drug, mariju:ana 1
also boosts olfaction . It was recently fm.md that activating en docannabinoid
receptors (the recepto rs that respond to the active component in marijuana) in
the brains of fasted mice increased their s mel l sensitivity and induced the mice
to eat more (Soria-Gomez e t al., 2014). Extrapola ting to htm1ans, the reason
for the "mtmchies" with a marijuana buzz may be tha t flavor (through smell)
is intensified, and thus w h at '"'e ea t tas tes richer and is more alluring. We get
a whiff of pizza and are immediately seduced. IndeedT increasing a ppetite
is one of the main reasons marijuana is p rescribed for patients undergoing
chemo therapy. Although this may seen1 like a contradiction to wh at was jus t
mentioned a bout grea ter flavor intensity decreasing food cons umpti on, the
ca nn abinol d effects observed were in fasted mice, not those w ho h ad recentl y

438 CHAPTER 1 4
trigemina (V) nerve The fifth cra- eaten. It may also be that the munchies ""'e experience are due to being less
nial nerve, which transmits Information aware of our inten-1a l ph ysical/ h o1neos tatic s tates w hen werre under the
about the ' feel" of an odorant (e.g., influence. TI1at is, we may no t be feeling whether o w· s to machs are hungry
mint feels coo, cinnamon feels warm).
as well as pain and Irritation sensations or full ; we may just be savoring the newfound fla vor intensity of pepperoni
(e.g .. ammCflla fees burning), pizza. This "mundlies" theory is currently speculative and based on anecdote
in humans, but it illustrates the multiple, sometimescontrad icto1y, and as yet
unknmvn ways that od or perception can affect our food intake. Stay hmed.
The Feel of Scent

As mentioned at the s tart of this d111.pter, our experience of od ors often has a
feel to it, as well as a smell. This is because m ost odorants s timulate the so-
rnatosensory system to some d egree through p olymodal n ociceptors (touch,
pain, and tempera hue receptors) inside the n ose. For example, m enth ol feels
cool and ammonia feels burning. These sensations are mediated by the
gemlnal nerve (cranial nerve V) (Figure 14.8a ). In many cases it is imp ossible
to distinguish between the sensations traveling up cranial nerve I from olfac-
tory receptors and those traveling up cranial nerve V from somritosensory
receptors. For exa mple, the nasal cooling (cranial ne rve \') and sped.fie scent
(cranial nerve I.) associa ted v1iiith the sme ll of peppermint fuse to produce a
holistic sensory experience. Tri ge.minal s timulation accounts for why our eyes
tear we ch op onions (as in Figure 14.8b)-and Vlh ve, sneeze. we
.sniff pepper. High levels of trigemina l s timulation can produce a severeD mn-
ing sensation, and trigeminal activity has been linked to the facial-head pain
felt in migraine headaches. "Smelling salts" (made from am mon ia combined
vvith e ucalyptus o il ) revive us because of their trigeminal activation.
FURTHER DISCUSSION of the trigeminal nff\'6, which also tel ls you about
fat in your diet, can be found ln Chaptet" 15 on page 488.
(a) (b)
Ophthalmic Mandibular Trigeminal
nen."t! nervekr.anial
nerve \f)
Ethrno1d \
FIGURE 14.8 The trigeminal nerve' s

rol9 In the PQrcgpticn of odors. (a) Th9
trigerninal ngrye carries information
from somatosGfisory r9Ceptors in the
nose an:::! face to the thalamus and
then on t o the sornatosensory cortex.
(b) The n;iason your eyes t9ar when you
c hop o nions is due to stimulation of the
I
lnfe1ioralveolar
trigillninal nerve. ne1ve(teeth)

OLFACTION 439
From Chemicals to Smells

Now tha t we know something about the physiologic.al basis of o lfuction, we shape-pattern theory The current
can ask the following crucial question: H ow does the biochemical interaction dominant blochemlcoJ theory fcr how
chemlcals come to be perceived as
between an odorant and an OR, and s ubsequent neurological processing in the
speclnc odors. Sl1apa-pattern theory
olfactory bulbs and later brain structures, result in the psychological percep- contems that different scents-as
tion of a scent s uch as wintergreen mint? Buck and Axel's seminal discovery a function of t11e It between odorant
of olfactory receptor genes prompted an explosion of research s urrotmding shape to OR shape-activate differ-
this q uestion over the pas t h vo decades, but a fully comprehensive theory of ent arrays o1cifaotory receptcrs In
how we perceive S\.."'ent.s has till realized\ tre olfactcry epltrella. These various
arrays prodl.!Oe spec1nc lnng patterns
of neurons In tre olfactcry bull>. v.tilch
Theories of Olfactory Perwption then determine the partlclllar scent we
At present, the best-accepted biochemicc1l theory (first proposed in its mcxlem perceive.
for m h1 the 1950s by the British scientist John Amoore) is on the match
between the shapes of odornnts and odorant receptors. It was dubbed "shape
theory" but is better deno ted as "shop<>-pattem theory." ln a nutshell, shape.
patte rn theory contends that odora nt molecules have differen t shapes and
o lfactory recep tors have different shapes, and an odornn t w ill be detected by
a specific OR to the extent that theodon:mt's molecul es fit into the OR {Figure
14.9). Gord.on Shepherd at Yale University and his s h1dents pioneered the idea
that w hen a given odorn nt is sniffed , a particular pattem is generated across
the g lomeruli. Differences in those spf\tial patterns provide the basis for the
arroJy of od o r s that\"""' perceive.
The m ost recent m olecular resea rch suggests that scents are detected by
tneans of a combinatorial code, where one odor<mt may bind to several differ-
en t receptors and one receptor may bind several diffi;;-rent odorants to varying
degrees (see Figure 14.9). This 1neans that different scents activa te different
arrays of olfac tory recep tors in the o lfactory e pithelia, produci ng specific
(•) Odor.ants Odoranl 1\!'ceptors (b) Odorant compounds Reo....--eptor arrays Hypothetical recepto r
activ.ation pattern
'iJ 'iJ C:'.J
DD t:H:='.l
c:: :J w t"'.l t::'.l
FIGURE 14.9 Odorant-receptor bind-

ing and odorant activation, as pr8dic1Qd by
shape-pattern theory. (a) Chembals of specif-
ic shapes fit r&CGptors with shapes that l>Qst
accommodate them. (b) Odoront m oleculeis
activate feature detectors on various roc.eptor
types. The specific pattern o f activity 91id tQd
• Best act iv a ti on
by a gtven set of receptors determines the
D Some activation
specific sc:ent p9rceiv9d.

440 CHAPTER 14
vibration theoty An altanattve to glorne rular activity in the olfactory bulb. TI1e s p ecifi c pattern of glorne rular
shape-pattern tlleo!y fcr des:::rlblng activity in the olfactory bulb then de termines the particular scent we perceive.
hmv olfactlon works. Vibration theory That is, different pa tterns are elicited for the pen::ep tion of rooe, rnint, urine, and
proposes that every cdorant has a
different vlbratlmal frequency, and sklmk (for example), and the various p a tterns for speci fic scents tum out to be
that molecules that produce the same highly consistent across individuals (Zou, Li, and Buck, 2005). 1his theory ''°'as
vibrational frequencies will smell the implied in the sec tion /(Neurophysiology of O lfoction ." Hm·vever, there are also
same. alternative exp la na tions of h ow olfaction works. The strongest alte rnative to
speclftc anosmla The Inability to sh'-'pe-pattern theory is vibration theory, championed mos t recen tly by Luca
s mell o ne specific ocrnpoond amid Turin (Franco et al., 2011; Turin, 1996). ln essen ce, v ibration theory proposes
otherwise normal smell perception. tha1t o f a_tomic :5tnicture, eve1y has a differe nt v ibrational
frequ-en y, and ·'rnOlecules that p roduce the sam e vibrntional frequencies have
the same smell.Turin reported th3t various ch e m icals that have predictably
s imilar vibrations beca use o f their m olecular compos ition also have simila r
s mells. For examp le, a ll ci trus odors foll in to the same vibrational-frequency
class. Bu t o ther, indep e ndent resea rchers have no t va lida ted Turin's claims
(Keller and VosshaU, 2004).
Mud1 less resea rch has focu sed on the vibration theory than on theshape-
pattem theory, so c urrent ad va n ces may unfairly bic1s our understanding .
Nevertheless, vibra tion theory canno t explain several conundnuns of olfa.ctory
p e rcep tion, s uch as specifi c anosmias and the different scen ts produ ced by
s te reoisomers, w hich sh ap e-patte.rn theory can explain.
A specific anosmla is the inability to sm ell one specific compound a rnid
otherwise nom1a.l smell perception. Specific an osrnias are due to fuu1ty odornnt-
receptor interactions, or the lack o f (or diffe rent variants of) specificORs, not
odorant vibrations. Most specifican osmias are to steroidal musk compcnmds,
and thecondition appears to be gen etic. TI1e m ost s tudied specific anosmia is an
inability to smell the compo und and.r osten one, w hich is fotmd in armpit sv..reat
and p ork. A significant proportion of the p opu.la tion has a s pecific a nosmia to
androstenonej estim.ates range between 11 % and 75%, but between 20% and
40% is m ore typical (Bremn er e t al., 2003). Interesti ng ly, a m ong those \Vh o
ca n s mell andsoste.n one, the majority find it to be unpleasant a nd " urin ou s,'1
w hile the rest d escribe it as a "sweet musky-floral'' scent . A recent cloning
s tudy of human o lfactory receptors s howed that the variability in de tection
of the od orant androsten on e, as "'"·ell as its perceived pleasantness, is dut:' to
genetic differences in O R expression betvveen individuals (Kelle r e t al., 2007).
Similarly, the diffe rent anosmia rates obtained for ad rostenon e in different
s tudies most likely reflect the fact tha t the gene governing the ability to smell
and.rostenone va ries rand omly w ithin a ny given sample of participants.
FURTHER DISOJSSION of specific sensory deficits amid otherwise nor-

mal pet'"ception-in this case vision (agnosias)-can be fcurd in Chapter 4 on
pageQ2.
Nonmus k od or an ts for w h ich s pecific a nosrn.ias have also been fo und

include the sulfur compound in asparagus-this is the " li..mny" s mell in urine
that some p eople d etect a fte r ea ting it- but in additi on to variation in the
ability to s mell th.is compow1d, the a moLmt of the compound tha t is excreted
in urine varies am ong individuals. 1l1e refore, if you don' t s mell
some thing 11 funny 11 after eating asparagus, it may be because you d on' t ex-
crete the sulforme ta boBte, not beca use you're m issing the recep tor to de tect
the odorant (Peld1at et al., 2011). If you really want to d 1eck w hether you' re
among the approximately 6% of Americans w h o a re anosmlc to aspa ragu s
pee, you'll n eed to ta ke a sniff after a friend w h o can sm ell it in her m-vn urine
and ate the sam e asparagus-vegetable course as y·ou goes to the bathroom .

OLFACTION 441
o..caivone (b) L<'an·o1le FIGURE 14. 10 Thesteireoisomeirs o-cruvo ne {a) and L-car-
vone (b) c cntain the same atoms, yeit they smell oompletely
different: o-carvone like cam.way; L-carvone. like spear-
mint. Shape-pattg-n th9ory can account for this fact.
Approximately l CYY.. o f the population is anosm ic to the scent of freesia flow-

ers, and sensi ti\dty to the sweaty-sock aroma of isovn leric acid also v aries
vastly among people, with about 6% of the popu lation lmable to de tect it at
<111 (Voc kl ey and Ensenauer, 2006). 1llese ex.a rnples all furth er how
individual genetic variability in receptor expression determines ead1 of our
own unique olfactory experiences. Altho ugh the differen ces in perception
may be sma ll, as s tated before, unless you have an identical n. . :tn, everyone
has a different nose.
Vibration theory cannot explain the presence of specific an osmias or why
the sam e od orant produces different scent sensations in different people,
but s hape-pattern theory can: these phenomena could arise fr om differing
odoran t-recep tor inte ractio ns or the absen ce of or variance i:n certain recep-
to rs . Another mark in favor of shape-pattern theory come:s from the s tud y o f
stereoisomers. Stereolsomers are molecules that are mirror-image rotations
of one an other, and a ltho ugh th ey contain all the s...'lm e atoms, they mn s me ll
completely different For exa mpl e, D-carvone(the right-handed isomer) sm ells
like c..'l.ra,vay (Figure and L-carvone (th e left-handed isomer) smells like
speam1int (Fig'Ure 14.10b). According to s hape-pattern this difference
arises because the rotated molecules do not fit the sam e receptors (as if you
were trying to put your right hand into your le.ft-hand gl ove); thus, differen t
receptors are activated for these h-vo m olecules, causing different scents to be
perceived. Vibration theory cannot explain why stereoisomers s mell different,
beca use the v ibrations of s tereoisomers are the s.ame. More recent, and m ore
direct, evidence for shape-pattern theory comes from in vitro experiments using
cloned olfactory receptors, which h ave revealed chem ical-receptor interactions
that spe.cific odorants binding with specific recep tors.
At the sam e fim ,':.\J I mo1ecn!es vib rate, and all receptors are made o f atoms
that vibrate. Moreover, recent work has shown that fruit Hies are capable of
disting u islUng between h¥o m o lecules that are identical in shape but have
different vibratio nal frequencies {Franco e t nl., 201 1). The sa me tests applied
to humans have so far failed, but a n interaction behveen. the vibration and
shape-pa ttern theories may yet be discovered for human olfoction. stereolsomers locmem (moleruloo
that can exist In different structural
The Importance of Patterns forms) In which the spatial arrange-
ments of the atoms are mirror-Image
You may have noti ced a potential discrep::mcy in h ow we' ve accotmted for rotations of cne another , like a right
odor perception. As a lready no ted, we can detect an infinite munber of od ors, and lett hard.

442 CHAPTER 14
O:lorant molecule yet mu· genes code for only C1bo ut 1000 olfactory receptors, and 600--700 of
..... .. ----··-... them are n onfuncti on al. How can we detect so m any different scents? To
s tart to see the way o ut of this conundl'um, reca ll that, in vision, we can tell
the difference beh11,1een thousands of differen t colors, even thou gh we have
only three types of cones. Each type of o lfactory receptor responds to the
s tructure of certain m olec ules onl y. However, a molecule may have features
th a t stinnJate several different receptors. Moreover, eadl receptor appears
to h ave vari ous 11 feature d e tectors 11 that contribute to furth er s pecificity in
O R activation. Thu s, in addition to differences in OR activation1 a diffe rent
FIGURE 14.1 1 1l"l9 hypothetic al role set of feature de tectors is activa ted when we smell dlOcolate or \'lli nterg reen
of OR activation timing and order. A or rose (Fig m e 14.9b).
slnQle m olecule b inds first to recep- As wi th color de tection, we detect od ors by the pattern o f ac ti vity across
tor type 1, then a split second later to various different receptor types. The intensity of an odor also changes which
receptor type 2, and then to r909ptor
3. Brains are especially well suitad
recep tors (and hen ce patterns} '"rill be ac tivated, w h ich is wh y weak a nd
to rQCognizing patterns o f responses s trong concentrations of an od oran t d o n ot s me ll qui te the sa m e. The fact
such as th B. so the number of odCfs that recep to r activation is affected by odora nt concen tration likely explains
we can rQCOgnlze gr9ally &Xceeds th9 why d ogs \vith ma ny more functio nal receptors than we have can perceive
number of types available. odorants o f cons iderably lowerconcen tra tions . TI1e timing of olfactory recep-
tor activatio n also seem s to be important; a n odorant that activates several
recep tors w ill also s timulate them in a. specifi c temporn. l sequen ce a nd speed.
Ano ther odora nt m ight sti mubte the sa m e receptors in a differen t order
and ra te, and the d iHerence might lead to th e perception of a different scent
(Figure 14.11). Thus, different odor perceptions can be due to diffe rent OR
firin g patterns, o r to firing of the sam e receptors at a differen t rate and / or
in a diHerent sequence.
l}1e flip side of the pattern perception 1nechanism is that if two o d orar1ts,
one molecularly simple and the other complex, activate the same recep tors
in the same way, we end up smelling the same th.ing. Por exam p le, the fea-
hlre detectors fo r the sing le mo lecule odorant phe-nylethyl alcohol (w+U ch is
artificial rose scent) and for a rea l live rose (whose scen t is composed o f m ore
than 1000 different m olecules) might both res ult in the sa me basic pattern o f
O R activation .md h en ce the same percep tion of "rose" (this phenomenon
sh ould be re1niniscent of metamers in color vision, d iscussed in Cha pter 5).
Patterns are also important for odor processing at early stages in the ol-
factory cor tex. Specific patterns of activity are prod uced by specific odors in
the pirifor m cortex, and odors that are perceived as smelling sim ifar produce
s in'lilarpatterns. For example, "minty" od ors like \•; inte rgreen and spear mint
produce patterns of activity that have a lot of overlap, but the pattern produced
by a "citrus'' odor like le mon d oes not overlap wi th the pattern elici ted by
wintergreen. Interestingly, distinctive patterns of activity by specific categories
of odors (e.g.? rnint)-icitrus}arenofseen fartherdmvnstream in olfactory proce.ss--
ing, such as in the amygda la or orbitofronta1 cortex Q. 0 . Howard et al., 2((19}.
Patterns may be lacking in these regions of the bra in because at these stages
of processing, person.a l and em otional associations mediate activity patterns.
Is Odor Perception Synthetic or Analytical?

8230336 Amr-.a Just as we rarely hear p ure tones outside of auditory perception experim ents,
we rMely smell "pure odorants' 1 outside of an o lfactory perception lab. Almost
a.II of the olfactory stimuli that \Ve encounte r in the re.U world are mixtures, like
the l CXX>-molecule rose scent emana ting from a flower bed that we discussed
in the previous section . How d o we process the compon en ts in an od ora nt
mixh.1 re? There are h vo brodd possibili ties: ana lysis and synthesis. Audi tory
mi xhues provide the classic example of analysis. A high note a nd a low n ote
played simultan eous ly on a piano each can be a nalyzed out of the mix and

OLFACTION 443
(a) Auditory mixture: .malysis (b) Color mix ture: synthesis (c) Olfactol}'"mixture: FIGURE 14. 12 The roles of a noJysls
.m.:l)'lsis and synthesis and synthesis in sensory perception.
We can SE:paratet,• perOOve the, thre.;i
tones of thiB musical chord being
played in (a), but not the high- and
medium-wavelength light rays mixing
in the ctfiter "M"teo we mix odor-
ants (c), W'3 perceive the mixture pri-
ma.rity syntheticaltij. but solTl9 degr99 of
anatyticaJ is possible. Ana·
lytical ability vari9S with prio r training
and with the o dorants that constitute
8230336 Am the mixture.
perceived separately (Figure 14.12a). Color mixtures provide the classic ex-
ample of synthesis. If we m ix red and sreen li5h ts, the resulting colo r that ""'e
see is yell ow (Rgure 14 .12b ). Red and green cannot be analyzed o ut, because
the h vo lights have been synthesized into some thing else. ls o Uaction {Fig ure
14 .12c} nn analytical o r a synthetic sense?
It seems tha t the an swer is both-olfactory p erception can be both ana lytical
and synthetic, although the synthetic component is '"·ha t we are more like ly
to perpetually experience. The arnt lytical qualities of olfaction are evident in
blnaral rivalry. As you read in Chapter 6, w hen a different object is presented
to each eye sfrnultaneously, you d on' t see a blend of the two objects; rather,
binocular rivalry occurs, and yo u see the hvo objects a lte rnating. A s imilar
phenom enon occu rs in hearing w hen discrepant sounds are presented to each
ea r (see Ch apter 10). lt was recently fo und that binaral rh·«1 lry in olfoction also
occurs: w hen hvo diffe rent odors are presented, one to each of our nostiils,
we alternate in our ability to sme ll one od or or the other (Zhou llnd Chen,
2009). Researchers presented a rose scent to one nostril an d simultaneous ly
an odor tlh'lt smelled like marke r pens to the othe r nostril a nd found tha t
the p articipan ts back and forth be tween saying tl1at they s melled
" ma rkers" o r "rose." The marker scent was perceived as more intense, an d
it was us ually perceived first, w h ich also occurs in binoc ular rivalry, w here
the "s tron ger-" image ten ds to d om in ate. What's interesting is tha t this rivalry
seems to occur in both the n ose and the brain. Whe n the sa me participants
were la ter presented vvith a m ixture scent of bo th rose and markers to both
n ostrils s imultaneous ly, rnost of them also reported s me lling firs t markers
and then rose and then marke rs ilgain, a nd so on . This observation s uggests
that odor rivalry is occurring in the brain as well as in the nostrils, or pe rhaps
only in th e brain.
New rese<'!rch regarding the analytica l component o f o d or p e rt..--ep tion has
also s ugges ted that as in vision , w here we have add itive and subtractive color
primaries (red, g reen, blue or cyan , magenta, yellow), o dor percep tions fall
into categories or perceptual d usters a nd there may be ten of the m (Castro,
Ramanathan, and Chenn ubhotla, 201 3). Resea rchers recen tl y reported that
usi.ng soph.is tica ted s tatis tica l a nalysis appli ed to 144 d.iverse odo rants, they
could categorize all of tlie ocloran ts i nto one of ten perceptual c.::1.tegories, loosely
labeled as follows: fragrant, woody / resin ou s, che mical, fruity (n ot lemon ), binaral rivalry Competltla1 between
sickening/ sour, minty. sweet, nutty, a nd s icke ning/sulfurous . This tre two rostrlls for odor perception.
When a different scent Is presented
is n ot a n ew idea, and in the 1950s Jolm A moore (who,as 1nention ed earlie r,
to each rostrll slmultaroously, we
first proposed the "shape" theory for odor perception) hypothesized tha t there perceive each scent to be alternating
i,.•.rere seven oder primaries: swe.. spermous, fishy, malty, urinous, rnusky, back and forth with the other. and not
ilnd ca mphoraceou s (1ike m othballs). Amoore's work was based largely on a blend of the two scents.

444 CHAPTER 14
olfactory white The dfactOl)I stu yi.ng specific a nosmias, but independent support for his theory \.Vas never
equtvalent of "v.t>te noise" cr the cdcr fou nd-until now ? Note that the prhnaries most recently discovere d a re
w l1lte. When at least 30 odofants o f somewhat different fro m those desc ribed by Amoore, so though tantalizing.
equal Intensity that span olfactcry
ptrfsloohemlcal and psychologic al the existence of od or primaries should be considered tenta tive tmtil furthe r
(perceptuaO space are mixed. they convergent evidence is obtained . Stay tlmed.
produce a resultant odcr percept that Although we are capable of discriminating thousa nds of different odors,
Is the same as eveiy othE!f mixture intuition tells us that most mixtures are perceived as tmitary ""·h oles. For
o f 30 odcrants meeting the span exa mple, the s mell of ba con is very dis tinc tive, a nd m ost p eople wou ld
and equivalent Intensity crltena, even
though the various mixtures do not
p e rceive it to be a unitary sensa tion. But, like na tural rose aroma, there is no
share any common odorants. sin gle "bacon " odo rant, and the sensation we recognize as bacon is made up
of a combination of ma n y diffe rent volatile che micals. To test the synthetic/
analytical n ature of olfactory perception, Laing and h is colleag ues (Laing a nd
Prancis, 1989; Laing and Glem a rec, 1992) conducted a series of experiments
in vvhich they asked untrained particip ants, participants w ho had received
preliminary "od or training,'' and ex perie nced p e rhm1e rs and fl avorists to
identify the constituents of mixtures cont:aining be hveen one a nd fi ve comm on
odorants. The average discrimination ra te from all the participants co mbined
was n o more than th ree compon ents in a five-component mixture.
the m ore training the participants had, the better they d id. \\lith more than
five com ponents i.n a rnixture, thoug h,even professional perfumers' an alytical
ability breaks d O\-\:n. Thus, it a ppea rs th a t olfactio n is prim arily a sy nthe tic
sense, but tha t a cert<iin a mo unt of a nalytic(1I ability can be d evelop ed .
TI1e predomim.mtly synthe tic quality of od or mixture perception was also
s upported by a recent experiment de mons trating the existence o f olfactory
white. Like w hite n oise, w h ere vari ous mi xtu res of many different frequen-
cies are pe rceiv ed as soundi ng Hke the sa m e kind o f m &1.nin gless bu zz, or
w hite in vision , w here m ixtures of many different waveleng ths p roduce
the pe rception of "whi te.'' Researchers in Israel found tha t w he n a t least 30
odorants (the m ore odora n ts in the m ix ture, the be tter, but 30 seem s to be
the tipping p oint) of equ a l intensity tha t span o lfactory physi oche mical and
psych ologica l (percep tual ) space are mixed, they p roduce a resulta nt odor
percept that is the sa me as every o ther mixture of 30 odorants meeting the
s pa n and eq u iva lent intensity crite ria, even though the various mixtures d o
n ot s hare a n y conunon od orants (Weiss e t 2012). Tha t is, three different
rnixtures o f 30 different o do rant molecules w h ere the components have the
same inten s ity a nd a re distributed across the range o f o lfac tory s timulus
space, but \•vhere none o f the od orants in the three mixtures are the same,
actu ally sm ell the same. The scent of this "olfactory w hite" has been described
quite variably (to som e it s m ells fl oral ) and is consiste ntly rated as bein g
of ne utral pleasantn ess. You m ay now be wonde ring why compl ex o d ora nt
mixtures that produ ce th e p e rceptio ns o f " bacon " or $'rose" do n ot ins tead
all s mell the same. 11'1.e a nswer is that the compone nt odorant m olecules in
"b;)con " and "rose" arom;) d o n ot s pa n the range o f olfactory s p ace, n or ar e
they of eq ual inten sity; rather, the o d orants wi th dominant inte nsi ties a nd
certain blends s ta nd out, thus pro ducing the specific perceptions o f rose,
bacon , coffee, a nd ind eed all of the specific o d or sensation s we ca n pe rceive.
Notably, th is exci ting new findin g sugges ts an underly ing comm ona lity
betw"een o lfaction , vision, and a udition-a uni fo rm perceph.1a l experience
can a rise frorn cornplex mixh.ues.
FURTHER DISOJSSION of color mixture synthesis can be found in Chap-

ter 5 on pages 129-130.

OLFACTION 445
The Power of Sniffing

When you s mell something passive ly, you are breathing in air containing
odorn nt m olecules and detecting th ose m o lecules. By contrast, wh en you
sniff, you me con scious ly and forcib ly inha ling bms ts of air into your n os-
trils. S.niffing increases the ability to de tect o d ornnts. Sniffing also produces
greater activation in some parts of the brain, s uch as the cerebellum, than pas-
sive inhalation d oes. Exci ting new researc h is curre ntly exploring how a ctive
sniffing may enable severely physically disabled people to commt.m..icate with
th e o utside ""'orld and to move independently. r' Locke d-in syndrome# is a
horrendous neurological condition cha racterized by comple tely intact cogni-
tion and rnentality but to tal physical paralysis, s uch that afflicted people are
li terally locked in their own bod.ib ::Brail\-cQmpi1te?, interfares would be an
ideal solution for s uch individuals, but they have not yet been s uccessfully
implemented, and up to n ow these patients have ha d to rely on b linking as
the sole m ea ns to conununicate (it was recognized tha t the eyeblink response
isstiU intact in these people).
Active sniffi ng hLls now .:tlso been discovered to be p reserved in locked-in
syndrome. Trudngadvantage of this intact physical hmclion , researchers recently
shO\ved that by using precise, de llberate sniffing-changing sniff magnitude,
durati on , onse t, and offset-l ocke d-in p atien ts can be s uccessfu lly trained
to control tex t-writing software and meaningfully conununicate \'\ith loved
ones. Us ing the sa me type of s niff technology, quadriplegic patients ha ve also
been trained to m ove their w heekh.a.irs in complex circuits by us ing their nose
al one( Plotkin e t al., 2010). This may be the dawn of a nasal ly p owered world.
Odor Imagery
On e area w he re olfacti on and om other senses diverge is imagery. We know
that vis ual and auditory imagery is easy and readily accessible a nd, tho ugh
in somevvhat different ways, imagery is also evident in touch (shivers, ting les,
phantom lim.b) and h1ste (e.g.1 the sotrr salivation reaction). By contrast, humans
appe..1r to have Httle or n o ability to conjure "odor images." For example, you
can probably see the visual im..1ge of a Hershey's ch ocolate kiss in your mind's
eye right now (you might even s tart salivating). But can you really rep roduce
the s mell of d-.. ocolate in yo ur " mind "s nose"? Brain-imag ing studies (e.g .,
Kosslyn e tal., 1995) have shown that many of the parts of the brain that would
be involve d in actua lly seeing the kiss are also involved in vis ually imaging
it; wi th olfaction, however, simi lar studies s uggest that the degree of overlap
between smelling an odor and ;'irnaging'' it is mud-.. weake r (Djordjevic et aL,
2005). Drea1ns 1,vith olfac tory sen sa ti on s are also very rare (Carskadon e t a L,
1989; Zadra, Nielsen, and Donderi, 1998).
Animals, s ud1 as rodents, tha t rely predominantly on s mell as the sense
by w h ich they n egotia te the worl d may well think and dream in s mell.
H owever, beca use we do not think in sme ll, it is not n ecessary to h ave stored
representations of olfactory exp e riences. Neverth e less, re cen t hmctional
neu.roimaging resea rch suggests that alth ough o dor imagery among average
peop le is very weak, exp e rt perhune rs may be able to acquire the abili ty to
image od ors throug h training. Function al magnetic reson an ce imaging (fMRl)
sh.owed th a t p erfumers produced activation in their pirifom1 cortex w hen
asked to image odors, and the greater their p e rfumery th e more
their brains appeared to be reorganized to a cconunodate this tas k
Delon-Martin, and Royet, 2011). This is another exa mple of h ow the olfactory
system is extremely flexible and capable of ne ural reorganization on the basis
of experience and learning.

446 CHAPTER 14
Olfactory Psyc hophysics, Identification,

and Adaptation
psychophysics The sdenC<J of The subfield of psyd,ology called psychophysics was in troduced in Chapter
defining quantitative relationships 1. 1lle go.:'ll of the psychophysidst is to quantify the psychological experience
between prryslcel and psycholo;Jical of our sensory world. In th is section some of the nuts-a nd-b olts quffitions
(Subjective) events.
addressed by o lfactory psyd'lophysics a re discussed, and then we'll go on to
consider how we identify and ad apt to odors.
Detection, Discrimination, and Recognition

A lthough we can theoreti ca lly d e tect over a trillion o dors, a number of fac-
tors limit m u d etection ability ln reality. One m ajor factor has to do w ith how
much s timulati on is required before we p erceive some th ing . As wi th o ther
senses, our olfactory detection thresho lds depend on a nmT1be r of factors. For
instance, odorrult m olecules with longer carbon chai ns, s uch as vanillin, are
eilSier to detect {have lower detection thresho lds) than those wi th shorter car-
bon chains, such as acetone (o therwise known as nail polish remover). O ther
limitations have to do wi th individual d ifference factors as you w ill see soon
in the '1ndividua l Differences" section.
1l1e abili ty to detect an odoran t can also be m ani pulated by experien ce.
Serendip itous observation revealed that throu gh repeated testi.ng, sensi ti v-
ity to androstenone ' the s teroida l musk compo und th at m an y people ha ve
a specific anosmia for) ca n be i.ndt11..""ed in ab out h..,lf of the people wh o are
initially unable to detect it (Wysocki et al., 1989). That is, a proportion of the
people who are a nosm ic to this particular chemical develop an ability to smell
androstenon e through repeated exposure. From other a reas o f bi ology, we
know that gen es can be "turned o n" by environmental factors, a nd because
eild1 o lfactory receptor is coded for by a specific gene, it is con ceivable that
the receptors for detecting androstenone ca n be activa ted th.ro ug h repeated
presentation of the chenUcal. Other studies have shown that in creased sensi tiv-
ity to som e common odoran ts, such as benzaldehyde (ch eny-alm ond arom a)
and Citralva (lem on-ora nge scent), can be induced through repe.."t ted exposu re
to these che micals, pMticularly am ong fema les (Dalton et al., 2002i Diamond
et al., 2005). Purthennore, s tudies in fruit flies have shown that genetica lly
hardwired OSNs can be altered by exp osure to specific ch em icals (Sachse et
al., 2007)_ Plasticity and modulation through environmental influen ces appear
to be basic principles in olfaction.
A healthy person can discriminate- tell the difference between-a huge
number of odors. Note, however, that is not the same thing
as recogniti on, the ability to remember w hether or n o t we've s me lled an
odor before. It takes being exp osed to up to th ree times M many o do ran t
molecules to recognize an odor as it d oes to simply detect the p resence o f an
odor. You've probably expe rien ced th is phenomenon yourself: you registe r
that yo u s me ll sornething before you know wha t the s m ell is. Interes ting ly,
we d on' t need to know w hat a s me ll is, or be able to name it, in order to
recognize it or even to have a memory triggered by it. The only requirement
is that we hnve previo us ly en countered the scent-that is, that the odor is
familiar (C leary et aL, 2010; Herz and Cupch ik, 1992). This is diffe rent from
other sensory experiences amo ng cognitively hea lthy adults. For example,
\\.·e wouldn 't be able to say that we 11 recognize," respond appropriately to,
or have a m em ory elicited by somethin g that vve see wi th out h avin g sorne
kind of label 1or lt, even jf tabel isx.ery vague or idiosyncratkr s ud1 as
"food " or " uncle's weird hat.1' Brain-damaged patients \vi.th visual agnosia

8230336 Amma Appa OLFACTION 44 7
80
,,! 70
60
30
RetO!ntipn intervals (days)
FIGURE 14.13 Long-t&rrn for smi;olls . Th9 first point on this graph, at al::out
70% . indic.ates odor recognition accuracy with only a 3()-second dgay from learning ,
Note. however. that this accuracy lewl is the same after a week and has dropped
only about 3% aft9r a month. This ret9ntion rat9 rQTiains re.lativ9ty constant owK inter-
vals at least as lo ng as onei year. (Frcm Engen and Roos, 1973.)
are the exception in th a t they o ften know how to use an object even if they
d on 't know w ha t it's called.
Another interesting fea h1re of odor recognition is its durability. ln controlled
experiments1 a 30-second d elay between o...i or presentation and testing produces
a precipitous drop in recogni ti on but w h at \ Ve re m ember after 30
seconds is very close to '"·hat we remember after 3 days, a mo nth1 o r even a
year (Figure 14.13) {Engen, Kuisma, and Eimas, l 973i Engen a nd Ross, 1973;
Murphy e t al ., 1991; Rabin a nd Cain , 1984)_ Ag..Un, you probably recognize
this phenom enon from yo ur own life. If you s melled a cert.i.in pe rhime onl y
once '"'hen you '\>Vere a child, you ca me upon that perfume again 20 years
later, you might very well know that the odor was familiar.. O ur mem ory for
staircase method A psychophysi·
odors is even m ore resilient if the initial exposure is accornpanied byen10ti on cal method fcr determining tile con-
(Herz, l 997). centration of a stlmulus required for
detection at the threshold level. The
Psychophysical Methods for Detection and Discrimination staircase method Is an example of
Resea rchers wanting to measure h ow perceptually sensitive people are to
a method of Hmlts. A stimulus (e.g.,
odorant) Is presentoo In an asoondlng
odors, or w he ther they can discl'imina te one od orant fro m an other, u se vari- concentration sequence until deta::::-
ous psychophysicaJ methods. A common o lfactory techniqu e fo r d etem1ini.n g tion Is Indicated, and then the con-
som eone's odor de tection thresh old is called the staircase me1hod (or 1'reverse centration Is shifted to a deooerdlng
staircase me thod "), and it is a n exa mple of are know n as " metho ds of sequence until the response charges
to •no detect Ien · This ascending
limits" (see Chapter 1). There are various versions of the s taircase m e thod. In
and descending sequence Is twically
a ty pical procedure, an odorant is presented in ever-i.ncreasingconcentration repeated several times, and the oon-
increments until the participant reports bein g able to "sm ell something" for centratkms at which revemals occur
severa l repeats of a con centration. Then the odora nt's concentration is d e- are averaged to determine the thresh·
creased increme ntall y w1til the participa nt reports no d etection . old detection level o f that odorant for
a lndMdual. AJso calloo ra'"""3e
These reversals are repeated a number of times, and the odorant con cen- staircase methoo.
trati ons at the p oi nt where reversals occur are averaged to d e termine the
triangle test A test In wh b h a
approximate concentra tion need ed for that person to detec t the odorant. As a participant Is given three odorants
rev-ersal point is re..1 ched 1 th e increments by which an od orant's concentration to smell, o f which two are the same
is raised a nd lmvered can be fin e-ttmed for precision. Staircase me thods ca n and one Is different. The paitlc lpant Is
be used to determine a genera l benchmark for o lfacto ry sen sitivity. They can required to state which Is the cdd cdor
also be used to d eter mine d e tection thresholds across a range of odors to test out. Typbally, the order In which the
individuals in specific ways. three odorants are given (e.g ., same,
same. different; dlffernnt, same, same;
To d etermine w he ther som eone can discrimina te between two cx-iorants, same, different, same) Is manlpueted
the m ost common psychophysical test u sed is called the triangle test. In a and the test Is repeated several times
triangle test, a p articipant is given three odo rants to smell, of which t¥.:o are the for greater aoourae')'.

448 CHAPTER 1 4
same and one is diffe l'ent. The participant is required to s tate whid1 is the odd
odor out. Typically, the order in w!Ud\ the three odor.ants are given (e.g., sa me,
same, different; different, same, same; same, different, same) is rnanipulated
and the test is repeated several times to establish accuracy.
Identification
Attt1ching a verbal labe l to a smell is a step beyond odor recognition . Olfac-
tion has a th sense" because we are so often lost for words
to descr ibe our olfactory experiences (D. Ackerman, 1990). All of us have had
the experience of not be ing able to come up with the name of something we
know; for example, what was the n::ime o f your fourth-grade teacher? This
experien re is known as the " tip-of-the-tongue" phen omenon. In the olfactory
d om.ain, s uppose you Ut ke a sniff of some thing from a bottle that prov ides no
visual clues to w h::it it contains and you immediately know that the scent is
extremely familiar, but you jus t can't come up wi th the name fot it. Borrowing
frorn the verbal scenario, we call this experience the tip-of-theRnose phenomR
enon (Law less and Engen, 1977), and it can be very frustrating.
There are some import<m t differences behveen the tip-of-the-nose state
and the tip-of-the-tongue s tate. For o ne, in the tip-of-the-tongue s tate we d on ' t
know the exact word we're looking for, but we might know its firs t le tter, its
general word configuration, the number of syllables in the word, and so on
("Starts Mth a K, two syllables, sotmds like a sand\-\ich rol l ... aha, my fourth-
grade teach er was tv1rs. Kaiser! "). By contras t, in the tip-of-the-nose s tate, we
typically kn ow n othing abo ut the label we' re searching for. Up until very
recently anthropologists th ought that in all languages there were fewer word s
thatrefe.r exclusively to o ur experience of s me lls than there o:ire for any othe r
sensation (Classen, Howes, and Synnott, 1994). In English , aromatic, fra grant,
pungent, and s tinky pretty mud1 exh aust the List o f adjectives that specifi-
cally describe olfactory s timuli and n othing else. Typically people use words
referring to the perceived SOUl'Ce of the scent 1-ike lem ony or ca tegory te rms
like floral or fruity. We a lso borrow terms from other senses (chocolate s mells
sweet,gmsssmells gl'ee:n, and so on) . New eddenre, however, s uggests that
tho ugh an impoverished olfactory vocabulary is still the a 1se for m ost bn-
guages, at leas t one language is the exception to the rule. The language of the
Jahai, a n omadic hWlter-gatherer society who live in the Malay Penins ula (the
rnotmtainous rain fores t that nms beti."'-een Myanmar, Malays ia, and Thailand),
has specific and discrete words for a panoply of olfac tory sensations, such as
their word p?us (pronotmced "pa-oos" ), which describes wha t dead branches
and some types of fur and feathe rs smell like (Majid and Burenhult, 2014). In
o ther words, the Jahai appear to be able to isolate basic o lfactory prope rties
the way we can isolate the color green from a leaf or a lime. It is not yet known
why the Ja hai have s uch an expanded olfactory vocabulary, but it is knm...TI
that scents play a very important role in their daily life, so learning and es-
tablishing an exact vocabulary for olfactory experiences s uch as the scent of
tiger versus 1nonkey sca t, or a poison ous vers us nutritious \·Vould
assist much more with their survival than it would for us.
tlp-ol-1he-nose phenomenon The
lnablllty to name an odor, even thcug h It is n ot full y knmvn why olfaction and language are generally so discon-
rt Is very familiar . Contrary to the tip-of- nected . Various possibilities include the fact thL1t unlike what happen s in othe r
the-tongue pherornen::n, cne has no sensory systems, olfactory i.nforrnation d oes no t need to be integrated in the
laxlcal access to the name of the OOor, tha lamus prior to processing in the cortex, and it is argued that the thalarnus
such as first rh)1Tle. number of has relevance for language. In addition, a large body of evidence indicates that
ard so on, when in the tip-
of-the-nose state. This Is an example the majority of olfactory processing occurs in the right hem isp here of the brain,
of how larguage and olfactory perc ep- whereas language processing is known to be dorninated by the left he misphere
tion are d0€PIY disconnected. {see Royetand 2004, fora review). Convergent data frorn brain-imaging

OLFACTION 449
studies have also sho'WTI th.at conscious odor perception interferes with language
That is, processing o f odors ll.nd processin g o f w o rds compete for
thes.ani.ecortical resources.. This meansthats imultaneous ly presenting an odo r
and a word \vill lea d to impaired word encoding s uch that namin g something
at the sa me time as s me lling an odor is espe cially difficult. Like,vise, processing
words a t the same time as s melling diminishes the pe rceptual quality of the
odor and the odor srnells weaker (Lorig, 1999; Heatherbell, and White,
2002; Walla et al., 2003; Walla, 2008). (See also Web Essays 14.1: Olfactory
Lateralization and 14.2: Verbal-Olfactory Interactions.)
Individual Differences
We do not all perceive all odors the same way oc e:qua lly actftely. One.of the
reasons for these differen ces is indi";dual variatio n in how many and w h ich
OR.s are expressed in our olfac tory epithelia (as discussed in 'The Genetic Basis
of Olfuctory Receptors'' earlier in this chapter). Genetic va riability accounts for
differences in pe rception among people wi th a "no mlal" sense of s mell and
those with a ''specifica nosmia." However, there are tv•oother major individual
difference facto rs that affect our o lfactory capabilities overall: age and sex.
Sex differences are reliably dem o ns trated in od or perception, \.vi th women
typically o utperfonning men on all meas ures (e.g., identificatio n, detection,
discriminatio n) a t all ages (Boesveldt et al., 2011; Doty and Cam eron, 2C09).
Otuing a woman's reprodL1ctive years, it seems, fe male su periority is medi-
ated by h orm ona l fluctuations, \·vHh women being especiall y se ns itive to
odors during ovulation but no diffe rent from men during m enses (Doty et
al ., 1981). However, a female a dvantage is also seen before puberty and after
menopa use; thus it has been speculated that endocrinologic effects exert an
organiz ing influence on the ner vous system during early development s uch
that baseline sex differences in olfactory sens itivity persist throu ghout the Llfe
span (Doty and Cameron, 2oo:J). Contrary to popular belief, however, research
has s hown that o lfactory sensiti v ity is n ot heig htened during pregn ancy
(Came ron, 2007; Hummel et al, 2002; Lask.a et al., 1996). Taste is another s torv,
as we'll see in C hap ter 15.
O ur D.bi!ity to d etect odorants declines with nge, because o f a ch.."l.nge in the
proportion of ce ll regeneration to cell in ORs. As we age, the number
of odorant receptors th at die off rises beyond the number that are regener-
ated (Kern e t al., 2004). This ratio continues to w orsen in fav or of cell death
as we grow o lder1 s uch that after the age of 85 it is estimated that about 50%
of the p opulation ha s effectively becom e anosmic (Hummel et al., 200'7i ]. C.
Stevens and Cain, 1987). Trigeminal pe rception d eclines \vi. th age as well,
and seems to be directly related to loss in olfactory function (Hummel e t al.,
2003). Sensitivity to tasta:nts dec.J"eases as we ge t older too, but not all tas tes
s uffer to the sam e extent; sour seems the most affec ted and swee t the least
(Boesveldt e t a l., 2011). These declines in chem osensory function explain the
increased use of condimen ts among the elderly (especially salt), w hich rnay
produce unhealthy outcomes.
Similar to detection, our ability to identify odors is best between our teen-
age years and otu 40s, but we are in our 50s it s tarts to decline fair ly
precipitously, sud1 that by age 65, about half of the poptJation have notice..1bly
impaired olfactory identification ability. For people age 80 or o ld er, abou t
75% s uffer from major impairments in odor ide ntification (Doty et al., 1984)
(Figure 14.1 4). A main reason for this sharp decline is that odor identification
is s trongly influency-d by verbal a nd semantic processin g. It is well esroblished
that as '"''e continue to age in adulthood, our ability to nam e everything gets
worse (Au et al., 1995). Because there is already a weak connec tion behveen

450 CHAPTER 14
FIGURE 14.14 Changes in o lfac -

tory ide ntification ablity as a functb n
o f a ge. Our ability to cprrectly identify 161 90
OdC>r$ peaks in our 20s and stays rela-
tlvety constant until o ur mid 50s w hen
71
it s tarts to d9clini;i. Mal9S tf!fld to have
sl ightly lowe< scores than females at
all ag9S.. How.wer, there is gr90.t deal
o f lnd i\lid ual variability and a number of
octogenarians still have high function.
1f4.ftG1r Doty et al. , 1Q84 .)
.
·' 15
-
-
....... Fein.ales (11 = 11:8)
Males (11 = 'JW)
Total group (11 = "1955)
pa
:D-29 40-49 W-69 S0-1>9

Age group
lan guage and olfaction, any increase in the v ulnerability of our narning abil-
ity is most readily observed in olfactory identification. That is, these "senior
mo m ents" are m os t obvious w ith o lfac tio n beca u se it' s ah\'ays been harde r to
identify the scent in the spice jar that is 11 rosemmy" than to MY that the name of
that guy in the movie is 11 Matf:hew McConaughey." Notably, h m,,.ever, people
wi th higher ed u cation levels (more yea rs of p ostsecondary schooling) have
bette r odor identificati on abili ty as they age tha n people \<\>i th less ed u ca tion
(Boesveldt e t al., 2011 ). TI1e correlation is presumably due to an associa ti on
betv-.·een education and naming abili ty or verb al fluency- the more ed u m tion
you have, the better your vocabulary and verba l fluency-and this buffers the
odor identification declines of aging. Notably, age-rela ted declines in ability are
not nearly as pronounced for odor detection and discri rni.nrition as they are for
identification, because they rely mud 1 lesson sen1antic processing (Hedner et
al.,2010). It s hould be mentioned, thou gh , tha t just as wi th hearing and vision,
there is con siderable indi vidual variability in age-rela ted declines in odor
perception, and there are m an y oc togenarians \·vi th excellent olfactory ability.
TI1e connectio n behveen namin g a nd olfactory identification turns out to
be very important in identifying certain ne urologica l diseases a t their earli-
est stages, in particular Alzheimer' s d isease. This is because AD is a disease
characterized by loss of memory, especia lly sem a ntic rne mory (mem ory for
the names of things and factual infonn ation), and s ince, as alrea dy explained,
odor naming (corn pared to naming any thing else)is most v ulnerab le to s light
cognitive p erturbations. Testing for s uch a deficit is crucial, because the sooner
peop le are correctly diagnosed wi th Alz heimer's disease and treated, the bet-
ter their quality and duration of life w ill be. (See "Sensation & Perception in
Everyday Lile: Anosmia" on pages 432-433.)
Adaptation
Have you ever had the experience of noticing coworkers or classmates w ho
seem to be po ming a bo ttle of co logne over their head every m onUng. leav-
ing you choki ng on the overpowerin g nroma? Can't they smell anything? O r
perhaps you've n oticed tl1nt, after having a bottle of perfume fo r a few months,
you can't sm ell the fragrance in it anymore. Or you've gone away on vaca tion
and retunied home to find that your house seei11s to have a "funny" smell that

OLFACTION 451
it didn' t have when you left. \Vhat's going on ? The answer has two parts: the G protelr>-eoupled receptor
first involves the nose, and the second involves the mind. (GPCR) Any of the class of recep-
The sense o f smell is a change d etection sys tem. When a new chemical tors that are present m the surface of
cifactory sensory neurms. All GPCRs
con1.es along, yo ur olfactory receptors fire in response to it, and yo u perceive are charactertzed by a common
a scent. For ex.ample, when you first enter a bakery, you n otice the mouthwa- structural feature of seven membrane-
tering arom as of the cakes, cookies, pies, and brea d s. But if you stand in.side spanning a-he Ces.
inspecting the s\oV-eet baked goods for a. while, you may find that by the time receptor adaptaUon The bk>-
you've p icked out what cake you want for d essel't, you can n o lon ger s mell chemical pheronenm, occurrlrg after
it.\ Vhat has h appened is that the odorant m olecule.s that make up the bakery contlnuaJ exposure to an odorant.
aroma have bound to the corresponding olfacto ry sen.sory ne urons in your whereby receptors stop responding to
an odaant am detection ceases.
n ose. When this h appens the O Rs retreat into the cell body {Firestein, 2001 ).
The receptors are the refore no longer physically available to respond to the
bakery scent m olecules. This response is a process in "receptor recycling.1'
Specifically, od orant binding to an OR 01 uses the OR to be internalized into
its cell body, where it becornes unbound from the odorant and is then recyded
through theceUand emerges again in a number o f minutes. Receptor recycling
is a mechanism common to all receptors in the class to which ORs be lo n g:
G protein-coupled receptors (GPCRs).
FURTH ER DISCUSSION of parallels to visual adaptation can be found in

Chapter 2 s tarting o n page 4 7.
This process i.s ca lled receptor adaptation. The precise len gth of time
req uired for ad apt.'l tion varies as a function of both the indi vidua l (Da lton,
2002) and the o do rant (Pierce et al_, 1996). On average it takes about 15-20
minutes o f continual expostue to an odorant for the m olec ules to s top eliciting
an olfacto ry response, but a daptation ca n also occ ur in less tha n a rninute.
Recep tor adaptation can also be und one relatively quickly. Stepping outside
the bakery for a few nUnutes g ives unbound olfactory recep tors a chance to
acc umulate on tbe cell surfuceagain, so when you wa lk back in, you can enjoy
the appetizing scents once rnore. The magnitude of adaptation is also affected
b y od or intensity {Kado hisa and Wilson, 2006). As the concentra ti on of a n
odorant increases, the degree o f adaptation d ecreases. For example, it takes
lon ger to adapt to the aroma \¥afting fronl an app le pie baking in the oven
than to the scen t emana ting from a cool slice of pie on the kitchen counte r.
Th is is beca use m ore rnolec ules of apple pie aroma become volatile and a re
thus available to acti va te ORs when the pie is hot than when it is cold .
One W<-' Y to prolong the effect of smelling a scent before adaptation kicks
in is to dispense an odo r internUttently. For example, bursts of air freshener
alternated w ith no scent to\--ill draw out the time before your receptors are
s mothered by the air freshener m olecules a nd duck for cover.
Dalton Ci996) also showed that presumed. danger can have an effect on
adaptation rates. In one experiment, half the participants were told that a n
odor they were being exposed to was "healthful,'' w hile the other partici pants
were told the od or was 11h..'l.Za rdous." Twenty minutes after initial exposure,
the participants sm elling the s upposedly healthful od or had a dapted to it,
whereas the participants who tho ught they were s me lling a haz..'U"dous che mi-
cal actually became sens itized-they reported the s mell as even m ore intense
after 20 minutes than at the s tart o f the experiment. H owever, when these
participants were given a psychophysical tes t of odor d etection, it was s hown
that they had adapted just as the people who were told th at the o dor was
healthful had adapted. Fear of odor exposure can even make peopl e perceive
an odo tbat d - n t e · h(Engen, 1972). Fo r example, people who be lieve

452 CHAPTER 14
cross-adaptation Th3 reduction tha t a factory mus t be e mitting r'da ngerous che micals" frequently complain
In detection of one cdcrant followlrQ that they can smell a 1:nalod or corning from the fac ility, rega rdless of w hat the
expC6Ure to another odorant. Cross· factory p rod uces or whether it is even ope.rating.
adaptation Is presumed to occur
because the ocmp:nents of the odors New findings shed light on why psychological factors often sup ersede
(cr odcrants) in questbn sh•re one physical reality in odor perception. Recent research (Krusemark et al.., 2013)
cr more otfactor1 receptors for their has s hmvn that when we are anxious, initially ne utral odors becom e perceived
tranOOuction, but the order In which as tmpleasant and these now negative odors corresponding ly elici t a ug mented
odorants are presented also plays a responses in highe r-order olfactory a nd e moti onal processing cente rs in the
role.
brain (orbitofrontal cortex and pregenua l anterior d ng ula te cortex). M ore-
over, anxiety stren g thens the connection between olfactory processin g in the
orbito frontal cortex and e motion .al processingln the:an,lygdala This-s uggests
that w hen we are worried tha t an odor may harm us, we perceive it as more
unpleasant, and it elicits more-intense emotional processing and potentiates
forthe r negati ve e motiona l response to it. For example, if you smell an odor
co ming from a factory that you believe is producing dangerous chemicals,
tha t odor will smell bad to you even though that same odor could be one you
like in a different context. Em o tional hypersensitivity can also create olfactory
illusions, and you may believe that you are smell in g the 1'bad " odor, merely
w hen seeing the factory, even w h en no odor a t all IB being emi tted (En gen ,
1972; Herz and Von Clef, 2001).
O ne of the benefits of o lfactory adaptation is th ..1t it en.."lbles us to filter o ut
stable background odors, and th is filtering a bility can be enhanced through
active s n iffing-ta king d e libera te, quick inha lations (Kepecs, Uchida, a nd
Mainen, 2007). Sniffing makes OR neu rons less resp onsive to st<Jble odors
and rnore responsive to new od oran ts (Verhagen et al., 2007). For exam p le, if
we're a t a car dealership and think we smell sornething bu ming, we typically
engage in active sniffing to (1) see if we're right and (2) detenni.ne w hlch BJ\ofW
is s mo lde ring. In vision a nd hearing, perceptual stimuli in the fo reground can
be segregate d from stimuli in the background thro ugh spatial ana lysis of the
scene. In o lfa.ction , spa tia l analysis is compromised because backgroun d an d
foreground od ors merge in the air. H owever, p rovided th at background and
foregrou nd odors are at least biieAy separated in time (new-car s mell firs t and
then the scent of bmning plastic), sniffing enab les u s to sepa rate compon ents
of a n olfactory scen e.
ln som e cases, exposure to on e o dorant can raise
the odor d etection thresh old for a second,com pletely
differentodorant. For example, w hen you're picking
out a perfwne in a depat'hnent store, yo ur nose may
become fairly useless a t differentia ting the fragrances
after severa l sampl es, despite the salesperson 's
insiste nce tha t the perhunes are quite different from
each othe r (Figure 14.15). 11-Us phenomenon is called
cross.adaptation, and it is prestuned to occur when
components of the o d ors in question, or s pecifi c
odorants, rely on si milar sets o f olfactory receptors.
However, this s imple explanatio n is complicated
by the fact that m ost cross-a da ptation relationships
are n onreciprncal . For example, sm e ll ing p e ntanol
FIGURE 14.15 Do th9S9 frve fragrances smell the

same? No, but b9c31JSoB of olfac tay cross-adaptation,
if you've smell-3d four of in succession, the subtle-
ties of th9 fifth one may b9 lost to your nose.

OLFACTION 453
(a chemical used in some paints) seems to have a s trong cross-adapting ef-

fect on next sm e lling propanol {used as an antiseptic and solvent)1 whereas
6rs t has 0nly a small cross-adapting effect on then sm elling
pentanol (Cai n and Engen , 1969). Furthermore, exp osure to the firs t od orant
can sorn etim.es enhance sen si tivity to the second o dora.nt. (See Web Activity
14.2: Odor Adaptation and Habituation.)
Regard less of w hy they occur, effects usually go away
after a few minutes. Profes.siona..I perfumers, w ho may have to smell hundred s
of scents per d ay and thus don 't have a few minutes to spare, use a trick of
sniffing thei r bare arm or cotton shirtsleeve between s melling odorantsj d o-
ing this effective ly dears the nose even a t a fas t pace o f odorant presentation .
Nobody knows why this works, but it does. 11\e n ext time you find you rself
s ufferin g from munb nose in a depa rtme nt store, try it.
Cognitive Habituation
Receptor ad apta tion explains w hy you lose the d eliciotLS aroma of the bakery
after you ·ve been in the s tore fo r a \'\.·hile but can s mel.l it again after a s hort
break outside. If you took a job a t a bakery, however1 a different process would
take place. This is the phenomen on tha t your friend ""·h o ca n' t s m ell his own
cologne i.s experiencing, a nd it's the reason w hy you d on ' t s mell the ambien t
scent o f your own h om e wi..less you go out o f to-wn for a couple of weeks. It
is a psychologkal effect ca lled cognitive l1abituatJon.
In s ho rt, when we Jive with a n odor, we cognitive ly habituate to it a nd no
lon ger react to it1 or we show a very dirn.inis hed response to it . For example,
textile workers exposed dai ly to acetone exhibited acetone detection thresh olds
tha t were eight times higher tha n thooe of a comparable gro up of control subjects.
H o·w eveJ, thresh olds to another che mical to w h ich neith er gro up
had been regula rly exposed, were no different for the two groups (Wysocki 8230336 Amma Appa
e t al. , 1997). \Ve habitua te (that is, our rece ptors adapt) to som e d egree to
s timuli p resen ted to all our senses (e.g ., you s top hearing the ticktock of a
gran dfathe r dock a fte r bei ng in the room fo r a w hile), but a ttention can bring
u s o ut of habi tuation with every sen se except smell. Unlike receptor adapta-
tion, w hid1 ca n be und one in a few minutes, cogni tive habituation req uires
weeks to reverse, even for pungent trigeminal s timulants like acetone (Da lto n
e t a l., 1997; Wysocki et al., 1997). Por example, if you stopped wearing your
cologn e for 5 d ays, you ""'ould .s till not be able to s mell it well once you put it
back on . But if you abstained for 2 \.\-'eeks or more1 you wo uld.
Dalton (2002) suggested tl>atat least three mechanisms could be involved-
either singularly or in combination-in producing olfactory habituation. Firs t,
the olfactory receptors tha t are internalized into their cell bodies during od or
ad aptation may be more hindered after continual exposure and take much
longer to recycle than they n onn ally would . Second, from continual exposure,
odorantmolecu les m ay be absorbed in to the bloodstream and then transported
to the olfactory receptors via nasal ca pillaries when we breathe out through
the n ose. As long as the od orant chemicals remain in the bloodstrea m, we
will be cons tantly adapted (Maruniak, Silver, and Moulton, 1983). Finally,
cogni tive-em oti onal factors, llke those d e m onstrated in the experiment in
which participants were told tha t a n odor was h a rmful and then perceived
that they did not adapt, may be involved (but in the reverse di rection) in
cognitive habituation. cognitive habituation Tl1e psy·
Another feature of o lfactory perception that hi ghlights the imp ortance of chologlcal procsss by which. attar
long-term exp:)Sure to an odor, one
conscious pe rceptio n is that we cannot sm ell while we' re asleep (Carska d on no longer has the ability to detect that
and He.rz, 2CI01). Unlike what happens with auditory stimu.li1 w hen trige minally cdor or has very diminished detection
activating odor.ants such as menthol and pyridine (also a d1er11.ical component ability.

454 CHAPTER 14
odor hedonics TheJiklnQ dlmenslal of sm oke) were p resented, even at high concentrations, to participants in slow-
of odor r;eroaptlon. l)lp!calfY meastKed '>'ave sleep (S\VS; deep sleep) or REM sleep, they did not awa ken or show
by ratings of an odors's perceived any electroencephalograrn (EEG) sleep pattern changes. More recently, it \"·as
pleasantness. ard Intensity.
fo und tha t expos ing s leepe rs to a rtificial sm oke during all stages of s leep had
no affect on arousal frequen cy o r EEG activity (Heiser et al. , 2012). These find-
ings underscore the .serious ness of using auditory sm oke detectors, and why
o ur sense of s mell cannot p rotect us from s moke inhalation and consequent
catastrophes w hile we s leep . In s pite o f these findings, a fon cti onal magn etic
resonance imaging (f11RI) s h1dy rep orted that p resenting a n odor during S\.VS
that had previously been present during an awake learning tas k produced
hippocampal d1an ge.s con sistent with those observe d in memory consolida-
tion (B. et al , 2007). Ho\·vever, the odor had no effect if it \"'as presen ted
during o ther sleep s tages, including REf\.'I s leep. This finding .suggests that
od ors may be detected by the brain during S\VS, but this d etection rnay no t
re.suit in overt arousal It is also known from ne uroim agin g research on awake
subjects that durin g smell tasks, paying attention to odors alters brain activ ity
and odor d etection ability (Plallly et al., 2008; Zelan oet al., 2005). \ Ve use more
of o ur brain and can sme11 more acutely w hen we conscious ly focus on s mell-
ing than w hen \.Ve d on' t. It may be that,, because attention is.effectively cut off
during sleep, so is o ur abili ty to respond to od ors. A further tantalizing piece
of eviden ce supporting the difference beh'\o·een olfaction during w ake ver sus
s leep states comes fr om the recent find ing in mice that g ranul e cell activity in
the olfactory bulb during s leep is s low, consistent,, and tuned to the brea th ing
cycle, whereas during \-vaking it exhibi ts much stronger, wilder, and spontaneous
firing, Tha t is, gra nule cells, w hid 1 may be the ha.sis o f odornnt identifi cation
operate in a consistent fa shion independent of the od or env ironment during
s leep, but by contrast appear to respond to specifi c o d or an ts as they enter the
environment while we are (Caz..'lkoff et al., 2014).
Olfactory Hedonics
The m ost immedi1Jte and basic response we have to an odor is whether \>Ye it
or not. Such affec ti ve e valuations are knom1 n.s odor hedonlcs. Jn tests of odor
hedonic evalu ation, people are typically asked to rate ho""' pleasa nt, fru1liliar,
and intense a gi ven odor is. TI1ese measures are then used to d etermine the
h edonic value of a specific s me ll. It is obvious that perceived pleasantness is
related to our liking for an cxior. But how are familiari ty and intensity related?
Familiarity and Intensity

As with ma ny other facets of life, \ •le tend to like odors tha t we've s melled
many times before. That is,, we tend to like familiar od ors better than unfa-
miliar odors. Moreover, we often perceive pleasant odors as familiar, even if
we ha ven 't smelled th em before (Moskowitz, Dravnieks, a nd Klarman, 1976;
Sulmont, lssa nch ou, and Koster, 2002). Thus, ratings of odor pleasanh1ess and
fami liarity s how a linea r relationship with od or liking.
Intensity has a m ore complex relati onsh ip to od o r likin g th a t is often
represented by an inverted-U function, but this depends on the odorant. A
rose scent1 like pheny lethyl alcohol, may be evaluated as more posi tive with
increas ing intens ity, up to a p oint; then the fun ction reverses, and as the
scent becom.es stron ger, it is judged to be m ore disagreeable (Figure 14.16a).
By contrast, a fi sh y od or, like trimethy lan1ine, may be acceptable at low con-
centr1Jtions, but as intensity increases, its pe rception becomes s teadil y rnOl'e
n egative (Figure 14.1 Gb )_ Note a lso that in di vi dual differences in the nwn be r
and type of receptors exp ressed may influen ce on e's sensitiv ity (i ntens ity

OLFACTION 455
FIGURE 14.16 Pleasantness ratings o f odorants plott9d against (a) (a)

The relationship between odor intet'"1$1ty and pleasantness is often described by an Phenylethyl ak:ohol
lnwrted-U func tion, if the odorant Is considtfed pleas.ant, as the synthetc
rose S09nt (ph9n)'iethyl alcohoQ usually is. (b) For an od orant that is initial ty c cnsidered
d
t ol€fable (but not necessatily pleasant), such as fishy smelling trimethylamine, the
relationship may more aptly bQ d9SCtib9d by a linear graph like this.
perception) and hence the predisposition to experien ce specific odors along

a pleasantness continuum.
Nature or Nurture? ;1; 11!!] Unplm•nt

A long-st.mding debate in olfaction centers around whether hedonicresponses
to odors are innate or learned . Researdlers on the inn.ate side of the d ebate l!!J Low --- - - - - -•H'ogh
daim that we are bo.m \vi th a predisp osition to like or dislike various s m e lls. Intensity
In o ther wo rds, rose is inherently a good s me ll and skunk is inherently a bad (b)
smell, the way bitter is inherently Lmpleasant to us and sweet inherently
Trimethylamine
pleasa nt (see Chapte·r 15). ln contras t1 researchers taking the Ier rnt¥i vJew
ho ld that we are born merely \vith a predisposi tion to learn to like or diSlike
sm ells, that w hether a sm ell is liked or not is d etennined by the emobon..1 1
va lue (good or bad) of the experiences that have been associated with it. Tha t
is, if we like rose and dislike sk unk, the reason is that we have a good and a
bad associa tion, respectively, with these hvo scents. \Ve need n ot have di rect c:
!
contact with a skunk to form s uch an associa ti on1 thou gh, because cultural j
lea mi ng provides mea ning to many unen countered s timuli.
If asked to take a position you rself, on the sole basis of your ov..'11 per.sonal
experiences, it's pretty like ly you '"''ou ld come d own on the ilm.a te side o f the
deba te. After a ll, wh o could like the s mell of sktmk1 and who wouldn' t like
the s me ll of a rose? [n fact1 however, a g reat deal of evi d en ce s uggests that l!!J
odor hedonics are cllmost exclusively lea rned. A good place to sta rt lookin g for
Intensity
such e vidence is wi th infa.nts. If odor preferences a re innate1 then newborns
should display them. However, researchers have repeatedly found that infants
and children o ften display very different preferenc-e:s from those of adults.
Fo r in.st.mce, infants d o n ot find the s me lls of sweat and feces unpleasant
(Engen, 1982; M. Ste in, O ttenberg, and Roulet, 1958), and toddlers often do
not hed onically differen tiate between odorants that adults find ei ther very
unp lea.s.a nt (e.g., butyric acid, wh ich s mells like dirty socks) or pleasant {e.g.,
amyl acetate1 which smells like bana na).
01e difficulty witl1 these typ<S of studies is that the olfactory system is fully
functional by the tlurd month of gestation (6 montllS before the baby is born)
(Schaal, Marlier, and Soussignan, 1995, 1998; and Porter, 1998), and
odorant molecules do find their way into the womb.So it is difficult to know
exactly how much exposure even a newbom infant has had to a n odorant. But
exposure to od ors in utero has led to yet another line of evidence in s upport
of the lea rned view of hed onlcs: .Mennella and colleague.!ii found that mothers
who conswned d istincti ve-smelling volatiles {e.g., garlic, alcohol, and cigarette
sm oke) dming pregnancy or breast-feeding had infants who showed g reater
preferences for these s mells th.an did infants w ho had not been expose d to
these scen ts (Mennella and Beauchamp1 19911 1993j Mennella, Johnson, and
Beauchamp, 1995). \'Vhat we learn about odors prenatally and during infancy
and ea rly childhood can also go on to influence our food and flavor preferences
in adulthood (Haller et al., 1999). lhis correlation s uggests that it might be
a good idea for women to eat lots of brussels .sprouts and salmon and other gestation Fetal developmenl during
heal thful foods during pregnancy and lactation . prt>;Jnancy.

456 CHAPTER 14
data provide furth er support for the idea

associative learning, and not h.a rd,-vired l'e'Sponses, is responsible
for olfactory preferences. No scientific s tudies to da te ha ve fotmd
cross-cultura l agreement for hed onic respon ses to cornm on
everyday odors, ei ther "good " or ''bad u (Ayabe-Kanamura e t
al., 1998; Schleidt, Hold, and Attil a, 1981). Anecdotal and obser-
\·ational examples a lso illus trate the many culturally p olarized
responses that \•ve have to s peci fic odors, especially food arom as.
For example, As ians typically con sid er the smell of ch eese to be
disgusting, yet most Westen-.ers consider it anything from comfort
food to an ex tra vagant indulgence. In contrast, the Japan ese often
enjoy a meal for breakfas t called nano (a fe rmented soybean dish}
(Figure 14.17) tha t most Weste rne rs wouldn' t bring near their
m ouths (some say it s1nells like burning rubber). Both d'Leese
and nottoare hi gh in protein and s imilarly nutTitious; the reason
they a re pl'efe rred, or not, has to d o with respective differe nce
in le ttmed associations to theiI scen ts.
In case you're thinking tha t the examples of odo rs g iven so
fu r aren 't that bad , and tha t there must be consens us on
really ho rrid this also doesn_,t seem to be the case. Fecal
FIGURE 14.17 The Japan9S8 r9gu-
larty eat naUO l)'lght) 1or brookfast. but smells a re not high on most North Am erican s' ''best s mells list/' but the Masa i
Westi9fners do not equatQ the smefl of Africa like to dress their hair wi th cow d\.mg as a co:sm.etic color trea hnent.
of this food 'With eating, In contrast, And in a s tudy unde rta ken by the US milita ry to create a s tink bomb (to be
c hee:se, which most W0stemers einjoy, used in place of tear gas to disband it was impossible to find an odor
Is oonsidered disgusting by m ost (induding "US Arm y-issu e latrine scent") th a t was unanimous ly considered
J apanese.
repelle nt across e thnic groups (Dilks, Dalton , and Beau champ, 1999). Labo-
ra tory studies dire ctly aimed a t testin g the learning h ypothesis have shown
tha t a n ovel odor can be m ad e to be pe rceived as good or ba d as a fun ction of
the experiences (good or bad) th a t are associated vvith it (He rz, Beland, and
He lle rstein, 2004). Note, however, that from an exp erime ntal p erspecti ve it is
much easie r to dem on strate tha t od or p referen ces can be learned th.-m . to prove
th.a t no exist to w hich ll1.nate responses may be s hown.
ln an a ttempt to support the vie w th:at odor pleasantness h.:""ts an inna te basis,
a recent stud y tested \..V he ther an od o.rant's physical and chemical s tructure
co uld predict.-1,ow pleasanL(or not) it would be perceived to be (Khan et al .,
2007). 11le researchers fo und tha t the m o lecul ar s1Tuch1re of odora nts could
accotmt for 300'u o f the varia nce in participan ts' responses (70?u was n ot ac-
cmmted for this way)a nd the refore argued th a t a portion of od or p leasantness
perception may be innate. More researc h is clea rl y needed , but as yo u '""ill
n ow read, it would actually be d is..1 dva ntageou s for l LS if o ur o d or prefe rences
were hard wired .
An Evolutionary Argument
Specialist anirnaJ species live in very specific habitats and thus ha ve a limited
number of food sources and predators. For s uch species, i.Jma te respon ses to
particular odors are ada pti ve. California ground squirrels, for example, exhibit
an ins tinctive d efen sive resp onse the first time they are exposed to the odor of
the ir natural predato r, the Paci fic rattlesna ke, but they d on' t s how the sa1ne
resp onse to the scent of Pacific go phe r s na kes, w hich a re no t their natural
predators (Figure 14.18) (Coss et al ., 1993; Pora n and Coss, 1990). Gene ralist
sp ecies (including humans, rats, a nd cockroaches), by contrast, ca n exploit
man y different habitats. For gene ral ists, the availa ble resources and p otential
p redators can va ry widely across env ironments, so it is n ot adaptive fo r these
species to have p red e termin ed olfactory resp onses to any particula r odor. For

OLFACTION 457
(a) (b)
FIGURE 14.18 The CaJifornia ground squirrel lives in a restrk;ted

habitat (a). so it has only a few natural predators. lnclu::Hng the Pacific
rnttlQS("lakQ (b). StudiQS h ave shown that, unlik9 humans. special-
ist species suc h as the Galifomia ground squirrel show innate odor
r9Spo nSEis, in this casQ to avoid th9 S09nt of animal that w ill try to
eat them.
exampl e, a certai n o d or in o ne locale may s ignal poisono us mus hrooms, but

in an other environme nt tha t sam e o dor co uld m e an nutritiou s food .
C lear evidence that learning i.s a critica l m echanism by whid l generalists
re odor responses is shown by learned taste aversions. Ra ts and hum ans
can be made to a void a fk·wor by bein g m a de sick See Chapter
15 for an explan ation of the cri tical role of olfaction in flavo r perception. For
example, p resenting a rat \vith a sweet-tasting, banM3·91.1elling drink and
then injecting the ra t wi th lithium, w hich l".:1\tses nausea, crea tes a condition
avoidance of this fla vor in the foture. Similarly, a pioneer ing "learned tas te
avers ion '' ex-periment in humans s howed th at children wh o experien ced
chemothe rapy after ingesting a novel flavor of ice cream (dubbed "nmpletoff")
su bseq uently refosed to eat mapletoff ice crea m b ut had no problem enjoy ing
a different novel-fla vored ice cream (Berns te in, 1978).
Researchers have s hovvn th at in humans the cond.i tio ned aversion is to
the smell, no t the histe, of the su bsta nce (Ba rtoshuk and WoLfe, 1990; also see
Miranda, 2012, for re\.;ew). In rodents, although discrete taste aversion can
occur {e.g., to the tas te of sacch arine; Mornga-Amaro, 2013), it is .al.so dear that
d iscrete (without a tastant) odor aversion learning can take pl.ice, especially
w hen the od orant is consumed (e.g., scented wa ter), and that a compO\md 's
odor is often the critical central fea ture of conditioned "taste'' aversion (Cap(1ldi
et al., 2004; Chap uis et a!., 2007). The lon g-tem1 effects of learned taste aversion
are d early adaptive. If p oison is ingested, it is best to learn to immediately
avoid it, rathe r than h aving to repeat the m istake until it kills you .
ll'te special im portance of initial associations in odor lea ming was further
shovrn in a recen t flvlRI s tudy wh ere it was found tha t the firs t associa ti on
m.ade to an odo.r is etd 1ed into the brain and produces a 1,mique neural signa-
ture in the a rnygdala-hippocampal complex tha t p redicts later memory fo r
that od or association. However, after the firs t association is made to an odor,
subsequent associations do notprodu1.."'e a unique ne ural s ignature (Ye.slnuun
et al., 2009). The bottom line is that the o lfactory system of gene ralists does
not come preprograrn. m ed but1 rather, is geared to very effectively learn the
meaning of s mells based on experience, especially first experiences. learned taste aversion The avdd -
ance of a novel flavor aftEf It has teen
Caveats pa/rad with gastrko illness. The srnEl l,
not the taste. of the substanoo Is key
Although a great deal of evidence suggests that odo r hed on..ics are we for the learned aversion response In
mus t no te two caveats. First, trigernfoally irrita ting odorants may el ici t pain humans.

458 CHAPTER 14
responses, and all hlUnans have an iimate dri ve to avoid pain (altho ug h even
this dri ve can be overcom e by cult\l ra l influences, as attested by the p opular-
ity of chili peppers in many ethnic cuisines). Second, as h as been mentioned
severa l times, the potential variability in the receptor genes and pseudogenes
tha.t are expressed l'lcross individuals m ay influence odor intensity; and con-
sel1u ently the perceived pleasanh1ess, of od ors. For example, peo ple w ho like
the s m el l of s kunk may exhibit th is resp onse in part because they are mis.sing
receptors for d etecting some of the more pungent volatiles, whereas people
w ho are particularly repulsed by this scen t may be e nd owed with a gre<Jter
number of receptors that are keenly ath1ned to the m ercaptan and s ulfide as-
pects pf this Tilat Cf., de tec tion of the full complement of chemicals
that make d r (e. 1,skunk, ci lantro)and how s trong or weak an odor
is perceived to be p lay a role in its perc:eived (lUl)pleasa ntness. Rec<'\ll tha t for
many odors, an inverted-U function describes the relationship between liking
an d intensity (see Figu re 14.16). Genetic differences in OR exp ression appea r
to exten d to ethnici ties as well (Menash e e t al., 2003), which may help explain
why it hos bft>n impossible to develop > universally effective stink bomb.
Associative Learning and Emotion: Neuroana-

tomical and Evolutionary Considerations
Ill the mid-1 960s in. Brita in, adult responden ts were as ked to provide hed onic
rntings fo r a ba tte ry of commo n odors (lvfoncrieff, 1966). A s imilar study was
conducted in the United Stares in the late 1970s (Cain a nd Johnson, 1978). In-
cluded in both stu d ies was the o d or an t m ethyl sa licylate (w intergreen ). In the
British sh1d y, w intergreen v11.;1S given o ne of the lowest pleas..1nh,ess ra tings;
in the American s h1dy, it was ra ted as the m ost. pleasant scent tested . There
is a historical reason for this diHerence. In Britain, the s melJ o f w intergreen is
associated w ith m ed icine; in particular, wintergreen was added to analgesics
used during \Vorld \Var 111 a time that the adults in the 1966 s h1dy would not
ha ve rem.e mbered fondly. Con versely, in the United States the sme ll o f winter-
green is exdusively a cR ndy sm e ll, so it has only s v.·-eet, p ositive connotations.
A5 this exa1nple d emonstrates, the key to o lfactory associa fo;e learning is the
experience when the od or is first encotmtered and, i.n particular, the emotional
conno tation o f thr1t experience (Bartoshu.k, 1991; Engen, 1991; H erz, 2001).
\.Vhen an odor is liked or disliked. because of w ha t it has been associa ted
wi th in o ur past, we a re al so recalling a m em ory when w e smell it. of
our odor experiences, such as most Britons' and Americans' experiences lvith
wintergreen, are too vague to conjure up specific m em ories, and only general
associations (e.g., "a happy, appetizing sme ll") and feelings of liking or disliking
are registered. H owever, one of the m os t d istinctive fea hues of o lfoction is its
abiJity to el icit o tu most em otional and evocative personal recollection s. For
orbltofrontal cortex (OFC) The more information see "Sens ation & Perception in Everyday Llfe: Od or-Evoked
part of the frootal lobe of the cortex Mem ory and the Truth behind Aroma therap y" o n p age 462.
that liea behind the bone (orbln con- Neuroa na tom y s upports th e proposition tha t o ur olfactory system is es-
taining the eyes. The OFC Is roopon-
pecially prepa red to lea rn the affective significance of odors. The a mygdala,
sible fcr the conoclous experience of
dfaction, as 1Mlll as the Integration of \v•hich syna pses directl y with the olfactory ne rve, is critical for e motional
pleasure and displeasure food; associ.a tfre learning (M. Davis and \ Vhalen , 2001 ). The orbttofrontal cortex
and it has been referred to as the sec- (OFC), wh ich is located behind our eyes at the very front of the front-a l lobes
ondary olfactory c onex and the sec- (see Fig ure 14.7b), is where we consciously e.x pe rience and perceive o d ors,
ondary taste ccrtex. The OFC Is also and it is consid ered to be the ''secondary olfactory cortex,'' just as the OFC
lnvotved In many oth€f functions, and
n Is critical for assigning affective value h as been labe led the "secondary tas te cortex" (Rolls, 2006) because it is where
to stimuli-in other'NCfds. determining pleasure <1.nd displeas ure from food are integrated. Indeed, the OFC h as been
l1edoolc meanlrg , ide ntified as the ne ural loc us for assigni ng affective va lue in genera l---that

OLFACTION 459
is1 our hedonic response to a wide range of s timuli {R. J. David son, Pub1an1 1 main oHactory bulb (MOB) The
and Larson, 2000). Notably, the right OFC plays a s ignificant role in conscious clfactory bulb: the blueberry-sized
o lfactory pe rception (W. Li et al., 2010). This is important b ecll. u se the right extension of the brain just above the
nose: the first region of the brain where
hemisphere of the brain is d on"Linant for emotional processing (R. J. David.son, smells are processed. ln humans we
1984) and odors are inherently hed o1Uc-that is, em otional--stimuli (He rz, refer simply to •owactory bulb(s)": in
2007). Furthem10re, the most ancient part of the bra.in, the rhinencephalon- animals accessory clfactory bulbs,
literall y, the "n ose brain"- w hich now comprises the primary olfactory cortex we distinguish between •main" and
and the specific s tructures of the limbic syste1n, developed firs t from tissue "accessory."
that was dedicated only to processing od ors. It \·v asn' t until later in evolution accessory olfactory b<Jlb (AOB) A
that limbic s tructures s uch as the amygdala and hippoca mpus emerged . It is smaller neural stn.cture located behirrl
the m_.n oWactory bulb that receives
interesting to consider that o ur hed onic and em otional reactions to s tirnuH in Input frcrn the vomerorasal agan.
general may have their origin in our sen se of s m ell.
Alm ost a ll species of animals use s m ell or chemica.l communication fo r vomeronasal organ {VN O) A
chemlcal senslrg crgan at the base of
the most basic beha viors necessa ry fo r s urviva l: recognizin g kin, finding the nasal cavity with a curved tubular
reproductively mates, loc.i tin g food, an d detem1ining w he the r an shape. The VNO evolved to detoot
anima l or object is dangerous (see the ne.x t section ). Only in humans h1n·e chemicals that cannot be processed
visual and auditory information mostly replaced .smell for imparting this kind by ORs. suoh as large and/or aqueous
of cm cinl knowledge about the world Yet our o lfactory system has reh.1i ned molecules, the types of molecules that
constltLne pheromones. Aoo c_.led
some of its basic functions. The most in-1mediate respo nses we have to o d ors Jacobson's organ.
are sim ple binary opposi tes: like or dislike, a pproach or avoid. Ernotions
con\"ey .similar messages: approach what is good, s..1fe, and joyful; and avoid
w hat is bad, d an gerous, or liable to cause grief. Thus, emo tions and olfaction
are functionally an<llogous. Both enable the organism to react appropriate ly to
its en vironment, 1naximizing its chances for basic survival and reproductive
su o:es.s. Viewed in this context, the hnm..1 11 e mo ti ona l system can be seen as a
highly evolved , abstractcognitiveversion of the basic beh..1.vioral motiva tions
by the olfactory system in animals (Herz, 200), 2004).
The Vomeronasal Organ, Human Pheromones,

and Chemosignals
In anirn.als that rely on s mell for s m viv"U, the olfac-
tory system consists of tw'o s ubdivisions: the main
olfactory b<Jlb (MOB) and the accessory olfactory
bulb (AOB) (Figure14.19). 11-.eAOB isattached totl1e
Co11ex
back of the MOB. Neurons from these two systems
do not interconnect, and the h'Y'O systems ftmction Accesso!i'
olfac tory.
separately in the integration of specific dlemicals. In
bulbs
order for the AOB to be activated, a structtue differ-
ent from the nose needs to be engaged. lhis stn1cture
is called the vomeronasal organ (VNO), sometimes
also referred to as 11Jacobson 1s organ'1 after the Dan-
ish anatomist who discovered it.
1he VNO is fo und in some amphibians, most
reptiles, and many mammals, induding New Worl d Oliactory
primates, but n ot in birds. 'When snakes open their muoos.1
mouth.sand appear to be licking the air, they are
actually m oving che micals from the air into the
vorneronasal organ. In mammals, the VNO is cigar-
Vomeronasal
shaped and located a t the base of the nasal cavity.
01gan
The VNOcan respond to some olfactory stim uli, but
it responds primarily to chemicals thc'l t are h igher in FIGURE 14.19 The olfactory system of a hamster, showing the
location of the vomeronasal organ and accessory olfactory bulbs.
m.olecular v.:eight tha n can be detected by the olfac- No1:9 that no functional vomeronasal organ or aCCQSsory olfac -
tory sensory neurons, as well as to chemicals that tory bulbs have been found in humans. t;A.ftt'lr an illustration by Dr.
are n on volatile. It also detects d1emica.ls dissolved Michael Meredith.)

460 CHAPTER 14
pheromone A c hemical anttted by in water (as opposed to only in air). Whether hmnans p ossess a fun ctioning
one member of a &iecloo that tnggern VNO has been the focus of mud1 deb.a te, but it is now generally accepted
a physldogical or behavioral response_ tha t a lth ough huma n embryos may have a VNO? this tissue is not neurally
In anotl1er manber of the same spe-
cies. Pheromones are signals for connected a nd disappears s h ortly after birth. Moreover, we d o n ot have a n
cheinlcal o:::mmunlcatlon and do not AOB for VNO ne urons to connect to in the brain.
nood to have any smell. ln aninwls tha t possess a VNO, its primary function is to d etect pheromones.
lordosis The position thal females Pheromones are n ot o do rs . They are d1emicc1ls tha t may or may n ot have a
o f some species (e.g.. pigs and rats) s mell. Th e word pheromone is derived from the Greek pherein, meaning ''to
need to assume In order to be Impreg- ca rry,'' and hormon, meaning "to sthnulate.'' It was first coin ed in 1959 by Peter
nated. It Involves tlie Inward curving of Karlson (a Gem1an biochemis t) a nd l\llartin li.ischer (a Swiss entom ologist) to
the spinal column and exposure o f the
genitals.
d escri be a che mical su bsta nce th at carries a rnessage about the physiological
or beh avioral s tate of one insect to another, and that in tum leads to a specific
releaser pheromone A pheromone reaction in the receiver insect. Tha t is, pheromones a re a means of ch emical
that tnggern an Immediate behavioral
respcnse among ocnspedflc.s. cornmlm ication . Today; the more genera lized definition ofa pheromone is "a
chemical compound produced by one a n.im al tha t elicits a specifi c behaviora l
primer pheromone A pheromone
that tnggEfs a physiological (often hor- or physiological resp onse in another anima l of the same species."
monal) charge among oon&iecincs. Pheromones are most important for comm unication in the social insects,
This effect usualt/ lnvot1es prolonged like ants, tem1ites, a nd bees, but they also convey important information fo r
pheromone exposure. ma ny non in sect s pecies, including primates. The re are ma n y examples of
this fom1 of chemical commtmication. Pheromones are used to identify terri-
tory. For example, a tiger w ill nib a tree with g lands from its cheeks to dai m
it-just as a cat nibs your leg with its eyebrow and rump glands to mark you
as its territoryl Pheromones also initiate a lam1 or defense reactions. \Vhen a
h oneybee stings us, the chemica l re leased from the stinger ('"'·h1ch h appens
to srn el l li ke banan a) is a cue to oth el' bees to join in-unforhmately for us.
Most important, for many a nimals pherom ones a re critically involved in
cornmunica tion (1bo ut reproductive be havio r. They can provide signals to
males about w hen a fem<Jle is fertile and pro\'ide s ign als to fe m..'lles to ini tiate
sex ual behavior. Fo.r example, a male rhesus m onkey w ill ignore a female
rhesu s in heat if h i.s n ose is blocked, and a female pig will not go into lordosis
(the p osition n ecessa.ry for impregnation) if s he isn't exposed to the male pig
pheromone and rostenone.
Phe romon es have tv.lO kin ds of effects. The hvo types of pheromones
tha t cause theSt:t ¢ff"e<i .ue. knqwn, as releaser pheromones a nd primer
pheromones. Primer pheromone effects are slow and produce a physiological
change in the recipient over time. For example, fema le rodents that are housed
together \¥ill come into estru s a t the same time after several cycles. Releaser
pheromone effects are fast and always produce behavioral responses. The
conseque nce of the banana aroma from a bee sting is one exa mple. Sexual
cues and beh avioral responses to those cues, s uch as lordosis, are other ex-
amples o f releaser pheromone effects. ln contrast, primer pheromones may
produce no behavioral consequences and m ay be noticea ble only in terms
of physiological changes.
1l1e most frequently mentioned example of a primer pheromone effect in
humans is known as the "Mc:Clintock effect" after the psychologist Martha
McClintock, w ho identified this phenomenon in college w hile purs uing her
tmdergraduate degree. l11e McClintock effect is shoi.vn when \•Vome n w ho a re
in physical proximity (e.g., live together ) over time s tart to have rnenstrua l
1..)'cles that coincide. TI1a t is, worn en w ho move into college donnitories toge ther
at the beginning of the school term often find tha t by the end of the semes te r
they're havin g their peri"ods in sync with one The McCl.intock effect is
believed to be ca used bychenUca.ls present in human swea t tha t are.capab le of
priming the ho nnonal syste ms of other individuals. However, other researd1
suggests tha t human pherornones a re a m y th (Doty, 2010), and even the

OLFACTION 461
McClintock effect has come lmder sha1p criticism. For one, McClintock's fom1er chemoslgnal Arry of various
collaborntor, Jeffrey Schank,. recentl y followed 186 01inese students living in chemicals emRted by humans that
d ormitories fol' an entire- year and saw n b evidence of m ens trual synchrony are detaoted by the oW
actory system
and that may have sane effect on the
(Yan g and Schank, 2006). Other researchers have argued convincingly that the mcod, behavior, h01mmaJ status. and/
McC lintoc::k effect is n o more than a s tatis tical artifact (H . C. \Vilson, 1992). a sexual arousal of other humans.
we d o n ot ha ve a functi oning VNO or AOB. Therefore, it is unkno"¥¥n
how th is effect could be processed, if indeed it truly is. Some have suggested
that, pre.smning the e ffect is J"ea l, hormonal infom1ation ma:y be trans mitted
from sweat th.roug h a nother system altogether, s uch as absorption through
the skin fro m physicaJ contact (Herz, 2001). Nonetheless, until the chemirn ls
responsible for menstrual synchrony, or any other puta tive pheromone, are
identified and their bioche:rnical pathways d etermined , the v alidity of human
pheromones remains dubious.
Since the exis tence of hrnna n pherom ones is highly uncerta in, a more
impartial way to discuss human chemical communication is to use the term
chemosignal. Here, chem osignal.s refers to d1emicals emittet.1 by hrnm\ns that
are d etected by the o lfactory system and that may have some e.ffect o n the
m ood or behav ior o f other hum..'l.ns.
ll1e chemical and_rostadienone is a steroid derivative of the male sex honnone
testosterone (it is chemically related to which was mentioned
ear lier), and it is present in body fluids (e.g., sweat) a t higherconcentration.s in
males than in females. fn several studies, androstadienone has been observed to
improve wornen 's rnood, but only when wornen are in the presence of men (facob,
Hayreh, and McClintock, 2001; Ltmdstrom and Olsson, 2005). fn tl1e p resence of
a female experimenter, androstadienone had no effec t on fern.ale participants'
tnood. An other study \o\rith a male experimenter and female participants fotmd
that androstadien one increased self-rated sex ual arousal and cortisol (a stress
and mousal hom1one) levels (\'Vyert et al., 2007). Note, h owever, that the levels
of and rostadienone women were exposed to in these st udies were a million
times higher than the am ount a no rmal man actually emits. Tims, the ecological
va lidity of androstadienone as an aphrod isiac chemosignal is questiona ble.
With respect to ch emicals em itted by females, a study (G. M_il1er, Tybur, and
Jordan, 2007) found that professional exotic: li e,.danc ts earned almost twice
as much in tips (averagin g $335 a nig11t versus $185 when th e performed
during the ov ulatory phase o f their mens trual cycles, compared wi th the
m ens trual phase of thei r cycles. Moreo ver, dancers w ho were taking birth
control pills (and were thus hormonally infertile) s h owed no change in tip
earnings over time and also ea rned less overall th an di d dancers who were
n ot us ing h ormonal contraception (ave.rage earnings $193 versus $276). Since
the dancers all claimed they perforrned the san\e way every d ay and that
thei r behavior to the patrons was consistent, the expl an ation offered is that
the women were perceived as rnore attractive by the male patrons when they
were most fe rtile, throu gh some mechanism othe r than behaviora l or visu al
cues. This oth er mechanism is proposed to be chemica l. Note, though, that
the degree to which conscious sm elling by the male patrons was involved W"5
not tes ted . However, a recent s tudy found that w hen men sme lled T-shirts
that had been worn by ovulating women, their testosterone levels vvere h igher
than they were after they sniffed T-shirts worn by n onovulating women or a
clean, unworn T-shirt (S. L. Miller a nd Mane r, 2010). It has also been s hown
that the odors emitted by a breast-feeding woman could increase sexu ..'ll desire
in another wom an--by 24% if the wornan s me Uing had a partner (whe the r
ma le or fe male) and by 17% if s he was single (Spen cer et al ., 2004).
H owever, n ot all female chem osignals increase sexual de.sire. Indeed , a
recent sh1dy found that chem icals p rese't1t in the tears of wornen dampen the

336 Am111a Appa
462 CHAPTER 14
sexual d esire of men (Gels tein et al., 2011). When men were exposed to a s trip
p laced und er their n ose containin g emotional tears from women, though they
knew nothing about th e source of the compound, a significan t d ecrease was
observed in self- rated desire, rated sexual a ttractiveness o f fe male fa ces,
tes tosterone levet and brain ac tivity associated V1oith sexual arousal. Are the
tea rs o f a distressed wonMn a d irect tun1off, or is the effect due to something
else? 11-te authors s uggested tha t th e d rop in tes toste rone and sexual a rou s.1!
may be a by-p rod uct of a d rop in aggression, \vh ic h is a lso manifeste d by
lowered testosterone, and th e re a re good evolutiona ry reasons to h ope fo r a
d ecrease in testosterone from potential aggressors when they are exposed. to
ymu tears. \'\'i. th tha t in min d , the au thors speculated that the effect w ould
also be seen if the tears came fro m a ma n.
ln s um , chem icals p resen t in h u m an bod y fl uids m ay mod ula te sexua l
aro usa l to som e extent. T h ese findin gs can be interpreted as il lustrating a
bio logical basis for olfactory influences on behavior, bu t they are also consis-
ten t w ith th e view that through pllst associations, od ors acquire meaning and
th en affect em oti on al and behavioral responses accordingly. Therefore, the
scent of ovu la ting wom en may be arousing to rnen beca use of the men's past
sexual experien ces; and d espi te n ot h>J \oing a d istinctive o dor, past as..socia-
tions w ith tears an d the ir connection \vi th sadness m ay exp lain why tea rs are
the anti-Viagra The d egree to w hkh perceptible odors and/ or imperceptib le
chemicc'l l s igmUs influen ce h um an sexuality is an exci ting field of curren t and
funue research .
Odor-Evoked M emory and the Truth behind Aromatherapy

You have probably had tile experience v.tiera an odor triggered a spe-
cific and special autot:i ographlcal memoiy. These occurrences are often
referred to as after the llterary anecdote described
by Marcel Proust 1Nhere the aroma of linden tea ard a madeleine cockie
suddenly triggered the recollectim of a lcng-1orgot1en event (Prous1.
1928). Prrustian memo lies have been mown to differ from episodic
mem ories triggered by ot her rues in several important vvays.
C011pared w ith \lisual and verbal cues, odors el icit more affec-
tive, old . rare. and evocat ive personal memoires (Chu and Downes.
2000: Herz and Schooler. 2002: Hinto n and Henley. 1993: Larsson and
Wil lander. 2000; Rub in. Groth . and Goldsmith, 1984 ; W illander and
Larsson, 2007; Zucco et al., 2012). o:tors also evoke more emotional
memories than musical or tac rn e cues (Herz, 1998). (See Web ActMty
14.3: Sensory Memory Cues.)
For examp le. Herz (1998, 2004) c ompared recollections stimu-
lated by a familiar sm ell-for examp le, p::opcom-w lth memo ries evoked
by the sight of popcorn , the sound of popcorn popping. the feel of pop-
corn kernels, o r simply the word popcorn (Flgure 14.20). Consistently,
memories that W€fe triggeroo by o dcrs were experienced as more emo -
tionally intense. and participants felt m ore transported back to the oligi-
nal time and p laoe of the event , than when mem ories were tri ggered by
rues in any other moctality.
The distinctive emotional features of odor- evoked mernay are ex-
plai ned by the uniquely direct connectlon between the neural substrates
of olfactlon. emoti on . associative learning, and m emory (Gahill et. al. ,

OLFACTION 463
1995). Only two synapses separate the olfac tory nerve from the amygdala,
a structure critical for tre expression and experience of emotion and hu- FIGU R E 14.20 T he smell i;.:"l),
man emoti onal memory; and only three synapses separate tre olfactory sight (/;>), sound (c), feel (d/. and
nave from the hlppocampus. involved in the selection and transmission of vert>al label (e) of popcorn elicit
information in v.t0rking memory, short-term and long-term mem ory transfe-, IT'19fT'IOrio9S that are equivalent in
terms of ther accuracy. Ho'Yverver,
and various declarative memory functicns (Elchenbaum , 200 1). The amyg-
til
odor-induC9d racollections ani
dala has also been shown to play a major role in stimulus reinforcement as- more €·rno tional, and this
soclatim learning in primates (Jones and Mishkin, 1972: Wi lson ard Rolls, q uality has earned odors a reputa-
2005), and neuroimaging studies In humans have shO'A'n a direct neuro- tion as particularly good cues for
biolog lcal correlatlon OOtv.teen recall of significantly emotional odor- evoked memory.
memori es and activity in the amygdala (Arshamian et al., 2013; Herz et al ..
2004; Varnetten et al .. 2007). (a) Sm• ll
Notatfy, although memories evoked by odors
have many unique and special characteristics, they
are not more accurate than mem ories elicited by other •
cues. Nevertheless. odors have earned the reputation • · ·
of being the "best cues" to memory. Part of thi s can be = •: · · · ' .
explai ned by the same mechanisms that make eyewlt- ' - '''
ness testlmrny so fraught. When there is a hi gh degree .
of emotion experienced during recollection. pecple are
much more confident that th0 r memories are oorrect,
r1i·
even thOL.gh they are oft en sho·wn to be inacx:::urate
(Herz. 1998). However, It Is also the case that as with
Proust. odors can remioo one of memories that might
otherwise be forever forgotten. That Is, odors may
unlock memor1es whose
only "mental tag" was the
odor that was present when
the memory v,ias enooded.
The reason for the special (c) Sou nd
capacity of odors to do this
may be twofold : (1) the low
frequency by w hich certain
odors are encountered, ren·
dering minimal intetiererce
from mUltiple associations.
and (2) resistance to being All
overvoJTitten-there are very equiv.ilent
strong proactive intetierence recoUecUon
(d) F"I
rui·
effects in olfactlon. so the """""'!'
first association you aa:iulre
to an odor is extremely hard
to undo, and subsequent as-
soclatims are very hard to make. Unfortunately, It is not
possible to test whether odcrs can ret rieve memories
t11at would othervvise never be remembered. but anec-
dote suggests that this may indeed be the one feature
that makes odors "better" memory cues (Herz . 2007). (e) \•\!rilten l.1b.;J
Odors that elicit specific emotio nal associations
can also produce ooncomltant changes in beha\'ior.
For exampe, it has been shown that an odor that was
associated ,,,;th a frustrating experience led to reduced
motivation and perfonnance when later smelled (Epple
and Herz, t 999; Herz. Shankler. and Beland, 2004). and
odors that have acquired connotations of being ener-

464 CHAPTER 14
aromatherapy The manipulation gizing, such as r:eppennint. cinnamo n, and grapefruit, can lead to
of cdas to Influence, mco:j, pfffor- heightened physical and mental r:ertonmance when participants are lat-
manoe, and well-being, as well as the er exp:>sed to them (Raudenbush, Coney, and Eppich , 2001: RaL.den-
physiological correlates of emotion bush et al.. 2000). Proponents of aromatherapy contend that odors
s uch as heart rate, blood pressure, alter mood, performance, well-being, and the physiological correlates
and sleep of emoti on (e.g .. heart rate, bloo:l pressure. and sleep) in a drug like
autcmatic manner. There Is , however, ro eviderce for phannacological
effects of odors in humans. Instead, so-cal led aromatherapeutic effects
can all be explained by the em otional memo!)' assooiation that was
acquired to the scent (Herz, 2009). The scent of can indeed
make you feel invigo rated, but o nty if you have associated uplifting and
energizing associations to this aroma It is also the case that the emo -
tions elicited by an odor can have downstream effects on physiology
and p erio!lllance. Therefore , an OOor associated to an arousing experi-
mce (e.g., can increase your heart rate and may also make
you run faster. However. If you hwe never smelled peppermint before,
or if you dislike the scent. it would p-cduce either no effect or a negattve
one (Herz, 2009).
8230336 AMma Appa

Summary
1. Olfaction is one of the hvo chemica l senses; the other is taste (discussed in
Refer to the Chapter 15). To be perceived as a scent, a chemical must possess certain
Sensation and Perception physical properties; however1 even some molecules that have these ch arac-
COmpanlon Website teristics cannot be smelled. Olfaction has some unique physiological proper-
ties, one of which is tha t only be tween 35o/o-4C1'/o of the genes that code for
Sltes.Slnauer.convwone4e olfactory receptors in humans are funct ional Another unusual ie.."l.tu re is
for aottvities, essays, study tha t most odorants also s timulate the somatosensory sys tem via the trigemi-
questions. and otrer study aids. nal nerve, and it is often impossible to distinguish the contribution of olfac-
tory senMtion fro m trigeminal stimulation.
2. Anos:mia is the comple te absence of a sense of smell. It is most frequently
caused by sinus disease, which can usually be treated and enable the return
of normal olfactory function. However if anosmia is caused by head trau-
ma, it is likely to 'be penn anent. Anosmia can lead to severe disturbances
in an individual 's quality of life. Gradual loss of olfaction is a normal con-
sequence of aging; however, sudden olfactory loss can be the fi rst sign of
Alzheimer's or Parkinson's disease and should be investigated.
3. The dominant biochemical theory oi odor percep tion- shape-pattern theo-
ry-contends that the fit behveen a molecule and an olfactory receptor (OR)
determine \,:hich molecules are detected as scents, and that specific odor-
an ts activa te a combinatorial code of ORs, producing specific patten\S of
spa tial and temporal n eural ac tivation for each perceived. scent. However1
this theory is not universally accepted, and alternate ex'Planations exis t (e.g.,
vibration theory).
4. Recently, researchers demonstrated a d o.ser oonnection behveen the visual
system and olfaction than has ever before been thought to exi.st. Two exam-
ples are bi naral rivalry and the discovery of "olfactory white.'' TI1ere is also
a difference behV'een active sniifin,g and passive inhalation of odors at both
neurological and functional levels. Active sn iffing may even have therapeu-
tic application s fo r patients suffering from extreme physical disabilities.
5. Although we can potentially detect over a trillion almost all odors
that we encounter in the real world are mixtures, and \Ve appear n ot to be
very good a t analyzing the discrete che mical components of scent mixtures.
OUaction is thus primarily a synthetic, as opposed to analytical, sense.

OLFACTION 465
Hm"·ever, analytical olfactory ability can be developed w ith training. True

odo r imagery is also \ <\l eak (or no nexistent) fo r most people, but training, as
in the case of odor experts (e.g., perfum ers),. appears to facilita te this ability.
6. The psycho physical study of smell has shm'111 tha t various odorant intens ity
levels and various cogni tive fun ctions are requ ired for odor de tec tio n, d is-
crimination, and recognitio n. Identificatio n differs from odor recognition in
tha t, in the form er, one m ust come up \\.ri th a nam e for the olfactory sensa-
tion . It is very difficult to name even very fam iliar odors, and as we age this
becom es even harder. 1h.is state is known as the tip-of- the-nose pheno m-
enon-one of several indica tions that lingu istic processing is highly discon-
nected fro m olfactory experience. H owever, new resea rch h as shown that
at least o ne culture may possess an enhanced verbal connection V>' ith odors.
Regardless, rmlike the case w ith o ther sensory experiences, \\'e do no t need
to access any semantic infor ma tion about an odor in o rder to respond to it
appropriately, as long as it is famil iar.
7. Ano ther in1portant d iscrepancy beh\'een the physical experience and the
psycho logical experience of odors is the d ifference between receptor adap-
tation and cognitive habit uation. Receptor adaptahon occurs a fter continual
odorant exposure over a nu mber of mi nutes, can be undo ne after a fe\-.'
m inutes away from the odorant, and is explain ed by a basic biochemi-
cal m echanism . By contra.s t, cogn itive habituatio n occurs after long-term
exposure (e.g ., in a living or \-Vork envi ron ment) to an odor and takes weeks
away from th e odor to undo. Psych ological influences can have stron g
effects o n both perceived odor adaptation and ha bituation also .
8. The m os t immedia te and basic response \Ve have to an odo r is wh ether we
li ke it o r no t; this is hedonic evaluatio n . Odor hedonics are measured by
pleasan h1ess, fam iliarity, and in tensity ratings . Pleasantness and fa miliarity
are linearly re lated to odor liking; o dor intensity has a more complex rela-
tionship w ith hedon.ic percep tion. Substa ntial evidence suggests that our
hed onic responses to odors are learned and no t inna te, even fo r so-called
stenches. llIBt we lIBve learned to like o r d islike vario us odors ra ther than
being born w ith hardwired responses is evolutionarily ad aptive for general-
is t species like h umans . The caveats to the leam ed-response proposition are
odors tha t are highly trigeminally irrita ting (pain inducing) and the genetic
variabili ty in the numl:::er and types of receptors expressed across individu -
als, wh ich m ay influence o lfactory sensitivity and hence o dor hedon ic
percep tion .
9. The key to olfactory associa tive learning is the e mo tional value of the con -
text in wh ich the odor is fi rs t encountered. If the emo tional contex t is good,
the odor w ill be liked; if it is bad, the odor w ill be disliked . Previously
acquired emo tional associa tio ns w ith odors also underli e valida ted aroma- 136 AMma Appa
therapy effects. Emotion al potency further dis ting uishes o dor-evoked mem-
ories from m emories triggered by o ther sensory cues. The ne uroana to my of
the olfactory and limbic system s and their neuroevolutionary d evelopment
illustra te how emo ti onal p rocessing and olfactory processing are uniq uely
and intimately interrelated .
10. Pheromones are chemicals emitted by individuals tha t a ffect the physiology
and /or behavior of other me mbers of the sa mespe::ies and may or may no t
have any smell In al l mamma ls tha t ha ve been shown to use pheromones
for rommunic3tion, d etection is medfa ted through the vomeronasal o rgan
(VNO) and p roces.sed by the accessory o lfactory bulb (AOB}. Humans do
no t possess a fun ctional VNO o r AOB, and evidence fo r human phero-
mon es is controversial. H owever, human chemosig nals that axe processed
through the olfactory system appear to have som e influence o n ho rmonal
status and sex ual arousal.

CHAPTER 15

Taste
CALVIN TRILLJN, A WRITER WHO MAKES WONDERFUL OBSERVATIONS about the

joys of eating, described his 4-year-old daughter' s reaction to 'jpolishing off
a particularly satisfyin g d ish of d1ocolate ice crea m ." She sa id, "'My tongue
is s miling."
What makes to ng ues smile? As with o ur noses1 the basic answer to this
question is m o lecules. Olfaction and gustation are often grouped together as
the chemical senses, and in ter1n.s of these two sensory systems
are in som e ways quite similar. But the che rnicals we taste have alrea.dy e n-
tered our mouths and are about to move even farther into o ur bod ies. Thus,
taste serves the m0t3t specific hmcti on of a ny of the senses: discerning w hich
chemicals we need to in gest because they are nutritious, and w hich we need
to s pit o ut because they tnay be poisono us. Pe rha ps this is w hy some thing
about our liking or disliking of tastes a nd fla vors seems to be very dillerent
from the liking or d isliking that one might associa te with the color red or the
sound of middle Co n the piano. Nature h as equipped us to care passionately 0336 AmMa Appa
about food because passion h olds the key to otu s un+val.
Taste versus Flavor

Before delvin g any farther into the gus tatory systein, we n eed to dear llp a
very old m isundersta nding. According to the ear ly Greeks, sensa tions per-
ceived from foods a nd beverages in the rnouth were tastes, and sensations
perceived by s niffing were smells. ln fact, however, food m olecules are almost
al ways perceived by both our taste and mu o lfactory systems. The molecules
we taste are dissolved in our saliva and stimulate the taste receptors on o ur
taste b uds, as we' ll discuss in thi s chapter. But when we d1ew and swa llow taste Sensations evci<ed by solu-
foods, othe r m olecules are released into the air inside o ur mouths and forced tions In the rnout11that oontact recep -
tors on the tongue and the roof of the
up behind the palate into th e nasal cavity; w here they con l:D.ct the o lfactory
mouth that then connect to axcns In
epithelium a nd stimubteour o lfacto1y receptors{ Figure 15.1 ). The brain then cranial nE<Vlls vu, IX, am X.
knits these retronasal o lfactory sensations together with our taste sensations
into a kind of metasen.sation that goes by the name flavor. retronasal olfactory sensation
The sensation of an odor that Is
It is quite easy to prevent the airfl ow that carries od ornnts through the perceived when chewing and swal-
retron a.sa l passage. C hildren do it all the thn e when they h old their noses lowing fcrce an odorant In the mouth
w hil e eating spinach. Try the following experi ment n ow-but use a piece up behind tl1e palate into the na;e.
of ch ocolate if you're no t crazy abo ut spin ad1 . Pinch your n ose before put- Such odor sensations are perceived
as originating from the mouth, even
ting the chocolate in your m outh, Md then d1ew it and no te the sensation, though the actual contact of cdcrant
which will be almost pu re taste (sweet \vith a bit of bitter). Then, jus t before and receptor occurs at the owactory
swall owi n g,. release your nose. The volati le m olecules responsible for the mucosa.
chocolate sensation vi.rill immediately flow up behind your palate and into the flavor ll1e ccmblnatlon of true taste
nasal and you wil l understa nd the difference between taste and flavor. (sweet, salty, sour, bitter) an::I retrona·
You've probably noticed before that flavor is sirn.ila rly impoverished when sal olfactlon.

468 CHAPTER 15
FIGURE 15.1 Molecules

into the rur Inside our m ouths as we
c hi9w and swallow food trav9 up
through the retronasal into the
nose, where they then m ov9 upward
and oontact the olfactory epithelium.
you have a s tuffy nose. Web Activity 15.1: Taste without Smell asks you to
test this phenomenon wi th o ther s timuli.
lf you are a very care ful o bserver, yo u will not1ce so me thing e lse tha t
happens w hen you do this retronasal o lfo.ction de mo nstration: the sweetness
of the ca ndy w ill increase w hen you perceive the choco la te sensation. This
results bec'1use retron asal olfactory input enh an ces the taste of sweet. Our
current tmderstanding of this comes frorn increasing awareness that the central
integrntion of re tronasal olfac tion .and raste is complex, a nd we have yet to
underst..1 nd all of the rules.
Foods a re also perceived by the somatosensory system via touch, tem-
perature, and pain receptors in the tongue and 1nouth. Some of these sensa-
tions have protective functions: the bum of add (which might d amage your
s tomach if swa!Jowed)1 the hea t pain from scalding coffee, the pain of biting
the ton gue, and so on. Soma tosensa ti ons also provide infor mation about the
nahU'e of foods and beverages. For exam.pie, we get llifom1ation about the fat
content of foods from tactile sensa ti ons sud 1 as oily, viscous1 thick, and crea my.
Localizing Ravor Sensations

You should have realized something else w hen you perfonned the ch ocolate
experiment described ii'\ the previous section: even though you now know that
the ch ocolate sensa tio n origina tes from the olfactory receptors in your nose,
you p roba bly still perceived the flavor as coming entirely from your mouth.
1l1is perception is due in part to the tactile sensa ti ons evoked by chewing
and swallowin g, and in part to taste. Because you taste a nd feel the food only
in your m outh {n ot in your nose), your brain concludes tha t the sensa.tions
mus t have '1risen entirely from the mouth. Exceptions hldude foods s ud 1 as
chorda tympail The branch of cra- h orseradish, wasabi, a nd sp icy mus tard, which give off volatile chemicals
nial nerve VII (tl1e facial nerve) that car-
tha t activate pain receptors a ll the way up through the retronasal passage. But
nes taste lnforrnatbn trom the anterior,
mobile tongue (the part that car be these exceptions prove the mle: when we ea t the.se foods, we experience the
stucK Olrt). The chorda tympani exits sensations as coining from our n oses as well as our mouths.
the tongue with the lln;iual branch of Now consider the fo llowin g . :-urious case. A patient with n orma l olfaction
the trlgemlnal nerve (cranl31 nave V) but d a maged and ora l touch reported that she could smell lasagna, but
and then passes through tl1e mlddk>
ear on Its way to the brain.
when s he ate it, it h ad no flavor. Asimilar effect was produced in a labora tory
using n s mall a mo m1t of lidocaine and a large am m.mt of blueberry yogurt.
cranial nerves Twelve pairs of Subjects in th is sh1d y had thei r left chorda tympani (a brnnch of the fa cia l
nerves (one for each side of the bod0
that originate In the brain stern and nerve, one of the cranial nerves that carries iofotillRtiqn.fTQm rei;:eptors
read! sense organs am rnusoles to the brain) anesthetized with the lid ocain e hile they the yogurt 1n
through openings In the skull. this situation the s ubjects reported tha t the blueberry se.nsation-whicl'\ is d ue

TASTE 469
entirely to retronasal olfaction ----seerne d to com e only fr om the right s ide of the
m.outh. Moreover, the intens ity of the blueberry sens ation was reduced, and
this intens ity was reduced even further \vhen both tas te nerves were bl ocked
(D. J. Snyder et al., 2001).
In both the pa tient and the experimental subjects, the pathway front the
tnouth to the "'lSal cavfly. - int.let. Why then were their lasagna
and bl ueber ry sensations reduced? This 1s a nother consequence of the central
links betv•l een retro nasa l o lfac tion and taste. When the retronasal olfactory
sens ati on of ch ocola te was added to S\l\·eetness, the S\veetness went up; when
sweeh1ess was re m oved (with anes thetic), the ret ronasal o lfactory sensations
(lasagna, blueber ry) \\.'ent dow n.
Recent brain-imaging research by Dana Small was key to revealing charac-
teristics of retro nasal olfaction: the brain processes odors differently, depending
on whether they come frorn the m outh or through the nostrils. This d istinction
ma kes good sense fun ctionally because the significan ce of odors in the m outh
is very different fro m that of odors s niffed from the outside world . 'Alithout
the p rope r cues to te ll us w here an od orant is comin g fr om , input from the
o lfactory recepto rs a pparently cannot be routed to the p rope r b rai n a rea to
connect the smell sensation w ith the food stimulus.
Some of the interactions behvee:n taste and retronasal o lfuction h ave been
understood by the food. indus try for many years (SjOstrOm, 1955). For example,
if a company is mar ke ting p ear juice and w ants. to intens ify the sensation of
pear, it c.an add s uga r. The increase in sw ee tness (a pure tas te sensation) w ill
increase the perceived olfactory sens.a. ti.on of On ly more recently have
we beg un to focus o n the reverse interacti on: intens ifica tion of
taste (see Bartoshuk and Klee, 2013). The commercial impli cations of this are
obvious. The addition o f the appropria te volatiles could a llow a ma nufacturer
to reduce the amotmt of s ugar (or a rtificial sweetener ) and sa lt. TI1e scien tific
puzzle 1·em.ains; do these complex interactions se rve som e biological purpose?
The pervasiveness of food a dditives such as ccurageen an, guar g\un, and
other thickening agents.sh ows tha t the food industry also has a good h andle
on how somatosensati on affects food perception. And the ingredient lis ts of
mo.st p rocessed food s include at least on e artificiaJ co loring, testifyin g to the
importance of yet an other sense, vision, in how we perceive foods.
Volatile-Enhanced Taste: A New Way to Safely Alter Flavors

Flavor dominates our ttves . Flavor is created in the train from the inte-
gration of retronasal olfactlon and taste , b ut the rules governing that
integrat ion are not vvell understood . As noted earlier, since the 1950s,
the focd industry has made oonside-able use of one of the rules: add ing
sugar to fru it flavors intensifies t11e fruit navors. It took a couple more
decades before hints began to appear that the reverse \'.Jas also true:
sorne fruit flwors V\l"ere sh:>wn to intensify sweet. At first this did not
seem to be practical because the effects vi1ere so small. H01i\'ever. a.
serendipitous finding during a tomato (botanically, a fruit) experiment of-
fered a different point of view.
The tomato experiment was aimed at a problem focd shoppers
are well aware of: it's hard to find a good tom ato in a supennarket.
Recently, heirioom tomatoes have been finding their way into market s.
Why would these tomatoes taste better? The answer is t hat intensive

470 CHAPTER 15
b reed ing to giJe tom atoes charac teristic s thought to b e desirable (uni-
form ripening time, fruit Sze, and so on) have in some cases led to a
d eterioration in flavor. Heirloom to matoes are genetically m ore diverse
b e::ause they cane from a time pri or to the intensive breedir"Q. Thus
some of them have tt-e flavors rem embered 1rom those earlier days.
Collaboration between plant bldogists and psycho log ists at the
University of Florida 100 to an experiment utilizing eighty heirloom to -
matoes g rovvn on university fatmlard. After harvest ing , half went t o a
chemistry lab where the sug._qrs, acids, and vci atlles v.iere measured :
half went to a psychophysic s lab where taste . flavor (I.e .. retro nasal ol-
fac tlo n), and preference W ff'e measured. Regression analyses ldentlfie:J
the comp:::>nents responsib le fo r the sensory properties of the toma-
toes as well as how much they were liked . The solutio n to the problem
turned out to be sim ple. Some tomato oonst ituents correlat ed positiJely
with liking; that is. the more of that const ituent that was in the tomato ,
the mo re it was likOO . Some did the reverse and some did not m atter.
To make a better to mato , Increase the c onstituents oontritutlng to liking
and decrease t hose contributing to dlel iklng. Knowing what to aim for.
cross-b reed ing \vi ii give us better t omatces.
The serendipitous result cam e from a mathematic al anatysls of
the to mato dataset. Multiple regression is used widely in the social sci-
ences to examine vartcus sources for a given effect. Fo r exam ple, an
investigator mig ht \o'tant t o look at contributions to 10 from a variety of
sourt::es (age, health , lnoome , educaticn , and so o n). Multip le regression
was appl ied to the tom ato data, to see If any oonstituents other than
the sugars were oontributing to sweetness. The resu lt w as startling. A
considerable amo unt of sweetness was c oming from the volatiles. It
was sLddenly apparent that smal l effects from individ ual volatiles were
adding up such that a oonslderable am ount of the sweetness of a given
tomat o was produced by the volatiles (perc eived retronasally). For ex-
ampl e, increasing the conc entrations of those volatiles could double the
s1,veetness of a tomato .
The Imp lic ations for sugar reduct ion are dear: adding the correct
volarnes can red uce the amount of sugar needed to sweeten foods and
b everages. However. the potential goes ei1en further. There are ot her
volatiles that can enhance salty and still m ore that can supp-ess t.:Ctter.
The d iscoveiy that plants use volatiles In this way is very new. Are
these enacts in the brain ? Are they acquired somehow from
experience? The original thinking about volatiles that oould enliance
sweet was that these would be limited to fruity flavors because we so
oft en experience fruit and sweet together. However. one of the tomato
volatiles that enhanced sweet was lsovalerlc acid, which smells like
S\Neaty seeks. Sweaty socks and sweet d o not seem to be a oombina-
tio n that would b e exp!rienced to gether very often!
Muc h is yet to be learned. However. better to matoes are surely
right around the comer. alo ng with safe new ways to sweeten or salt
foods and reduce unw anted bitter (e. g .. bitter ve;ietablas , m edications)
8230336 Amna A to follow.
tast e bud A globul..- cluster of eels Anatomy and Physiology

that has the ft.netlon of aeatlng neural
sk;Jnals c onveyed to tl1e brain by the of the Gustatory System
taste naves. Some of the cells In a Ta ste percep tion co nsists of th e follow ing seq uen ce of events (the stm ctu res
taste bud have specla.lized sites on
their apical projections that Interact involv·ed are illus trate d in Figure 15.2): Che \vi.ng b reaks d o vvn food s ubsta nces
w ith taste stimuli. Some of the cells into m o lec ules, w hich are disso lved in saliva. The sa li va-b o rne food m o l-
form S')'Tlapses with taste nerve fibers. ecu les flow .into a taste pore tha t lead s to the tas te buds h oused in structures

TASTE 471
FIG URE 15.2 Th9 locatbns of ea.ch

type o1 taste papilla ar.:. Identified in the
of the t Cf"lg ue at lo wef right.
N9Ural signals from the t aste buds in
those papilla€! are transmitted via cra-
nial nerves Vll, IX, and X to the bra.in.
T.tSterec-eptor.s
V tb_. .o..,
CircumvaJl,1te
papillae
, Faliate ._..,., pillae

C!ol'oSopbaryngeaJ
nen -e(cranial
nen•eIX)
cal led papillae (singulm- papifla) that are loca ted mostly on the ed ges of the
ton gu e in a rough ov al (ii the olfactory epithelium is the retina of the nose,
the tongue is the retina of the m o uth). Taste buds, in tum, multiple
taste receptor cells,each of which resp onds toa limited number of molecule papilla Arr/ of multiple structures
types. When it comes in contact with one o f its p referred molecules, the taste that gll/e the tongue Its bumpy appear-
recep tor cell prod uces action p o tentials that send informa tion along one of ance. From smallest to largest the
the crania l nerves to the brain. See Web Activity 15.2: Gustatory Anatomy papilla types that contain taste buds
for a n interactive overv iew of the sys te m, w hich is described in gre.:1 ter de tail are follate, and clrcumval-
late; flllform papillae, which do not con-
in the sections that foll ow. tain taste buds, are the smallest and
most numerrus.
Papillae
taste receptor cell A cell v.;thln
Papillae give the tongue its bumpy appearance a come in four major va- the taste bud that contains sites on
rietie.s: filiforrn, hmgifom1, foliate, and d rcumvallate. The last three of these Its apical projection that can Interact
contain taste buds. with taste stimuli. These sttes fall Into
Flllform papillae, th e ones w ithout nny taste hmc ti on, are Joc,1 ted on the two major categories: those lnter-
actlrg with charged particles (e.g.,
anterior p ortion of the ton gue (the pa rt we s tick o ut when giv ing someone n sodk.rn and hydrogen lms), and those
raspberry) and come i.n different shapes in diffe rent species. In ca ts, they are Interacting v.;th specific chemical
shaped like tiny sp oons with sh.c'1 rp edges. The filifonn papillae on our tong ues structures.
d o no t have these sharp ed ges, which is why you will find lapping milk from tiltforrn papillae Small structures
a bowl considerably more difficult than your cat does. m the tmgue that prO'Ade most of the
Fungiform papillae, so named because they resemble tiny bu tton mus h - bumpy appearance. Fillform papillae
rooms, are also located on the ante rior part of the tong ue. They are visible to have no taste functlm.
the na ked eye, but blue food colorin g sv•rnbbed onto the tong ue makes the m fungiforrn papillae Mushroom-
particularly easy to see (blue food coloring sta ins filliorm papillae mu ch be t- shaped strwtures (maximum diameter
ter than fun.g iform papillae, so the fungifonn papillae appear as light circles 1 millimeter) tl1at are dlstnbuted moot
densety on the edges of the tongue,
again.st a darker blu e background )_ Ftmgiform papillae in di.a.meter, but espedally the tlp. Taste buds (an aver-
the maxi mum is about l millimeter (mm). On average, about six taste buds age of six per papilla) are burled In tt'e
are buried in the surfare o f each fungiform papilla. If we s tain the tongues surface.

472 CHAPTER 15
FIGURE 15.3 EM"llYlples showing

typical varlabit ity in the density of fun-
giform papil!M from one indMdual to
the next. 1h9 circles show the 6-mm
template area u sed for counting fungi-
form papllaeon tongues stained with
blue food coloring. (a) Tile tongue o f an
averagQ taster has 16 fungifon-n papil-
lae. In extrQrne cases. normal Individu-
als may have as few as 5 fungiform
papitlaG in that area or as marry as 00.
(b) A supertaster's tongue (supertasters
are discuss9d latier in the chapti9f) has
60 funglfotm papiUM in that area.
of many individuals, we see a large amount of variation (Figure 15.3). Son'!e

tongues have so few hmgifom1 papillae that their stained tongues appear to
have p olka d ots on the m. Other tong ues ha ve so many that the p olka dots
are wa ll-to-wall.
Foliate papillae are l.ocated on the sid es of the tongue r1t the point where
tl"l fu ngue 4rf<len m:>gnifica tion, they look like a series of folds.
Taste buds are buried in the folds.
Finally, clrcumvalate papllaeare relatively lu-ge,circular slmctures fom1ing
an inverted Von the rear o f the tongue. 1l1ese papillae look like tiny islands
su rrounded by moJts. The tls te buds are buried in the sides of the moats.
Although most people don' t real ize this, there are also taste buds on th e
roof of the mo uth where the hard and soft palates meet. To d em ons trate these,
wet your finger and dip it into salt crystals. Touch the roof of your mouth and
move your finger back until you feel the bone end (the margin be tw"een the
hard and soft palates). You will experien ce a flash of sa ltiness as you move
foliate papillae Fc>ds of tissue the salt crystal s onto the taste buds arrayed on tha t margin.
containing taste buds. Foliate papll lae In stun , the taste buds are distributed ina line acros.s the roof of the mouth
are looated on the rear of the tmgue and in papillae distributed in an oval on the tong ue. Fungiform papillae make
lateral to the drcumvallate papillae,
where the tongue attachoo to the up the front of the oval, a nd foliate and circum valla te papillae make up its :re.., r.
rnouth. Note that we have no subjective awareness of this distribution of tas te buds.
(For m ore on this topic, see Web Essay 15.1: Scientific Urban Legend- The
clrcumvallate papillae Circular
structures that form an inverted Von Bogus Tongue Map.)
the rear of the tongue (three to I ve m
each side. with the largest In the 001- Taste Buds and Taste Receptor Cells
ter). O rcumvallate papillae are mound- Tas te neurons are pseud ounipolar: a sing le process exi ts the cel l body and
llke structures, each surrounded by a
then splits into peripheral and central limbs. The periphe ral axons make up
trench Qike a moat). Thooe papillae are
much larger than funglform papillae. the n erves that project into the tongue (e.g., ch o rda tympani and glossophd-
ryngea l ne rves). The central axons project to the brain.
microvilli Slender projections of the
cell membrane on tre tips of some Ec1ch taste bud is a clus te r of elonga ted cells, organized much Hke the seg-
taste bud cells trat extend Into the ments of an orange (Figure 15.4a). Th e tips of som e of the cells end in s lende r
taste pae. mlcrovilll (singular microvi1lr1s) containing sites that bind to taste s ubstances.

TASTE 473
FIGURE 15.4 Tas1"' buds. (a) A taste bud buri€1d in the tissue o f a fungitorm (a)
papllla. (/J) Stimulation of 1h9 three taste receptor cell types results in responses Microvilli
in th9 taste nerves. T)ll)Q IH ceUs have Neurotransmitters {e.g., ATP
(adenosine t1iphosphateD may be secreted by cells 'Without TuesQ neu-
rotransmltte:irs may act on adjet.oent as we 1as taste nEf'le fibers, to ultimatel'j
produce nQUral signals that \llr'ill trav91 to the brain. (Aft'1t'" Ropgr and Olaudhati,
2009.)
Taste bud
sen.sorycelJ
In an ear lier era, these microvilli were mistakenly thought to be tiny hairs;
we n ow know that microvilli are extensions of the cell membran e.
Som e years ago, t.:'lste receptor cells \'\-"ere th ought to have sites sti mulated
by taste stimuli at one end and synapses with taste axons at the other end.
But rnore recent work has identified taste receptor cells that do not synapse (b)
with taste nerve fibers. H ow is information from these receptor cells con- Typi> I Typi> II Type Ill
veyed to the brain? Details of how this happens are begi.Jming to emerge
Sdty? Sour?
•
(Hemess et al., 2005; Roper, 2006, 2013). Taste bud cells a:munun.imte with Swn""f, Nttcr,
l l
um:.1mi
each other within the taste bud. An.atomically, taste bud ce lls fall into three
different groups with different functions (Figure 15.4b): Type I ceUs appea r
to be ho usekeepin g cells th.at excrete potassium through the taste pore al-
lowing the other ce ll types to maintain their resting membrane potential,
but they may also play a role in sal t taste. Type il ceUs are true receptor cells
that tend to respond to only one of s\-1,reet, bitter, or lllTh:'Uni s timuli. These
cells have G protein""'oupled receptors (GPCRs) on the microvi lli but do not
have synapses. Type III cells are nm ...· called "presynaptk" cells: they have
S}1'apses. Presynnptic cells may also play a role in sour taste. Many deta.ils
of the ce ll-to-cell comnnm.ication are yet to be determined, but the overall
m essage is that different tastants exci te different tas te bud ce lls utilizing
different transmitters. The contributions of this within-taste-bud processin g
to our taste experiences are not yet knO\Yn (Roper, 2013 ).
ln fun giform papillae, the taste ne rve fibe rs that ente r the taste buds
branch, so an individual cell can be innervate d by mo re than o ne taste
fiber, and a n individual taste fiber can irmervate more than on e cell. Taste
receptors haven limited life spa n. After a matter of days they die and are
replaced by new ce lls. This ronshmt renewal en ables th e taste syste m to 30
recover fron1 a variety of sou rces of damage, and it e.xplains why our i.:'lste
systerns remain robust even into o ld age. Recordings from taste nerve fibers
show that different receptor cells contacted by branches of a single fiber
sho'"'. simila r specificities to taste sti muli. In other words, it appears that
the ne rve fibers are somehow able to select the cells 1<vi th which they will
sy napse so that the message they convey rem.ains s table, even though the
receptor cells are continually replaced.
The mechan isms that p ermit a taste cell to recogni ze a taste stimulus
(knmvn as a tastant) that is contacting its microvillus can be divided into
hvo large ca tegories. One class of tastants is made up of s mall, charged
molecules that taste salty or sour. Small openings, called "ion channels,'' in
microvillus mernbranes allow some types of charge:d particles to enter cells
but bar o thers. When the d1arged particles in sa lty and sour foods enter sa lty
and sour receptor cells, these cells signal the ir respective tastes.
Tastants in the second cl ass, which produce sensations that we labe l as
sweet or bitte r, are perceived v ia a mechanism si m.ilar to that in the olfac-
tory sys te m, using GPCRs. These wind back and forth across microv illus
rnembranes, and when a particular tastant molecule "key" is fitted into the
''lcx::k" portion of a GPCR on the o utside of the m embrane, the portion o f tastant Any s timulus that can be
theG PCR inside the cell s ta rts a cascade of mo lecular events that evenhially tasted.

474 CHAPTER 15
causes nn action p otential to be sent to the brain. GPCRs a lso contribute to

what is called the ''umami" taste. More abo ut this later.
FURTHER DISCUSSION of Gl'CRs in the context of olfaotion can be

found in Chapta- 14 on page 451.
Taste Processing in the Central Nervous Sx.stem

insular cortex The primary cortical After le..'l'\.;ng the taste buds through the crania l ne es, g wtatory info rmi\-
processing area fcr taste-the part of tion tra vels through W'-'Y s tntions in the medulla an d thala mus before reach-
the cortex that first receives taste lnfor· ing the insular cortex (also refe rred to as the gustatory cortex) (Figure 15.5)
rnatlon. Aloo call:ld lnsula or i;,tJStatl)()'
con ex. {Pritchard and Norgren, 2004). 111e functional sep:lration of the taste quali ties
s uggested to early investiga tors tha t there woul d likely be a gus toto p k ma p
in the cortex, but evidence for this h;)s been elusive. However, two recent
contributions have s hed n ew light on earlier findings. One study (in mice)
using a calci um imag ing technique identified clus ters o f cells showing neural
resp onsiveness to bitter, sweet, and sodium compo unds. In a dditio n, some
cells we !'e s timul ated by m on o po tassium g lutama te (the use of p o tassium
was intend ed to evoke g lutama te and no t salty in o rde r to locali ze urnami
taste) (Che n, 2011 ). Oddly, this p rocedure d id no t e voke responses to acids,
but the authors suggested that a sour ch.1ster might be outside the area they
FIG URE 15.5 Taste info rmatio n projects from the tongue to the m€ldulla (Via
OQl'V9S VII , IX, and X), then to the thalamus (shown in cross sQCf:lon 1 of the brain) ,
then to the insula (cross section 2), and finally to the oratofrontal cortex (cross
sectbn 3').

TASTE 475
sampled. The separation of th ose clusters is consistent with a g ustotopic map.

Even m ore exci ting, a meticulous new lesion-mapping strategy ha s localized
ta ste function ally by utiBzing conditioned taste ave rs ions {Schier et al., 2014).
The areas that underlie conditi oned tas te aversions fall into a "subregion en-
cmnpassing the posterior gustatory cortex and the s urrounding ins ular cortex
cons idered to be visceroceptive." Earl ier contradictions across s tudies may
reflect less p recise lesion mapping.
The orbitofrontal cortex receives projections from the insular cortex. Some
orbitofrontal neurons are multhnodal; that they resp ond to temperature,
touch, and s mell, as ' "'·ell as to taste, s uggesting that the orbitofr ontal cortex
may be an integrati on area.
Inhibition plays an important role in the processing of taste in.fonnation in
the brain On.e of the functions of th.is inhibition m ay be to protect our whole-
mo uth perception of tas te in the fuce of injuries to the taste system . Our br(lins
receive taste input from several nerves (see Figtue 15.5). Damage to one of them
diminishes its contribution to the whole; however1 that da mage also re.leases
the inhibition that is normally produced by the damaged nerve. 1l1e result is
that whole-m outh taste intens ities are relatively tmch•.-mged. Unforhmately, this orbitofrontal cortex (OFC) The
p reserved who le- mouth perception comes at a cost in some cases. Localized part of the frontal lobe of the cortex
that Ilea behind the bona (orbit) con -
taste damage is often accompanied by " phantom. taste 11 .sensations {recall the taining lhe eyes. The OFC Is respon-
phantom limbs experienced by many limb amputees, described in Chapter sible fa tha ccnsclous experience of
13), as if the release o f inhibition permits eveJ1 noise in the nervous system to dfactlon, as mll as the integration of
be perceived as a taste. pleasure and displeasure from food;
It has be€fl referrecl to as the second-
Descending inhibition from the taste cortex to a variety of o ther s tructures ary olfactory eo<tex and seconctl!y
may also serve o ther ftmctions, For exa mple, mouth injuries th.:1 t lead to oral taste oonex. The OFC is also lrwolved
pain make it harder to ei.lt. 11-'te inhibition o f s uch pain perceptions by taste- In many other functions. and it Is
processing parts of the brai n would m ake ea ting easier a nd thus increase crltlcal for assigning affective •ralue to
the likelihood of s urvh'al (beca use no mrttter h ow mu ch the mouth hurts1 stimuli -In other words, determlnlrg
we still ha ve to eat). Consisten t with this P-atients with a serio us h€donlc meanlrg.
oral pain d1sorder (burning mo uth syndrome) sh wt\ to hnve.Joccilized basic taste l>l'rf of the four taste
ta ste d amage as \'\-ell (Gn1shka and Bartoshuk, 2CKX.l). Purthem1ore, women qualltles that are generally agreed to
describe hlman laste expeneme:
w h o h ave tas te damage are m ore like ly to suffer from severe nausea and sweet, salty, sour, bitter.
vomiting during pregnancy (Sipior"1 et al., 2000); and can cer patien ts, whose
chemo thera py and radfatio n the.rnp y is known to damage the taste sys te m, salty One of the four basic tastes;
the taste quaftty produced by the
are m ore li ke ly to experience coughing, gagging, hiccups, and pain. In all cations of salts (e.g .. the sodium In
these cases, inhibitory signals from the taste cortex that n ormally help prevent sodium chlcrlde prOOuces the salty
ea ting-disrnptive sy mptom s (oral pain, vorn.iting, hiccuping, and so on) may taste). Some cations also produce
have been turned off beca use of the d amage to the taste systeJ:n. other taste qualities (e. g., potassium
tastes bitter as well as salty). lhl pur-
est salty taste Is proouood by sodium
The Four Basic Tastes chlalde (Naa), commm table salt.
sour One of tlie four basic tastes;
We learned in the Chapte r 14 that we are able to dis tinguish 1na ny different the taste quality produced by the
odorants. Already in this d'!apter, h owever, '"1e have seen that when olfaction hydrogen Ion In acids.
is taken out of the equation 1 much o f the complexity of the sensa tio ns evoked bitter Ole of the frur basic tastes;
by foods vanis hes. Tirns we are led to believe that the number of basic taste the taste quality, generally considered
qualities is qui te s mall. In fact1 the current universally accepted lis t includes unpleasant, produced by substances
only the four basic tastes previously mentioned in the "Taste Buds and Taste Ilka qulnlre or caffeine.
Receptor Cells 11 section o f this cha pter: salty, sour, bitter, and sweet. As '""·e sweet One of the four basic tastes;
di scuss these in the sections that follow, n ote that one of the mos t important the taste quality produced by some
features of these basic tastes is that om liking (or d isliking) for them is hard- sugars, sum as glucose. fructose,
w ired in the brain; that is, we are essentially born liking o r disliking these and sucrose. These three sugars are
particularly useful to us,
tastes. This is very different from the way we lea rn to like or dis like odors. and our sweet receptors are tuned to
Incidentally, the num ber of basic tastes h as been the s ubject of a rgmnent that them. Sane ot11er cornpoun:::Js (e.g.,
goes back even before Aristotle (see &1.rtoshuk, 1978). Various investigators saccharin, aspartame) are also sweet.

476 CHAPTER 15
FIGURE 15.6 Diagram of a taste (•) NaCl (b) HO

receptor cell, Illustrating the diffi9f'-
Outside cell Outside cell
'9nt receptor mechanisms for ionic
stirnuli (salty and sour}. {a) SaJty taste J;NaC H + ion-selective
Is produced by the cation in a salt. In ch.UUlel H"'
Naa. the sodium cation is admitted to
tl"la receptor cell by sodium
(ENaC). (/)} Sour taste is produced by
hydrogen io ns (H•), which can errter
tl"le csll in tw'O ways tile t9Xt for
details). C haudhari and Roper.
2010.) H• .............. H•Ac
lnsi.de cell blSid€'cell
have su ggested adding to the Ii.st of basic tastes. The two m os t l'e cent contenders
(umami and fatty) are discussed later because o f their special as.sociati.o n \vi th
leaJlle.d preferences. Since there is no de finition o f ' 1basic taste,'' it is easy to
call particular oral sens.:i tio ns ''basic.'' Hmvever, th e authors o f this text prefer
to reserve '1basic taste" fo r those with hardwired affect, since that is arguably
the characteristic of taste that m os t distinguishes it from the other senses.
\Vill huther d etails about a gusto topic map in the cortex settle the argtmwnt
over what we s ho uld ca ll "bask tastes" ? Sadly, no they \.\.' ill not. Consider the
new d ata on dlli"iers of cells in the cortex that responded to bitter, sweet, salty,
and t11na mi. If tas te quality is coded bya gustotopic m ap, then different taste
qualities must create different dusters.
Salty
Salts are made up of tv.·o charged particles: a cation (posi tively charged) and
an a n.ion (neg..1. tively charged). For example, common table salt is NaO; th e
sodi mn is the ca tion (Na+), and the chloride is the anion (Cl -). The source of
the s.a lty taste is the cation (Figure 15.6a). Alth ough all sa lts taste at a
little salty to humans, pure NaCl is the sa ltiest-tasting sa lt arotmd . Sodium
mu st be available in relatively large quantities in the body to m::iintain nerve
and m usc le fun ction, and loss of body sodium leads to a swift death.
O u.r ability to perceive sa ltiness is n ot static. Gary Bea uch amp a nd his
colleagu es (Bertino, Beauchamp, and Engelman, 1982) s howed that di et CL\l\
affect the perception of saltiness. Fortunately for those on low-sodium die ts,
reduced sodium intake increases the intensity of saltiness overtime. lndividu-
als who are initially su ccessful in reducing their sodium intake will find that
foods they used to love may now taste too sa lt): This adjustment in perception
helps them keep their soditm1 intake d own.
Our liking for sa ltiness is n ot static either. Early experiences can mod ify salt
preference. In 1978 a nd 1979,several hundred infants were fed soy fomrnlas
that were accidenta lly deficient in chloride beca use of an error in formulati on .
Chloride de ficiency has effects on human physiology that mim ic the effects
of_ sodiu111 d fi cie.ncy. the infants who \Vere dtloride-deficient offered
an important way to study odium deficiency in humans. The Centers for
Disease Control and Prevention (COC) in Atlanta monitored these infants,
and a of s tudies were done to assess a ny potential damage. On e of the
consequences was that the salt preference of the child.ren increased (l. J.Stein
et al., 1996). Experiences during gestatio n can also affect salt preference. For
example, college studen ts whose moth ers had experienced rnod era te to severe
morn.mg sickn ess during pregnancy sh owed an increased preference for salty
s nacks (Cryst.al and Bern.stein, 1995). The exact mechanism s by which these
abnormal me tabolic events enh .."1.nce salt pre fe rence are still not unders tood.

TASTE 477
Sour
As you may reme mber from high school chemis try, a solution containing
hydrogen ions (H+) and hydroxide ions (OH-) in equal proportions produces
water (HOH, or As the relative proportion of H+ increases (decrea sing
the pH level), the solution becom es m ore acidic. Why do you need to be re-
nUnded of all this? Beca.usesou1· taste is produced by hydrogen ions. Hydro-
gen ions enter the receptor cell throuohJon c;harmels; an ad ditional
mechanism for sour allows undissociated aad molecU.les (infuct molecules that
have not split into h.vo charged pmtides) to enter as well. The undissociated
acid molecules dissociate inside the cell. Ultirn1.1tely, the s timu.Jus that triggers
sour taste is the hydrogen concentration inside the rece ptor ce ll (Ftgure 15.6b}
(DeSim one et al., 2011).
Some illdiv;duals like the sourness of acids in relatively low concentrations.
fvtany adults enjoy pickles and sa ue rkraut, both of w hich get their sour tastes
from acids . In addition, many children in particular like sour candies (Liem
and Me1mella, 2003). At high con cen trations , h owever,acids "'ill damage both
externaJ a nd internal body tiss ues.
Bitter
The Human Genome Project revealed a mtJtigene family responsible for about
25 different bitter receptors. The HUGO Gene Nomenclature Committee has
established the rules for naming genes. (HUGO stands for Hmnan Genome
Organisation, an international organization of scientists involved in human
genetic and genomic research.) The bitter gene family is TAS2R ("TAS" stands
for tas te, with the "2" indicating bitter); numbers following the R indicate the
particular gene that is a member of that farnily. The25 bitter genes are loca ted
on three differentduomosomes: 5, 7, and 12. TI1e receptors these genes express
are designated without using it.alics (e.g., TAS2RHor T2R#).
The bittet receptors expressed by these genes face a formidable task.
Wolfgang f\·1 eyerhof, one of the world '.s experts on bitter genes, estimates that
there are thousands of bitter m olecules (many coming from plants that protect
from predators by tasting bitter). How can only 25 bitter receptors
handle the job? Part of the ansvver is tha t som e of the T2 receptors
only to specific compounds (e.g., th e propylthiourncil [PROP] receptor [see
Figure 15.lOb] responds to a s mall group of chemically related compo unds),
but o thers (bitter "genern.lists" ) respond to many compounds (Figure 15.7 ).
Th is also means that some bitter s timuli s timulate many bitter receptors {e.g.,
qui.nine stim ulates 9 o f the 25 bitter receptors).
Tonic wa ter (which cm1tains quinine) was originally fornmlated as a tre..1t-
1ne nt for malaria; now, however, we know that tonk water does not contain
enough quinine for that purpose. H owever, tonk water does contain enough
quinin e to taste very bitter, a nd for this reason lots of s ugar was added to
make the tonic water palatable. This approach works because sweet and bitter
tastes inhibit on e another; tonic water tastes much less bitter than the quinine
FIGURE 15 .7 BittGr l"o9CQptors ar9

designated byTAS2Rit, where# Is the
nurnbo9r o f tho9 r9ceptor. Th" rQOBP·
tor numbers arg shown at the top o f
the tabl9. Exampl9S of cornrnon bitte.r
compounds are list{ld at th9 l9ft. A fill9d
box indlcat9s a ri909Ptor that responds
to the bitter compound on the left. The
total numbQr of stimulat9d
by a gtvQi compound is shown on the
right. (After MQYQffiof et 31 .. 2010.)

4 78 CHAPTER 15
content alone wouldr and it also tastes much less sweet th.:"U1 the s ugar content
alone would Tonic water actually contains about the same amount of s ugar
that' sodas have.
Although a great rnany different compounds taste we genera ll y do
n ot d isting uish between the tastes of these comp mmds; we s imply avoid them
a ll. ll-1e diversi ty of receptors for bitterness enables species or even indiv iduals
in a given species to ha\'e va rying responses to an array of bitte r comp ounds.
One of the most fam ou s of these is 11 taste blindness'1 to phenylthiocarbamide
(PTC) found in human s--a phe nomenon we \.Yill revis it la ter in this chapter_
A.I though bitte r taste usuall y s ignals p oison , some bitter stimuli are actu-
ally good for us. For bitter com poun ds in some vegetables he lp
pro tec t against cancer. We would like to be ab le to "turn off'' these bitte r
sensati ons to make it easier for people to ea t their vegetables. Jn pursuit of
this goal, Robert Margolskee, a p ioneer in s tudies of bitter tran sduction, used
his understanding of the bitter system to identify a s ubstance that can inhibit
.sorne bitter sensations: adenosine mono phosphate (Afl.1P) (Ming, Ninomiya,
and Margolskee, 1999). Al\llP may actua lly hmction as a natura l bitter inhibi-
tor in mo ther 's milk. A number of compounds in m.iLk, su ch as casein (milk
protein ) a nd calci um sal ts, taste bitter; and, as we will see aversions to
bitter tastes are present a t birth. The presence of AMP in mo ther 's milk may
s uppress those bitte r taste.s enough to a llow milk to be pa latable to babies \.vho
are particul arly respons ive to the m .
Bitte r perception is also affected by hormone levels jn women. Sens itivity
to bitterness inten sifies during a nd diminis hes a fte r men opause
(Duffy et al., 1998). These d ifferences ma ke sense in the context o f the foncti on
of bitte rness as a p oison d etection med1anism. Intensifyin g the pe rcepti on of
bitter ea rly in p regnan cy, when toxins exert thei r maximum e ffects, has clear
biological value. Consistent with this coi-relation, some o f the aversions during
pregnan cy occur w ith foods o r bevera ges tha t h.ave bitter (e.g., coffee).
One of the m os t interesting recen t develop ments oon ceming bitter is the
discovery th.a t bitte r receptors CE\ll be found on locati ons o ther th.an the tongue.
For exam pier bitter receptors in the gut appear to s low absorption Qeon et al .,
2008). flliusf f tQxlns get µasl l)ie-n101)th, tl>e g ut can still preven t p oisoning.
Sweet
Sweebless is evoked by s ugars, simple carbohydrates that generally con foml
to the chemical fomm la (CH 20),,, w here n isa number between 3 and 7. G lu-
cose, o ne o f the S\'\-·eetest-tasting sug.1rs, is the principal source o f e nergy in
FIGURE 15.8 The molecular struc- humans (as wel l as nearly every o ther livi ng thing o n Earth). Common tab le
ture of sucrose, commo n table sugar. s ugar, su crose (Figure 1 5.B)--w hich is a oombin ation of glucose and yet an-
This disaccharide is formed from a oth er su ga r, even sweete r.
combination o f a glucose molecule and
The bio logical fonction of sweet is diffe re nt from that o f bitter, a nd the
a fru ctoSiQ moh;icule. G lucose, which
is easily extracted fro m sucrose by the way taste receptors are h.med suppo rts th at bi ological diffe re n ce. Many dif-
digestive system. is the maJn fuel that ferent molecules taste bitter. Our bio logical task is not to distinguish among
powers ahrost every biologic al engln9
(including the human brain c urrently
reading this b ook). c.c-1 ,2 Glrcosidic linkage
,...-->----.,
vf-":"1, :6)--4'"'"'""
+
OMH
H
GI H
OH
H
t
0
2
M O
H
CHj:)H
H OH OH H
a-0-Glucose F11.lc tose a:-o-G\uoost.> Fruc tose

S ucrose

TASTE 479
Outside cell FIGURE 15.9 Structure of the T1R2-

T1 R3 heterodltner sweet receptor ,
showing binding sites for both large
and small SVV99t rnol9CUIQS. (Afto9r
T ernussl, 2007 .)
the m but ra ther to avoid them a ll. Thus, we have multiple bitter receptors to heterod mer A chain of t1MJ mol-
encompass the chemical diversity of poisons, but they alJ feed into comm on ecules that are different Iran each
lines le1ding to rejection. With regard tosvi,•eet, sorne bio logica lly llseless s ugars other.
have structures vel'y similar to those of glucose, fmctose, and sucrose. [n this
case, then, the task of the taste system is to tune recep tors so specificall y that
the biologically important sugars stimulate sweet taste but the others do not.
Consis tent w ith the biological purp ose, only h'lo G protein -co upled
recep tors are involved wi th sweet taste (Tl R2 and TI R3) , These two proteins
combine to form a singl.e recep tor ca lled a heterodlmer (Figure 15.9). ll1.e
oute r p or tions of the two proteins resemble the shape of Venus fl y traps;
large sweetene r molecules can enter the fly traps to stimula te the receptor. A
variety of othe r binding sites accommoda te s maller molec ules like su ga rs,
sacchari n, a nd aspartame. Lnitia lly, the TI R2-Tl R3 he terodimer was thou g ht
to be responsib le fo r a ll 5'"/eetness, but it introduced a new puzzle: No matter
how the heterodfo1er is stimulated , the receptOI' produces on ly one signal.
Th erefore, we \vould expect a ll sweeteners--sugars and artifidal sweeten ers
alike-to produce the same sweetness. However, artificial sweete ne rs like
sacch ari n and aspartame d o no t taste exactly like su gar; if they did, there
would be no need to continually search for better artificial sweeteners. Some
claim tha t arti ficial swee teners Produce additio na l tastes tha t -account for the
difference. For example, saccharin tastes bitter as well as S\Veet to many. But
some of us (the au th ors included. ) do not taste the bitterness of saccharin a t
all; \-V"e are quite ronvin ced tha t it is the na hu-e of th e sweetness that differs.
Geneticsh1dies may offer some help he re . TI1e receptor TlR3 appears to be able
to fun ction alone to respond only to high of sucrose (Zhao et
al., 2003). In addition, Margolskee and colleagu es (Yee et al., 201 1) have identi-
fied yet an other mechanism in addition to the h eterodi mer that can mediate
sweetness. Thus we can exp lain w hy m.."l. ny of us perceive diffe rences between
th e sweetness of s ucrose and those of artificial sweetene rs.
Sin ce our taste system can produce sweet receptors so precisely tuned to
the biologically usehtl su gars, \vha t is going on wi th artificial sweeten ers?
Are the.se nonsu gar molecules sweet because they accid enta lly stimula te the
S\\' eet heterodimer? We don ' t know. Are artificial sweetene rs medically useful?
Certainly they en able diabetics to enjoy a swee t taste wi thout the d an gers of
sugar, but the ea rly hopes tha t artificia l sweeteners wo uld be a p an acea fo r
weight loss now appear VvTon g,
Saccharin w,'l.S discovered in 1879 w hen the chem.ist [ra Re msen, working on
coal tar d eriva tives, failed to wash up before dinne r and subseciuently no ti ced
tha t the ta r residue on h is han ds tasted sweet. Another artificial sweetene r was

480 CHAPTER 15
nontaster (of PTC/PROP) An lrdl- d iscovered in 1937, by a graduate student working in w ha t must have been a
vldual born "1th t1M> recessive alle>is messy lab. When he tas ted sweet while s1noki.ng a cigarette, he realized that
for the TAS2R38 gene and unable to son:1ecom pound in the lab must be responsible. In th is way, cydamate \"·as
taste the ccmpounds phffiYlthlocarba-
rnlde and prcp)1thlruracll. d iscovered . Cyclam.ate was very p opula r for a few years, but s us picions were
raised that it was carcin ogenic.A lthough those Slb"'Picion.s were ch allenged and
taster (Of PTC/PROP) An Individual
cyclam.ate remains lega l in a munbe r of othe r cotmtries, the Food and Drug
bcrn with one a OOth dominant alleles
for the TAS2R38 gene and able to Administration (FD A) banned it in the United States. Artificial sweeteners
taste the ccmpounds phffiYlthlocar- are attractive to dieters because their s\veet taste comes \'\oi th essentially no
bamlde and prop)1thlouracll. PTC/ calories. Millions of dieters count on this p roperty to help the m watch th eir
PRO P tasters wro
also hao;e a high weight1 but a 1986epidemiological s tudy showed that wome n w ho cons tuned
denslrf of funglform papillae are PROP
supertai::.ters. artificial sweeteners actually gain ed weight (Ste!Lnan, 1988).
Also in 1986, John Blundell, an English expert on weight regula tion, published
a provocative article s u ggestin g thilt asparta me (the arti ficial sweetener sold
as NutraSweet) in creases app eti te (B!tmdell and Hill, 1986). \Vas the benefit of
the redttced calories lost w hen ind ividuals using rtSprut a me actually increased
thei r ca loric in rake in subseq ue nt meals? Terry Davidson and Susan Swithers
elaborated on this kind o f thinking lNi th s h1dies u si ng rats (T. L. D,1vidson
and Swi thers, 2004). l11e work wi th humans was d iscounted by m an y;
after all, those wi th weight problems m ay be the individuals w ho choose to
consume a rtificial S\•1..-eete.ners a nd it is not s urprising tha t they gain weig ht.
H owever, the sam e cri ticism could no t be made w hen the rats fed artificial
swee teners gained weight. Davidson nnd S""ri thers point to the fact that the
"obesity ep idem ic" occurred over the sa me years as the introduction of low-
calorie foods into the Am erican m arket. They argue tha t the tmcouplin g of
the sensory properties of these diet foods from their metabolic conseq uences
d isrupts regu lati on , lea ding to '""eight gain.
lncide n tally, theso--called obesi ty e pidemic deserves closer study. Katherine
Flegal, an epid emiologist w ho m onitors weight cha nges in the United States,
s howed that the drita telJ a story not well char<1cterized by epidemic (Flega l
et al, 2002). \ Veight is roughly normally distributed . TI1a t distribution is n ot
mo·dn g as a w hole toward h..ighe r weigh ts. Rn.ther1 the upper a rm of tha t
distribution is eleva ting. That is, we are n ot all getting heavier; those w ho are
(a) Pheny lthiocarh1n-Ud.e already overweight are ge tting hea\rier.
H
H I
H 1 N - C- NH, Genetic Variation in Bitter
II - Jn 1931 a ch em ist named Arthur Fox discovered tha t we do no t all live in the
II I s sa m e taste world (A. L. Fox, 1931). Fox was synthesizing the compound PTC
(Figure 15.10a) w hen some spilled and flew into the air. A colleague nearby
I n oticed. a bitter taste, but Fox tasted nothing . A test of add itional colleagu es
H revealed a few m ore nontasters like Fox w h o tasted little bitterness in the
(b) Pmpy lthiourocil compound1 but most (tasters) perceived it as bitter. The next year, Albert
H Blakeslee (a famous geneticist of the day) and Fox took PTC crystals to a meet-
"c,,. .
t CJ.L,CH 1 CH3 ing of the A merican Association for the Advancernent of Scien ce and se t up
a votin g boo th for attendees to register their p erceptions . About on e-third of
I I those polled fo und the crystals to be tasteless, whi le two-thirds found them
to be bitter. These proportions captured the imagina tion of many research ers,
111n
ti f'or severa l years the /ournf1f oJHereditysold papers impregn.ated with PTC
II for further studies. Family s tudies eventually confirmed tha t t(1ster status is
an inherited trai t (e.g., the Dionne quintuplets were all f01.ind to be tasters in
FIG UR E 15.1 o The chemical struc-
1941). Nontasters carry hvo recessive alle les, whereas tasters have either one
tur€<S of PTC (•) IXld PROP (b) . The o r both domimmt a lleles.
portions shadOO in bkJQ arg thosa lnitiaU)i ind ividuals were si mply classified according to w hether they co uld
responsibl9 for the bitter taste. taste PTC, but evenhmlly threshold s h.1dies came into vogue. In a threshold

TASTE 481
me thod invented specifically for PTC s tudies, subjects \Vere given eight cups,
four containing and four contai ning a given concentration o f PTC.
Correct sorting de termined the thres hold. The distribution of thres h olds was
bimodal, with non tas ters showing very high thresholds and tasters showing
low thresholds . Thi s distribution varied by sex and race: women had lower
thresholds than 1T1en, and Asians had lower thres holds than Caucasians.
Jn the 1960s, Roland Fischer shifted studies to a chemical relative of PTC
that was safer to test-propylthiouracil (PROP) (Figure 15.1 Ob)-and focused
on the nutriti onal implications of the genetic variation in taster s tatus (Fischer
and Griffin, 1964}. Fischer suggested that tasters were rnore fin icky ea ters:
because bitter tas tes are m ore in tense to these individuals, they tend to d is like
foods hi gh in bitter cornpounds,such as m any vegetables, that n on tasters find
m ore palatable. Fischer also rela ted tas ter s tatus to body type (e.g., weight)
and health . ..\lcoh olics and sm okers were found to contain a lower proportion
of tasters than would be expected by chance, pres mnably because unpleasant
sensations (e.g., bitterness) produced by alcoholic beverages and tobacco act
ns deterrents. The effect o f genetic variation in ta ste is even related to c.mcer
risk, as will be described s hortl y.
In 2003, Dennis Drayna and his colleagu es d iscovered the location of the
gene that exp resses PTC / PRO P recepto rs (Kim e t al., 2003). This gen e is a
member of the bitter family introduced ear lier and is designated TAS2R38.
lndividuals with two recessive alleles are nontasters; those wi th either one or
both d omir1ant a lleles are tasters.
Supertasters
By the 1970s, the "direct'' psych ophysicaJ methods introduced by H..1 rvard's
S. S. Stevens led to a n ew look at gene tic varia tion in taste. Ins tead of measur-
ing thresh olds-the dinunes t could look at supra-
thresh old tas te and plot the psycho physical functions sJ1o·wing how perceived
taste intensity varies w ith concentratio n.. You m ay reme mbe r from C hapter
1 that Stevens nnd his students d ocumented what is now kno"",' as Stevens'
power law. It h olds that perceived se ns ations rise as the .stimulus raised to
som e power. Written ns an equation, this is
5
w here S is sensation, I is stimulu s intensity (concentrati on of a solution if we' re
talkingabo11ttaste, and we are), and btakes on different values for different sen-
sory m odalitie; (Stevens and Galanter, 1957). Of special interest for our present
purposes, b takes on different values for different taste qualities. For example,
bitterness grows more s lowly v"·i th concentration th.an swee t d oes (Figure
15.11 ). Th'1t is, the value of b for bitter is smaller th an the value of b for S\"·eet.
0.00001
Molarconcentr.1tion
7 FIGURE 1 5.1 1 Psychophysical func tions for quinine
and sucrOS9. The logarithm o f the perC0ivad taste intansi·
ty is Potted against the logarithm of the ccncerrtrntion. In
this plot, bis the slope o f the function. The value of b for
quining Is 0.3; for suc rOS9, 0.8. (After Bartoshuk, 1Q79.)

482 CHAPTER 15
cross-modality matching The Unfortunately, know ing h ow taste intens ity grows with concentration
ability to match the Intensities of d oes n ot permit comparisons o f perceived taste intensity across individuals;
sensations that come from different b can be equal for two people even if one experiences taste in tensities h .vice as
soosory modalities. This ability enables
Insight Into sensay differences. For great as the othe r. Fortunately, t\.vo of S.S. Stei.-·en.s' s rudents-Joseph Stevens
example, a Hstener might adjust the and Ut\'\<Jence Marks-mad e an other ftmdamental discovery in th.is era . As
brightness of a light untll It matches the we d iscu ssed in C hapte r 1, humans are s urpris ing ly good a t crossRmodaltty
loudress of a tone. matching (Stevens, 1959 ). For example, we can match the loudness of a sound
supertaster An individual whose to the brightness of a Light, and we can m atch both of these to the intensity of
perception of taste sensations Is the a taste. This finding ultimately led to the discovery of su pertasters. Katherine
most Intense. A variety of fac tcrs may Fast, on e of Barto.shuk's s rudents, asked s ubjects to matd1 the bitterness o f
contribute to this helJhtened percep-
tion: among the most lmpottant is the PROP to other sen sa tions compl etely unrelated to taste (see Figure 1.9). For
density of funglform papillae. example, nontasters matched the taste of PROP to very \llteak sensations (the
tick of a watd1 or a w hisper). Tasters proved to be a he terogeneous lot. For
some, PROP '"'as likened to very intense sensation s, s uch as the brighh1ess
of the s un or the most intense pa in ever experienced . 1}1ese individuals were
labeled supertasters. uf\·1edium tasters" m a tched PROP to weaker s timuli,
su ch as the .smell of frying bacon or the pain o f a mild head ache (B.utoshuk,
Fast, and Snyder 2005).
Armed wi th genotypes, ratings of the bitterness of PROP, a nd C'O\.Ul ts of
8230336 Amma 1ungtform papillae, Valerie Duffy (ci. pion eer in the m odem m ovement among
nutritionists to eval uate food behavior in terms of th e sensory properties of
foods instead of only their nutrien t content), along with he r studen t John Hayes
and coll eagues (Hayes e t al.1 2008), was able to show that a combinati on o f
PROP gen otype and tongue anatorny divided indhriduals into three groups:
nontasters, tasters, and s upertasters of PROP. PROP s tatus and number o f
hmgiform papillae turned out to be indep endent. The combin ation of a bility
to tas te PROP and p ossession of a very large number of fungifonn p ap illae
produces a PROP supertaster; PROP trtsters vv·ith fewe r fungiforn1 papiJl ae are
medium PRO P tasters. Am ong PROP non tasters, the number of fungifom1
papillae va ries across the w hole range. A PROP nontaste.r m ay be a su per tas ter
for othe r tastes, but can never be a PROP su pertaster.
TI1ose \vi th the most munerous fungi fo rm papillae experience n ot only the
most intense taste sensa ti ons in genera l, but also the m ost intense se.n:s3tions
of oral bum (e.g., from chili p eppers) and oral touch (from fats or thickeners in
foods), because fung iform papillae are inn ervated by nerve fibe rs that con vey
burn a nd touch sensations, as well as those tha t con vey taste sensation s. In
addition, because of central conn ections between taste and retronasal olfoction,
those who experien ce the most intense taste sensa tion s also perceive more
intense retronasal olfo.ctio n and thus more intense flavor.
Pathology adds ye t ano the r sou rce of varia tion in our oraJ sen sati on s.
Surprisingly, removing input from one of the taste n erves (by an esthesia or
damage) can actually intensify taste sensa tions (Bartoshuk e t 2012). This
intens ifi cation rem its beca use inputs from the different tas te netvesinhibit one
another in the brain . Loss of input from one nerve releases o thers from that
inhibition. In some cases, the result is actu al intensification of w hole-mouth
tas te. When da m age is more \o\"i despre3d, not s urprising ly, taste pe rception
din1inish es.
In s mn, taste experience is the outcome of genetic variati on, pa thology, and
interactions among all of the cranial nerves innervating the mouth.
Health Consequences of Taste Sensation

With these new insig hts, the potentia l links beh\reen resp on siveness to PROP
and hea lth ha ve become the focu s of ren ewed interest. The new psychophysi-
ca l metho ds p ermitted Duffy to s how tha t variation in the sensory prope rties

TASTE 483
of food s and beverages affects food preferences and thus diet. Because diet
is a major risk factor for a variety of diseases, genetic variation in ta ste plays
a role in th ese diseases. For in.S tance, some vegetables produce unpleasa nt
sensations (e.g., bitter) to medium tas ters and s upertasters, leading these
individuals to eat feiA.•er of the m. Reduced vegetable intake is in turn a ris k
factor for colon cancer.
Sure enough, Duffy and her coll eagi.tes found that, in a sample of older men
getting routine colon oocopies at a Departrnent of Veterans Affairs hospital, those
tasting PROP as m ost bitter had the m ost colon polyps, a precursor to colon
ca ncer (Basson e t al., 2003). On the other hand, fats can produce unpleasantly
intense sensations in s upe rtasters, leading thern to eat fewer high-fat foods and
thereby lowering their risk of cardiovascular disease (Duffy et al., 2004). These
sensory links to behavior that affects health are not limited to diet. Fischer 's
early sugges tion that n on tasters are more likely to smoke and co1tstm1e alcohol
has proved correct (Duffy et al., 2004; Snyder et al., 2005).
More generally, varia tion in the sensory p ropetties of foods/ beverages is
linked to health because that variation affects wha. t V\'e like to ea t, and diet
affects he...1lth. ln addition to variation due to genetic differen ces, there is taste
damage due to common pathologies. The pathways of the t::iste nerves from
the tongue to the brain are v ulnerable to damage. The d 1ord a tympani taste
n erve pa!is:es through the middle ear, w here it is v ulnerabl e to middle-ear
infections (otitis media). The g lossoph.arynge...11 taste ne rve is physically n ear
th e tons ils. A layer of muscle can protect the glossoph aryngeal n erve dur-
ing tons illectomy, but that muscle is bcking in some individuals. Both the
chorda ty rnpani and the glossopharyngeal nerves can be darn.aged by head
injuries even if they are rela tively mild . Taste d amage can have w1expected
conseque n1..-es because of inhibitory connecti ons be h>1.1een taste nerves and
between taste arid other oral sensations. Recentl}'i indi viduals with his tories
of o titis media (usually in infancy ordllldhood), tonsillec tom y, o r head injury
were fo und to gain weight (Bartoshuk, 2012; BartoshtL.k., et al. 2013). Sensory
tes ting revealed a likely scena_ri o. The localized taste damage released inhibi-
tion on und ..unaged nerves s uch that whole- mouth taste, perception of tou ch
(fats in foods), and retronasal olfaction all increased. T11ese sensory changes
were associated with enhanced palatability o f high-fut foods, which could
potentially lea d to weight gain.
Wisdom of the Body:

How Do We Solve the "Omnivore's Dilemma" ?
Some species h ave few food ch oices to ma ke; for exarnple, koala bears eat
primarily leaves from the eucalyptus tree . However; humans (and rats1 rue Amma Appa
omnivores; we are confronted with an rtJTa.y of choices, and we must
foods and avoid poisons to s urvive. Paul Rozin coine<.i the term "omnivore's
dilemma" to describe this biological imperative (Rozin, 1981). Michael Pollan
subseq uently applied the tem1 to the modern human 1s need to find a healthy
diet amidst the dizzying dloices available to us today (Pollan, 2006)_How d o
the chemirn l senses ope rate to help us? TI1e specific properties of ta ste and
olfaction cooperate to help us s urvive.
Llke olfactory receptors, taste receptors detect specific features of rnolecu.les.
But the two associated senses evolved to serve quite different fonctions . In
olfaction, specific fea tu res of m olecules interact vvith receptors tuned to the m.
All ra--eptors of a given type projec t to indh.;du,al glomeruli in the olfactory
hr.'\in. The pattem across the glomemli paints a picture of the chemical structure
of the m olecule. These pictures help us identify objects in our environment.

484 CHAPTER 15
Indeed, fo r many animals (e.g., roden ts), olfaction is the primary means for
knowing what s urrounds them in the \.\'Orld. Consis tent with this purpooe, the
olfactory system is capnble of distinguishing among a large number of differ-
ent mol ecules, and an individual a nimal can lea rn about wh atever olfactory
stimuli exist in the environment where it happens to live.
FURTHER DISCUSSION of the sensing of molecular structure as it relates

to olfaotion can be found in O.apter 14 starting on page 43Q.
On the other hand, we've just seen that th.e taste system responds to a fixed
and much s maller se t of m olecules thatn ahue knows we will encounter. This
precise tuning is consistent \'lri th the role of taste as a system fo r de tecting
nutrients and 11 antinutrients" (substances that are eithe r helpful or harmful,
respectively, to our bodies) before we ingest them.
Each of the four basi.c tastes is responsible for a different nutrient or an-
tinutrient, and h as evo lved accordin g to its purp ose. For exa mple, the bitter
taste subsystem is nature's poison d e tector. In terms of structu re,
p oisons are quite d iverse. T11LLS the bitter receptors must be di verse as well
On the o ther given th..1.t we d on ' t really care if we can discriminate
among p oisons (since we just W;)Ilt to avoid them all), we could hook all of
those receptors up to a few common lines to the brain. As we saw earlier (see
Figure 15.7}, this is exactly how the bitter subsystern is set up.
Simila rly, the sour s ubsys tem is configured to reject any highly acidic
solutio n wi tho ut dis ting ui shing exactly \vha t is ca using the aci dity of the
solution to be so high. TI1e other hvo tas te .subsyste ms en able us to detect,
and therefore selectively ingest, foods tha t our bodies need: sodi um (salty)
and s ugars
Som e of the most m1p eevi'den""e for this hardwired affect with taste
came from the work of Jacob Steiner on facial expres.sions in newborn infan ts
(Stein er, 1973). Steiner found tha t infants responded with s tereotyped facial
express.ions w hen S\\teet, saJty, sour, .ind bitte r solutions were applied to their
ton gues. Swee t evoked a 11smilelike"' expression fo ll owed by s ucking (Figure
15.12a). Sour produced pursing and protrusion of the lips (Figure 15.12b).
Bitter p roduced gaping, m ovemen ts of s pitting, a nd in some cases, vomiting
m ovements. Even infants bon1 wi thout cerebral hemis pheres (a condition
(a) (b)
FIGURE 15.12 The two toddl.;irs' faciaJ expressions reveaJ the taste qualities that
th9y're 9>::p€1rfenc ing. (a) Sw00t potato produces the typical smil0 associated with the
acoe.ptancg of (b) Green ai:;ple produc es the puckefy fac.e associated w ith

TASTE 485
known as anenceph aly) s howed the sam e facial expressions, s uggesting tha t
these expressions are med ia ted. by very primitive parts of the brain,
The fact that the basic tastes provide both information and pleasure (affect)
enables organism s to solve critical nutritional problems irnmediately, without
having to learn {w hich takes time) . The newborn baby can nurse because the
sweet taste of m othe r 's rnilk is pleasurable. The baby can also reject poisons
because tbe bitter tastes they evoke are aversive. The athlete who sweats or
the new m other w ho loses blood can replace lost sodi urn because the taste of
salt is pleasant.
How Do We Regulate Nutrients? Early Belief in "Specific
Hungers" Gave Way to Identification of Conditioned
Preferences and Aversions
Early on, Cwt Richter p roposed a simple mechanism for the regulation of speclnc hlllgers theory The Idea
nutrients: the spectflc hungers theory. According to this view, the need for that de11clency of a given nutrient
a nutrient causes the body to crave it. For example, a boy '\vith produces c raving (a specific hunger)
fo r that nutnent. Curt Richter first pro-
an intense cravin g for sa lt died when his salt intake was restricted during posoo \hlo theory and demonstrated
a hospital stay. An au topsy revealed a hm'!Or o f his adrenal, gland tha alAngs for salty Cf fcr sweet
caused his body to lose sodium. Hi s salt craving had en abled his body to re- ,..e associated v.1th deftc lendes In
tain enough sodi um to keep him alive (Wilkins and Ri chter, 1Q40). Another those substances. However. the Idea
proved wrong for other nutnents (e.g. ,
source of s upport fo r specific hungers was a treatmen t for schizophrenia tha t
IAtamk'ls).
\";as p opular in the 1940s. At the tirne, some experts believed that the brain,
which depends on glu cose for fuel, could be forcib ly rested if blood g lucose
""'ere driven to very low values with insulin. ln tense cravings for svveet were
an unex pected by-product of the thera p y. Subsequent labora tory s tudies
confirmed that insulin injections produce increased liking for sweet. Rich ter
concluded that ingesti on of the nutrient reduces the craving a nd brings the
body back into balance.
Even more s uppo rt for the idea of s pecific hungers seem ed to come from
the '\Vork pediatrician, Clara Da vis. She allowed a group of 6-mon th-old
infants to ea t wha tever they Hked, to see if they would choose '\visely (C. M .
Davis, 1928). T11e infants thrived, leading Dav ls to concl ud e tha t, wh en a l-
lmved to choose am ong a variety o f healthy foo ds, infants had the ability to
select a healthy diet.
The early .succes.s of the s pecific hungers theory spurred furthe r investiga-
tions that ultimately proved that the theory was limited only to sweet and.
salty. In one of the early s tudies, rats were fed a diet deficient in vitamin B1,
w hich made them sick. When the rats were offered a choice of remaining on
the sa me diet o r switching to a diet containing B11 they immediately svvitched.
But Paul Rozin conducted a crucial control. He gave the control rats the choice
of the origina l diet or a different diet tha t was also deficient in B.i· These rats
also innnediately switched to the different diet. ll1 us 1 the rats in the original
s tud y had no t specifically sought B1; they had simply learned to avoid the
diet tha t made them ill (Rozin, 1967).
Rozi n' s work ended belief in specific hungers as an explanation of dietary
regulation for anything beyond s ugar and sa lt. In retrospect, we can see tha t
th e theory lacked an important ingredient. For craving to cause an animal
to seek out and take in a needed nutrient, a senso ry cue wo uld h.:1 ve to be
unambigu ous ly a ssociated wi th the nutrient. The saltiness of s..1. lt and the
sweetness of s ugar could as s ud1 cu es, but the B1 molecule d oes not
produce a detectable cue in food (Figure 15.13).
\Ve were left vvith a problem, though. H ow did C lam Davis's infan ts know
how to select a healthy diet if specific lnmger s d o not opernte for all nutrients?
It turned out that they '"'·ere not selectfoga healthy diet at all. They were simply

486 CHAPTER 15
FIG URE 15.13 In our QVOIUtionary past, when food was sc=:-trcQ
in
which foods are P'9ntiful &"Id ieaslly obtained, thes9 sp9cific hungers
(combined with the profit motive fa the food industry) lead many to
consumG too much junk food. The nutr\Qnts in wgeUlb49s are. alas,
largety undetect;Jb(Q, so wa cannot dw91op specific h.JngQfS fa
them.
e ;Jting a variety of the foods presented beca u se they got bored

eC1ting single foods. l11is phenomenon is called "sensory specific
sa tiety" (Rolls, 1986). Because all of the choices were he<'l lthy,
all the infants needed to d o was eat a variety. In the mo dern
\'l1orld, ea ting w hatever we like w ill not produce good heal th,
because too m any of the available foods are not hea lthy. In fact,
the specific hungers that are genuine can do us conside rable
ha rm; just think about our love for sweet and salty jtmk foods.
Rozin ended the belief that the brain was hardwired to te.LI t ts
w ha t to eat. But he offered us a mud 1 m ore fle..xible guide: learn-
ing. We come to Ii ke or dislike foods based o n the conseq uences
of cons umin g the m. \ Vhen the con.sequences are negative (e.g.,
nallSlea), we disl.ike the food and it (conditioned aversion);
\vhen the con.seque nces a re positive {e.g., ca lories), we like the
food and consume it (conditi oned preference). As we S.."\W in
C hapte r 14 1 olfactory likes a nd d islikes a re n ot hMdwired as
those for taste are.
1l'le associa tion of negati ve or positive affect with an experie nce is ca lled
"evaluati ve conditioning .1' The phenomenon has received remarkably little
attention given the powerful role that affect plays in our lives. The exp erience
of va lue is important in our social worlds (e.g., in religion, politics) as well
as in our food worlds. The classic paradigm of eva luative conditioning is
the trans fer of affect from a stimulus wi th positive (or neg..'1.tive.) valence to a
stirnulu.s. Por exa111ple, Debra Zellner and Paul Rozin conducted an
experi me nt ostens ibly about preference for teas. Subjects were asked to rate the
palatability o f a variety of teas; some had s uga r added, some did not. Tested
a second time, the teas that h ad originally been paired with sugar were rated
more positive.
Up to n ov.,.., resea rch h as su ggested that very yow1g human children are
n eutral to odors that attract or repel adults. Consequently, grandma 's cook-
ies m<.'ly lead one individual to love butterscotch, and a bou t of s tom ach flu
may lead another to hate it. Once n given odornnt has taken on v alen ce, it
ca n transfer that valence to other odorants by evaluative conditioaing.: tlds
pn:x::ess ca n p resumably lead to the rich and v aried pleasure we experience
from our food worlds.
1l1e gastrointestinal tract (conunonly refe rred to as the gut) plays a crucial
role in this learning. The rnacronutrients {nutrients that provideca lo1·ies) are
ca rbohydrates (long d1ains ot sugar molecules), proteins (chains of amino
acids), and fats (d'la ins of fatty acids att,1ched to glycerol). Except for the spe-
cial case of sugar (a very short chain), these macronutrient molecules are too
large to be sensed by taste or olfactory receptors. \ Vhen foods containin g these
macronutrients e nte.r the gut, how-ever, the macronutrie nts are broken into their
constituent pieces. These piecesstimulate d1emoreceptors in the gut, and the
brain makes u s like the sensory prope rties (prinlarily the retrona.sa l olfactory
sensation s) of the foods {conditioned food p references). On the other hand,

TASTE 487
if we experien ce na usea after eating, the brain makes us dis like the senso ry
properties of the foods (conditioned food avers ions). Thus, we regulate our
intake through a combination of ha:rd"t>vired basic tastes and learned responses
to food flavors.
The Special Case of Umami

Umaml arose as a candidate for a fifth basic taste as part of advertising claims umami The taste ·s ensation pro-
by manufacturers of monosodium glutamate (MSG), the sodium salt o f glu- duced by morosodium glutamate.
tamicacid. Identified by Japanese chem ists in the early 1900s, MSG was initially monosodium glutamate
marketed as a flavor enhancer, said to suppress unpleasant tastes and enhance (MSG) The sodium salt of glutamlc
pleasant ones. Tas te experts expressed skepticis m, MSG manufacturers went acid (an amino acid).
on to claim tha t MSG was a fifth basic taste, speculating that it signaled pro-
tein and thus played an important role in nutrition. Unfortunately, although
having special receptors for proteins might be nutrition.ally h elpful, protein
m olecules are too large to s timulate taste or olfaction . Glutam..<i.te isa part of a
protein m olecule, but m os t p rotein in the mouth is intact and not broken into its
parts. Further, the taste evoked by glutama te is not perceptible in many foods
containing protei ns. Finally, som e indivi duals like umami and som e do no t.
The four basic tastes, in contrast, are distinguished by their ha rdw ired affect.
There is an e1n erging and interesti ng d isti nction between the phys iology
and the psychophysics o f taste. Physiologically, it seem s th at there are gluta-
m ate receptors in the mouth and the g ut and that recepti on of glutamate has
consequences. Psychophysica lly, it seen'5 less likely that s timula tion of these
receptors gives rise to sensa tions that would qualify tuna mi as a basic taste
al ong with the classical sweet, sour, salty, and bitter:
Because g lutama te is a n important neuro trans mitter, receptors for the
m olecule are common throughout the body. The argument that sud1 receptors
might have been harnessed by the tas te system to s ignal uma m:i gained respect-
ability when neuroscientists Nirupa Ch;Judhmi and Steve Roper identified a
version o f a glutamate receptor in mt trlste papillae. Lnter, another receptor for
was identified: a heterodim er made up of two G protein--coupled
receptors: Tl Rl and TIR3 (remember that Tl R2 and T1R3 fo rm the sweet
heterodin1er) (Cha udhari, Pereira, a nd Rop er, 2009}.
Robert Margo lskee {whose pioneering s tudies foci.JSed on not only the bitter
receptor but also the sweet receptor) was an ea rly advocate for the importan ce
of taste receptors in the gut. We n ow know that taste receptors are not limited
to the mouth but rather can be found througho ut the gastrointestinal tract. This
exciting discovery suggests a different function for umami. Protein mo lecules
are too large to be sensed by taste or smell, but proteins are m ade up of amino
acids, including g lutam.ic acid . When eaten, proteins are broken d ow n into
providing stim uli fo r the glutamate receptors in
ihe gu t. These recept.tors an signal the brain that pro tein h as been cons umed .
ln this way, glutamate receptors ca n fulfill the function attributed to umarni,
but this is done in the g ut, not the 1nouth . . Consistent with th.is finding, Jolm
Prescott (a cognitive psychologist who s tudies the che mical senses) s h owed
tha t consu ming a novel-flavored soup wi th added to it p roduces a con-
ditioned preference for the novel fl;;ivor, "vhile simply holding the soup in the
m outh d oes n ot (Prescott 2004). Note that usinggluram ate receptors in the gut
to signal pro tein makes biological sense. This allows many different proteins
{that d o not tas te of tunami ) to evoke pleasure: not hard w ired p leasure but,
rather, learned pleasure.
Because glutamate is a ne urotrans mitter, con cerns have been raised about
its safety in the human diet. became particularly notorious in the 1960s.
Firs t it became associa ted with 01.inese restaurant S}Tid rorne---a constellation

488 CHAPTER 15
of symptoms including numbness, flus hing, tingling,s,veating, and

tighb"iess in the chest-that was reported by som e individuals after oonsmning
MSG (R.H. M. Kwok, 1968). Then Dr.John Olney,a toxicologist, sllgges ted that
MSG mi ght induce brain lesions, particularly in infants (Olney and Sharpe,
1969}. In response to these concen1s, MSG was removed from baby foods in
the 1970s. The fina l conclusion (see Wal ke r and Lupien, 2000) is that MSG in
large d oses may be a p roblem for some sensitive individuals, but appa rently
it does not present a serio LLS proble m for the p opulation . For those
w ho are sensitive, h m·vever, MSG can p ose a serio us risk.
The Special Case of Fat

Like pro tei n, fa t is a very important nutrient. Also Hke prote in, fat molecules
are too large to stirnulateeither taste or olfaction but are broken into theircon-
stihlent parts by digestion in the gut. Fat molecul es are ma de up of fatty acid s
attac hed to a s upport s tructure. 'A'h ole fat m olecules s timulate the trigeminal
nerve in the m outh, evoking tacti le sensations like oily, viscous, cream y, and so
on; but some fu t molecules m ay be partially digested while still in the m outh,
thus releasing fatty acid s. Neurobiologis t Tim G ilbertson1 s d iscovery of fa tty
acid receptors o n the to ng ues of rats led to the idea tha t stim ulation o f these
fatty add receptors might play a ha rdwired role in our love of fat.Just as wi th
glut.a.mate, however, nature uses a more general meth od to ens ure that we love
fa t-containing foods. Anthony Sclafani (n learning theorist who is an expert on
conditioned food preferences)showed that fat in the gu t produces cond itioned
p references for the sensory properties o f the food con taining the fat. O n ce
we understand the role of con di ti oning in food preferences, we s h ould have
a heal thy s kepticism about the value of so-called d.iet food s. Mimicking the
$ensory properties of n ormal foods but reducin g the caloric content disrnpts:
n orma l regu lator y m echan isms, as noted by Davidson and Swithe.rs (see the
''Sweet" section earlier in thJ s chapter).
Is All Olfactory Affect Learned?

The eviden ce th1.1 t a ll o lfactory affect is lea rned is still p rovis ional. The number
o f od.orants tested on young child ren is limited by the difficulty of d oing these
expe rimen ts. Could the re be a fe,..v o d ora.nts tha t, like taste, have ha rch..,·ired
affect? One su ggestion makes cons iderable evolution ary sense. Certain od or-
a.nts are de.rived from important nutrients in fniit.s and vegetables and are thus
cues to those nutrien ts. Hardwired liking for these odorants would lead to a
h ealthier diet (Goff and Klee, 2006).
Another reason to consider the possibility o f ha rd wirin g of the pleasure
associated with some o d orants is the eviden ce fo r hard wired olfactory aver-
sions in some species. Once we consi de r the possibility that some o dorants
are innately dis liked, it is reasonable to reevaluate the p ossibility that some
are innately liked. The problem with innate ave rsion to od ors is tha t s tudies
claiming this mus t d o a crucia l control. Since pain and its mild er cousin,
irritation, a re ilmately disliked, any od orant claimed to be inna tely dis liked
mus t be s h own to lack irritMion sensations . For example, cons ider s niffing
am monla. Amm onia h as a n odor, but it also bums the nose. TI1e burn is medi-
ated by the trigerninal ne rve. \'Vith humans there is an easy way to identify an
odorant that also s timulates the trigeminal nerve throug h the localization of
sens ntion . lf a pure odorant is ins tro duced to just one n ostril but bo th nostrils
are stimulated, the odor canno t be localized to th e appropriate n ostril. If the
odorant contains trigeminal stin1ulan ts, the sensatl on c.an be localized . 11-1ere
are n o final an s\vers yeL Stay tuned....

TASTE 489
The Nature of Taste Qualities

Although we take for granted that sweet, salty, sour, bitter (and whatever else labeled lines A theory of taste ced-
ma kes its way onto the list) are ta stequaJitiesr historically that was not always ing In which each taste nerve lber
the case. Herinan:n von Helmholtz, a physician/ physicist of n ote in the nine- carries a particular taste quality. For
example, the quality evoked from a
teenth century (see 01apter 1), tau ght that modality a nd quality .should be sucrose-best fiber Is ffi-veet 1 that frcrn
d istinguished; if h ¥o sensatio ns are so different th a t there are no transitions an NaCl-best flber Is salty, and so on.
be tween them, the n they should be con side red separate modalities. Two s ht-
d ents of Helmh oltz (\Vill1elm Ww1dt and Frithiof H olmgren) in tum produced
students (Friedrich Kiesow and H}almar Ohnvall) w h o saw Helmholtz's v iew
differently. Kiesow, a s tudent in Wundfs lab in Leipzig (con side red to be the
first labora tory of experimental psychology), saw th e differen t taste sensa-
tions as analogous to colors and so concluded that different taste sensations
are differen t qualities wi thin the tas te sense. On the other hand, O hnvall, a
student of H olmg re n, saw taste qualities as independent of o ne another and
so as sepa ra te modalities. The d etails of their researd1 duels (see Bartoshuk1
1978) ach1ally s upported but Kiesow obv ious ly won the battle. O ne
important fuctorwas tha t Wundt's la b hos ted a variety of prominen t psycho lo-
gists, among the m ni.any Americans. The proximity o f Kiemw to Americans
w ho wrote influentia l texts may have contribu ted to tha t victory.
Reminiscent of this old deba te is a 1nodem d eba te over how taste qmUity is
cod ed. A major source of his torical controversy in the taste literatu re revolved
around w he ther tastes are coded mainly via labeled lines, in w hich ead1 taste
neuron would W1ambiguously signal the presence of a certa in basic t..'\Ste, or via
patterns of activity across man y different taste n euro ns. We've seen examples
of both types of codin g in other senses. For example, color vision and olfac tio n
use pa tte rn coding . A single type o f cone cannot te ll us the wavelength of a
Bght ray, but the p a ttern of activity across our three cone types can give us
this infor mation. Hearing, on the othe r hand, u ses ri med"lanism more akin to
the labeled -line 3pproach: certain neu rons a lways respond best to 5000-hertz
(H z) ton es, others always l'esp ond bes t to 5100-Hz tones, '1nd so on . Which
scheme is used in the gus ta tory system?
FURlHER DISCUSSION of the coding of =lora, sourds, and odors can

be found in Oiapters 5, Q, and 14, respectivety.
Given \·"·ha t we' ve al rea d y lea rned about the functi ons of the four basic
tastes, it is easy to cons truct i.' ln evolution.."lry arg ument for labeled-line cod-
ing. Recall tha t in olfaction, which uses pattern codin g, mixhues o f different
compounds very often produce a new smell sensatio n; the components a umot
be recogn ized. l)uch a cOdiltg wo<tld be disastrnus for the purp ose o f
the taste system. For example, poisonous p la n ts contain componen ts with a
variety of tastes . If bitterness were to synthesize with these other tas tes, we
would not be able to parse it ou t and thus avoid the pojson. The ftmction s of
the four ras tes are well served by their independence from each other. A final
argument involves taste mixtures. Studies have shO\vn that we are, in fact, very
good a t analyzing taste nUxtures . For exa mple, in tonic '''a ter, w hid1 contains
a combin ation of quinine a nd we have no difficulty identifying its hvo
cornponents-bitter and swee t. In a pattern cod ing theory, bitter and swee t
would each be co ded by a differen t pattern; combining those patterns would
produce a new pattern and a new taste.
The historical controversy arose because initial research see med to indicate
that m ost neurons coming from tas te buds responded to m ore than one of the
four basic tas tes. How could s uch a system code sweet, so ur, sa lt, and bitter

490 CHAPTER 15
w ith o ut confus io n? C arl Pfaffmann initially conclude d that tas te qualities

mus t be coded by a pattern across taste fibers, but he ch anged his nUnd w hen
additiona l research showed that much o f the earlier confusion was n oise
(Pfuffmann, 1974a, 1974b).
Taste Adaptation and Cross-Adaptation

As we' ve seen throughou t this book, a ll sensory syste ms sh ow adapta-
tion effects, in which cons ta nt application of a certain stimulus temporar-
ily weakens s ubseq uent perception of tha t stimulus. In taste, o tu constant
adaptation to the salt in saliva affects o ur ability t o taste s alt; in addition,
ad a pta ti on to certain compo n ents in one food ca n chan ge the perception of
a second food. (See Web Essay 15.2: Water Tastes.) You've experien ced
cross-adaptation you rself if yodve ever n ot iced that a beverage like lem-
tastes too sour after you eat a sweet d essert. The sugar in the d essert
adapts the sweet receptors so tha t the lemonade tastes less sweet
and more sour than normal.
Pleasure and Retronasal versus Orthonasal O/faction

There is s till mud1 that is not understood about the links beh¥een retrona-
sal and orth.o nasal (th roug h the nostrils) olfaction, including the pleas ure or
d isplensure associated with these sensations. We know tha t we le.a m to like
or disli ke smells, but d o we learn these preferences separately fo r retro- and
orth.onasal olfaction? Bartoshuk recounts getting carsick on a childhood va-
cation '"'hile eating chocolate-co\·ered cherries. She now no t
only avoi ds the cherries, but finds cherry-scented soaps disgusting as well.
The distinction between retron asa l and orthonasal o lfaction has only emerged
as important in recent years , \Ve h;:ive much to learn in thi s area.
Chili Peppers
Th e pleasu re that some peop le experience fro m chili peppers deserves special
attention . \Ve are not born liking chili peppers. Rozin studied the acquisition
o f d1ili pepper preference in Mexico and fo und that the process depends on
social influences. Chili is gmduaJly added to the diet of ymmg children begin-
ning at abo ut age 3, and the dUld.ren observe their family members enj oying it.
By age 5 o r 61 child1-en voluntarily add chili to their own food. At some p oint
the chili is liked for its oV\o-·n sake.
A variety of argu me nts based on presumed health benefi ts have been
b1troduced to accmmt for our love o f chili peppers. For exa1nple, some have
argued that chilis kill microorganisms in food, thus acting as a prese rvative.
Oth ers have a rgued that d 1ilis contain vitamins A a nd C, vdtlch give them
adaptive va lue (in other words, the pain of the chili serves as a cue for the
p resence of the vitamins). 1l1e p leasure that some people experience from dtilis
has also been linked to the idea that the resulting bun1 leads to the re lease of
end orphins , the brain' s internal painkillers.
One of the most interesting features of the liking for the bum of d1i1i peppers
is its near tota l res triction to humans. Rozin has documented a few cases on
record of anima ls showing a liking fo r chilis, but in a ll cases these \'\·· ere pets
fed chili pepper by their human companions. \ Vh en Rozin tried to produce
liking for chilis in rats, h e failed. But on e of Rozin's s tudents, Be1mett Galef,
w ho is famous for the s tudy of socia l interactions a mong rats, was finally
able to get rats to like a diet season ed 'vith a ntlld leve l of rnyenn e pepper, by
expo.sing the rats to a "demonstrator" rnrthat hddjust aten the diet seems rt
that growing to like chili peppers is a social p henomenon for rats as well

TASTE 491
8230336 Amma Appa
FIGURE 15. 14 Do thQSQ irnagQs inspire foar or d'-llight in your mcuths?
111.e bum that we experience from dilli p eppers is highly variable across
individ u a ls (Figure 15.14). The variability has h.1,·o sou rces. First, as noted
earli er, individuals '\>1.-i th the largest number of hmgifo nn papillae (supertasters)
have the most fibers rnediating pain, and thus they perceive the most intense
oral bum from chills. Second, capsakin, the che mical that prod uces the bun1 in
chilis, desensi tizes p<iin recep tors. This m ea ns th a t individuals w ho consume
d1ilis quite often (once every 48 hours is sufficient) are chronically desensi tized.
C hili peppers produce considera bly less btrrn to those w ho are desensitized .
Desensiti zation ca n com e to your rescue if you accidenta lly order a m eal
tha t proves to be overspiced. for yom pa late. After the first m outhful, wa it
until the bum has subsided . The mis take many diners make is to keep trying
to eat. As long as the capsaicin continues to be applied, desens itization d oes
not occur. Desen sitization ocnu s during the decline o f the burn (B. G. Green ,
1993). Once the initial burn h as faded, the res t of the mea l can be consumed
with relative comfort.
Capsaicin desensitization has importa nt clinical value. l11e ancient Mayans
u sed a concoction made of chilis to treat the pain of mouth sores. In the 1990s,
Wolffe Nad oolman, then a medical s tudent a t Yale working in Bartoshuk's
laboratory, cre..1 ted a s imilar remedy by adding cayenne pepper to a recipe for
taffy. Can cer patients often develop painful mouth sores fro m che motherapy
and radiati on therapy, a nd if these patients suck on the capsaicin candies,
c.ipsaicin is brought into contact with the pain receptors s timulated by the
sores. The pain receptors are then desens itized a nd th.e pain is dramatically
reduced (Berger et al ., 1995). Although capsaicin can be used to re duce pain a t
any body site, the s kin is a potent barrier that prevents capsaidn from contacting

492 CHAPTER 15
pain receptors. capsaicin remedies for disorders like arthritis are rarely
very satisfactory. In the mouth, the mucous m ernbrane pem1its capsaicin to
easily contact p ain receptors, so there, desensitization is fast and powerful
Summary
1. Flavo r is produced by the combination of taste and retronasal olfaction
Refer to the (olfactory sensations produced when odorants in the mo uth are forced
Sensation and Perception up behind the palate into the nose). Flavor sensations are localized to the
Gompanim website mouth, even though the retron.asal olfactory sensations come from the olfac-
Sltes.Slnauer.comlwotte4e tory receptors high in the nasal cavity.
for essays, study 2. Taste buds are globular dusters of cells (like the segments in an orange).
The tips of some of the cells (m.icrovilli) contain sites tha t interad with
queslons. and otrer study aids .
taste molecules . TI1ose sites fall into h"o groups: ion channels that medi-
ate responses to salts and acids, and G protein-coupled receptors that bind
to swee t and bitter compounds as well as compounds d escribed as having
"umam.i ·• taste.
3. The tongue has a bumpy appearance because of structures cal led papillae.
Filiform papillae (the most numerou s) have no tas te buds. Taste buds are
found in the fungiiorm papillae (front of the tongue), foliate papillae (rear
edges oi the tongu e), and circumvallate papillae (rear renter of the ton gue),
as well as on the roof of the mouth.
4. Taste projects ipsilaterally from the tongue to the medulla, thalamus, and
cortex. It projects firs t to the insula in the and from there to the orbi-
tofrontal cortex, an area where taste ca n be {ntegrafed with other sensory
input (e.g., retronasal olfaction).
5. Taste and olfoction play very different roles in the perception of foods and
beverages. Taste is the true n utritional sense; taste receptors are tuned to
molecules that func tion as important nutrients. Bitter taste is a poison detec-
tion system. Sweet taste enables us to feSJX>nd to the sugars that are biologi-
cally useful to us: s ucrose, glucose, and fructose. S...<1.lty taste enables us to
identify sodium, a mineral crucial to survival because o f its role in nerve
conduction and muscle function . Sour taste permits u s to avoid adds in
concentrations that might injure
6. Urnami, the tas te p rod uced by monosod ium glut-ama te, has been s uggested
as a fifth basic taste that detects protein. However, umami lacks on e of the
most important properties of a basic tas te: hard\vired affect. Some individ u-
als like umami, but others do not. Taste receptors are not only in the mouth
but also in the gut. Digestion breaks dm\.n proteins into their conshtuent
amino acids, and the glutamate released stimulates gut gluta mate receptors,
leading to conditioned. preferences for the sensory propertie.51 of the foods
containing protein.
7. The importance of taste to survival requires tha t we be able to recognize
each taste quality independently, even \..•hen it is present in a mixture. By
coding tas te quality with labeled lines in much the same way that frequen-
cies are coded in hearing. na.hue has ensured that we have this important
capability. These labeled lines are noisy. For example, adds are able to stiln-
ulate fibers medii.lting saltiness, as well as those medii.lting sourness. Thus,
acids tend to taste both salty and sour.

TASTE 493
8. Foods d o no t tas te the sam e to everyone. The Human Geno me Project

revealed tha t we carry about 25 genes for bitter taste. ll\e mos t studied
bitter receptor responds to PROP -and sho\\'S allelic \·aria tion in human s,
leading to the d esignations "PROP nontaster'' for those who taste the leas t
bittern ess and " PRO P tas ter'' for those \vho taste the most. In addition,
humans vary in the number of fungifom1 papillae (and thus taste buds)
they possess. Those with the mos t tas te buds are called su pertasters and live
in a " neon" taste v.rorld; those "" ith the fe\-.•est taste buds live in a "pastel"
ta5te world. Psychologists discovered these differences by testing peop le's
ability to match sensory intens ities of stimuli from different modal ities. For
example/ the bittem e.ss of black coffee ma tches the pain of a mild h eadache
to nontasters but resembles a severe headach e to s upertasters. 1l1e ''"ay
foods taste affects palatabili ty,. wh ich in tum affec ts die t. Poor die t contrib-
utes to diseases li ke cancer and cardiovascular d isease.
9. Fo r taste, t.mlike olfaction, liking and d isliking are hard wired; for example,
babies are born liking sweet and salty and disliki ng bitter. When we become
defic ient in salt o r sucrose, liking for sal ty and sweet tastes,. respec tively,
increases. Junk foods a re cons tructed to appeal to these preferences. Li.k-
ing the burn of chili peppers, o n the o ther h and, is acquired and, w ith the
exception o f some pets,. is essentially limited to humans. Taste buds are s ur-
ro unded by pain fibers; thus supertasters perceive grea ter bum fro m chil is
than do no n tasters. In additio n, fungi fo rm papillae, structu res that house
taste buds, are innervated by to uch fibersi thus supertasters perceive greater
touch sensations fro m fa ts (e.g., creamy, viscous, thick) in foods.
8230336 Amma Appa

lq
8230336 Am111a Appa

Glossary
Ntunbei·s in brackets refo1· to the chapter(s) '"''het'l2 the term is introduced.
A In the wor1d (e.g., the sides of two aerial perspectfve or haze A

A-beta fiber A wide-diameter. lrldependent cb]ects llnlng up per- dspth cue based on the implicit under-
myellnated sensory nerve fiber that fectlYJ. (4] that light Is scattered by the
transmits slgnals from mechanical accommodation The process by atmosphere. Mere light Is scattered
stimulation. (1 3] whbh the eye changes Its focus wt1en we loci< thrcugh more atmo-
whbh the lens gets fatter as gaze Is sphere. Thus. more distant cbjects are
A- delta fiber An Intermediate- sub;oot to more scatter and appear
sized, myellnate::t sensory nerve flbef directed toward nearer objects). (2. 6]
fainter, bluer. and less distinct. (6]
that transmits pain and temperature achromatopsla An lnablltyto per-
signals. (1 3] ceive coors that Is caused by damage afferent fiber A neuroo that carries
to the central nervous system. (5] s€f"lsay Information to the central ner-
abducens (VI) nerves The sixth vous systan. Compare efffrent fiber.
pair of cranBJ net'Ves, whicl1 Innervate acoustic reflex A renex that pro- (9, 121
the lateral rectus muse"' of the eye- tects the ear from Intense sounds, via
balls. (1 J contraction of the stapedlus and ten - afferent signals Information nOW'-
sor tympani muscles. (9] lng Inward to the central nervous
absolute metrical deplh cue A sys.tern from senSCfs In the periphery.
depth cue that pr<Mdes quantifiable acoustic starUe reflex The very Passt;e sensln;i would rely exclust;ely
lnfe<matlon about distance in the third rapid motor response to a sudden on such sensory Inflow, providing a
dimension (e.g .. his nose stkJks out 4 soum. Very few neuroos are lrwolVed traditional view of sensation. See also
centimeters In tront of his face). (6] In the basic startle retlex. which can Bfferenl fiber. (1 2]
absolute pitch A rare ability also be affected by emotional state.
(1 0] age-related macular degenera-
whereby acme people are able to very tion (AMO) A disease associated
accurately name or produce notes acttve sensing Sensing that with agln;i that affects the macula.
without ccmparlscn to other notes. Includes self-generated probln;i of AMD gradually destroys shalP central
(11] the erMronment. Besides a.ir sense vision. making It difficult to read, drwe,
of equlllbnum. other active human and rec(>iJntze faces. There are two
absolute threshold The minimum
amount of stlmulat bn necessary ta senses lnckde vision and touch. An- forms of AMO: wet and dry. [2]
mal active sensing Includes the use of
a person to detect a stimulus 50% of echoes by whales and bats, the use agnosla A failure to recognize
thet&ne. (1] objects In spite of the ability to see
of sig)als by SOfne flsh. and them. Agnosla Is (\plcally due to brain
absorb Tot e too use of whlsl<Bfs/antennae by flsh.
such as light. noise. or energy-and damage. (4. 5]
Insects, and nocturnal rodents. (121
not transmit It at all. (2] aklnetopsla A rare neuropsycho-
acuity The smallest spatial detail
acceteratton A charge n vetoc- that cao be resolved at 100% cm- loglcal disorder in which the affected
lty. Mathematk:alty, acceleration Is the trast (3] lndlvldual has no perception of motion.
derlvatwe of velodty. In WOfcis. linear (SJ
acceleration lndlcatoo a change In lin- adaptation A reduction In respcrnie
caused by Pl1e< or cmtl nulng stlmuta- amacrine cell A retinal cell found
ear veloclty; an:;;iular aa:eleratlon Indi- In the Inner synapt b layer that makes
cates a change In angular velocity. (1 2] ttn. (3] synaptic ccntacts with bipolar cells.
accessory olfactory bulb adapting stimulus A stim ulus ganglion cells. and other amacnne
(AOB) A smaller neural structure wh:::ee removal produces a ch:tnJe In cells. (2]
located behind the main dfactory bulb visual perception or sensitivity. (5]
ambiguous figure A visual stimu-
that recetJes Input from the vomerona- additive color mixture A mixture lus that gives rise to two or more Inter-
sal organ. (141 of lights. If light A and light B are ooth pretations of Its k:lentlty or structure.
accidental vtewpoint A viewing renected from a surface to the eye, In 141
poeltlon that produces some regularity the perception of color the effects of
t hose two lights add together. (5] ambtyopia A devEJcpmental disor-
In the vlsU9.l Image that Is not present der characterized by reduced spat lal

496 GLOSSARY
vislcn In an otherwise healthy eye, even aperture problem The fact that acoustlo signal Into separate auj ltory
with proper carectlon for refractive when a rnovng object is viewed events for which each stream Is heard
errcr. Also known as l!IZY '*·
[3] throligh an aperture (or a receptive as a separate event. I1OJ
amplitude or Intensity The mag- field). the direction of motion of a local autonomic nervous system The
nitude of displacement Qncreese or feature cr part of the cll]ect may be part of the nervous system that Inner-
decrease) of a sound pressure W"<£Je. ambiguous. [8] vates heart. digestive system,
Aniplltude Is as loudness. [91 aperture An opening that allows only and so on, and that Is responsible for
a partial ,;ew of an cllject. [8] regulating many lnvoluntaiy actions. [1 2]
amplitude The size Oncrease or
decrease) of a head movement (e.g., apparent motion The illuocry azimuth The angle of a sound
angular veloclty, linear acceleratlon, t11t). Impression of smooth motion resulting source on the horizontal plane rela-
[1 2] from the rapid alternatbn o f cllJects that tive to a point In the center of the head
ampulla An expanslcn o f each semi- appear in different locations In rapid between the ears. Azimuth Is mea-
c ircular-canal d uct that lnclud"" that succe8"on. [8] sured In degre"". "1th 0 degrees bl>ng
canal's rupula, crista, and hair oolls, aqueous humor The watery fluid In straight ahead. Tile angle increases
wl1ere transdu::tlon occurs. [12] the anterior chamber of the eye. [2] clockvAse toward the right, with 180
degrees being directly behind. [1OJ
amygdala-hlppocampal complex aromatherapy The manipulation of
The conjoined regions of the arnygdala odors to lnnuence, mood, pertormance, B
and hlppocampus, which are key struc- and well-being, as well es the physi- balance system The smscry sys-
tures '1 tl1e limbic system. This com- ological ccrrelates of emotion such as tems. neural pro::::esoos. and muscles
plex Is c rltlcally In the unique heart rate, blood pressure, and sleep. that contnbute to postural oont rol. Spe-
emotlcnal and associative properties o f [14] cmc components Inc lude the vestibular
olfactory cognition. [14] artlculatlon The act or manner of organs, kinesthesis. vestlbulo-splnal
analgesia Decreasing pain sensation produc ing a speech sound using the pathways, skeletal bones. and postural
durtng conscious experience. [1 3] vocal tract. [11] ccntrol musdes. E?ecause of Its crucBJ
ocntributlcns to balance, some even
anamorphosls or anamorphic astigmatism A visual defect caused informally refer to the vestibular systerr
projecUon Use of the rules of Unear by the unequal curving of one or more as the system" and the ves·
perspective to create a two-di menslonal of the refractive surfaces of the "fe, tlbular organs as the "balance organs."
image so distorted that it looks correct usually the cornea. [2] But the 1::0.lance system is much more
only when viewed frcrn a special angle attack The part of a scund during than jllSt the vestltlular system. and the
er with a mirror that counters the distor- v.nlch amplitude lncreas"" [10] vest lbular system contributes to more
tion. [6] than just balanoe. [12]
attenUon deficit hyperacUvlly
angular acceleration The rate of balance The neural processes
disorder (ADHDJ a quite common
change of angular velocity. Mathemati- c hild hood disorder that can continue of postural control by whloh weight
cally. the Integral of angular acoeeratlon into adulthood. Symptoms lncludedlffl- Is evenly distributed. €flabll ng us to
Is angular velocity. and the Integral of c ulty focusing attention as well as prob- remain upright and stable. [1 2]
angular veloclty Is angular displace- lems controlling beha•Aor [7]
ment. Angular aoceleratlcn, angular basal cell One of tlie three types of
attention Mr/ of the very large set oells In the olfactory eplth<>lum. Basal
of selec11ve processes In tl1e brain. To cells are tl1e precursor oells to olfactory
motion. [12] deal with the lmposslblllr1 of handling sen oory neurons. [14)
al Inputs at onoe, the nervous system basic color terms Color words
angular moUon RotallonaJ motlcn
has evolved mechanisms that are able t11at are monolexemlc (single wcrds
like the rotatlcn of a spinning top or
to bias prooesslng to a subset of things, "blue• not. "sky blue'), used with high
swl'lgng saloon doors that rotate back places. Ideas, or moments In time. 171 frequency, and have meanlrrJS that
and forth. [12]
attenUonal blln k The tendency not are agreed upon by speakers of a lan-
anlsometropla A condition In which to pa-cetve or respond to the seocnd guage. [5]
the have dlffennt refractive of two different target stlmull amid a
errors (e.g., one "fe Is farsighted and basic taste Arf'f of the four taste
rapd stream of distracting stimuli If the qualities that are geneially agreed to
tl1e other noQ. [3] observer has responded to the first d...cnbe human taste experience:
anomia An inability to name objects target stimulus "1thln 200-{)00 mll- sweet . salty, sour. bitter. [1 5]
In spite of tl1e ablllty to see and recog- llsecords before the seccnd stin-.ulus Is
nize them (as shown by usage). An:::mla prooented. [7] basilar membrane A plate of fibers
that fcrms the base of t11e cochlear
Is typically due to brain damage. [5] audibility threshold The k:west partition and separates the middle and
anosmla The total Inability to smell, sound pressure level that can be rellatJly tympanic canals In tl1e oochlea. [9]
most often resultlng from sinus Illness or detected at a g"en frequency. [9]
l1ead trauma [14] Bayesian approach A We.ff of for·
auditory nerve A oo lectlon of neu- mallzlng the Idea that our perception Is
anterior cingulate cortex (ACC) rons that Information fran hair a combination of the current stlmulus
A region of the brain associated "1th cells In the oochlea to (afferent) and and our kno1-.edge about the condi-
the perceived unpleasantness of a pain from (efferert) the brain stem. [9] tions of the world-what Is and Is not
sensation. [13] auditory stream segregation The II kely to occur. Tl1e Bayesian approach
perceptual organization of a complex Is stated rnathanatlcaily as Bayes·

GLOSSARY 497
theorem-P(AIOJ = P\<\) xP(O jA)/P(O)- blood oxygen level-<lependent from the Greel< for "large" refe<rlrr;i to
whlch enables us to calculate the prob- (BOLD) signal The ratlo of oxygen- the size of the cells. [5]
ability ip) that the world Is In a particular ated ta daoxygenated hemoglobin that clrcumvallat e papillae Circular
state !Ai given a particular observat k::<1 permits the localization of brain neurons structures that form an Inverted Von
(0). [4. 6] that are most lnvolVed In a task. [1] the rear of the tongue (three to rrve on
belt area A region of = tex. directly body Image The Impression of our each side, with the largest In the cen-
adjacent to the prtmary auditory cortex bodies In spaoe. [1 3] ter). Circumvallate papillae are rnound-
(t>.1), w ith Inputs from A1, w here roo- c llke structures. each surrounded by a
rcns respond to mere complex charac- trenoh (like a moat). These papll lae are
teristics of sourds. [9] C fiber A narrow-diameter, unmyeUn- much larger than runglform papillae.
ated soosc.ry nerve flber that transmits (1 5]
blnaral rivalry Competition between pain and temperature signals. [1 3]
the two nootrlls fcr odor perneptlcn. closure In reference to p€fceptfoo,
When a different soent Is presented to C tactile (Cl) afferent A narrow- closure is the name of a Gestalt prin-
eac/1 nostril slrnultanecus/y, we percetve diameter, unmyellnated sensoty nave ciple that hods that a closed = itcur Is
each scent to be alternating back and fiber that transmits signals from pleas- preferred to an open contour. [4 J
forth w ith the other, and not a blend of ant touch. (1 3]
coarticulatlon The phenomenon In
the two scents. [14] cataract An opacity of t11e crystalline speech whereby attributes of su:ces-
binding problem The c halenge of lens. (2] slve spaech units overlap In articulatory
tying different attnbutes of visual stimuli categorical perception For speech or acoustic patterns. (11]
(e.g. , cdor, orientation. motion), which as well as other oomple.x sourds an::l cochlea A spiral structure of the
are handled by brain circuits, to Images, the phenomenon by w l1lch the inner ear containing the crgan of Corti .
the apprq::rlate object so that we per- dlsc:rlrnlnatlon of Items Is no better than (9]
celVe a unmed object (ag., red, vertical, the ability to label Items. (11j
m<M ng rlghQ. [7] cochlear nucleus The first brain
change blindness The failure to stem nuclaus at w hich afferent auditory
binocular depth cue A depth cue not ce a cliange between two scenes. If nerve fi bers synapse. (9]
that relies on lnfamatlon from both the gist. or meaning, of the s:ene Is not
eyes. Stereopsls Is the prlmaJY example altered, quite large changes can pass cochlear partition The combined
In humans, but convergence and the unnotloed. [7] basilar membrane, tectorlaJ membrane,
ablllty of two eyes to see more of an and c<gan of Corti, which are together
c haracteristic frequency (CF) The responsible fcr the transduction of
object than one eye sees are also bin-
frequency to which a particular auditory sound waves Into neural signals. (9]
ocu lar depth cues. (6]
nerva ftber Is moot sensitive. [9]
binocular disparity The dlffereroes cognitive habituation The psy-
chemoslgnal Any of varb.Js chological process b y "'11ch, after
between the two retinal Images of the
chemicals em ltted by humans that loog-term exposure to an c:dcr. one no
same scene. Dlspanty Is the basis for
are detected by the system lcnger has the ablllty to detect that odor
ster0q)sls, a vivid perception of the
and that may have scme effect on the or has very diminished detection ability.
three-dimensionality of the v.ald t110t Is
moo::!, betlavlor, horrncnal status. and/ (14]
not avallabla >Mth monocular vision. [6]
or sexual arousal of othE< l1umans. (14]
binocular rivalry The cornpetl- cold fiber A sensory nerve fiber that
tlcn between the two eye.s fa contrd chord A comblnatlcn of three or fires when skin temperature decreases.
more musical notes with different (1 3]
of visual pe<ceptlcn, which Is evident
played simultaneously. (11]
when completely different stimuli are color assimilation A cdcr percep -
presented to the two eyas. (6] chorda tympani The branch of tion effect In which two colors
cranial ne<ve VI I (the facial nerve) th3l Into each other. each taklrr;i on some of
binocular summation The combi-
carries taste Information frcm the ante- the chromatic quality of the other. [5]
nation (or •summation'] of signals from
nor. mob Ila torr;iue 11he part that can
each eye In ways that make perfor- color constancy The tendoocy of a
be stuck out). The chorda tympani exits
mance on rnany tasks better both surface to appear the same oda ur-.:ier
the tongue with the lingual branch of
eyes than with either eye alone. [6] a wide range of illumlnants. [5]
the trlgeminal nerve (eran k1l nave V)
binocular With two eyes. (6] and then passes through the middle ear color contrast A color perception
biological motion The pattern of en Its way to the brain. (1 5] effect In which the cdor of c:ne re:;Jlcn
movement of llvlrr;i bengs (humans ard chromophore The llght-catchlrr;i the opponent color ln a nelgh-
animals). (6] part of the visual pigments of the retina< borlrg reg ion. (qJ
bipolar cell A retinal cell that syn- 121 color space The tlY -dimensional
apses with either rods or cones (not cilium Any of the hairlike protrusions space. established booause ccior per-
botry and wltl1 hortzmtal oells. and then ception Is based on the outputs of tt'ree
en t11e dendrites of olfactory eansory
passes the signals on to ganglion cells. oone typos, that describes the set of all
neurons. The receptor sites for cx1crant
[2] molecules are on the cllla, which are the cciors. [5]
bitter One of tlie four ba$c tastes; ftrst structures lnvo.,ed In olfactory sig- color. anomalous A bettet' term for
the taste quality, generally considered nal transdLctlon. (14] what Is usually called •coor-bllnd." Most
unpleasant, produced by substances circadian Referring to cells In the 'color-blind" lrdMduals can still make
like quinine cr caffeine. 11 5] magnocellular layers of the lateral gerlc- dlscrlrnlMtt:ns based on waveloogth.
ulate nucleus of the thalamus. Magno

498 GLOSSARY
111000 dlscnmlnatlons are from helpful in sd'ling the corresponder>0e crlbriform plate A bcny struoture
the norm-that Is. anomalous . 151 problem. 161 riddled with tiny holes, at t11e levet of tile
column A vertbal arrangement of ccntralateral Referring to the oppo- eyebrows, that separates the """' trom
neurons. Neurons w1thln a single col- site sl:le of the body (or brain). [3] the brain. The axms from the olfactory
umn tend to have similar receptive fields senOCfy neurons pass tt1rough the ti ny
contraleslonal field The vlsual field hoes of the crlbrlform plate to enter U1e
and similar orientation preferences. (3) on 1110 side opposite a brain loo/on. For braln. 114]
comparator An area of the visual example, points to the left of fixation are
system that reoeves one copy of the contraleslonal to damage In the rlgl1t crlsta AJ-'f of the specialized detec-
command Issued by tile motcr system hemisphere of the brain. 17] tors of angular motion located In each
when the eyes move other copy semicircular canaJ In a swe llng caJled
contrast sensitivity !Unction (CSFJ the ampulla. 112]
goes to the eye muscles). ll1e corn- A funct bl describing how the sensltlvtty
parata compares tile Image motion to contrast (denned as the recprocal of criterion In slgnaJ detection theory,
slgnaJ with the eye motion signal and the contrast threshokj) depends on the an Internal threshod that Is set by the
can compensate for the image changes observer. If the lnternaJ response Is
spatial trequency (size) ol the stimulus.
caused by the eye movement. 181 above crlterlcn. the obsefVer gives one
13) response (e.g .. 'yes. I hear that}. Below
complex cell A OCftlcal neuron contrast threst1old The srnaJlest
whose receptive leld does not have crlt€fion, the observer gives arother
amount of contrast required to detect a response (e.g .. ·no. I hoor nothlng1. 11)
delned excltatcry and Inhibitory pattern. 131
regbns. 131 critical bandwidth ll1e range of fre-
contrast The dlffererce ln lumlnance quencies conveyed ''"thin a channel In
computed tomography (CT) An between an object and the bacl<(lround,
Imaging technology that usea X-rays the audltcry system. ]9]
or between llghter and darker parts of
to create Images of sllces through crlllcal period A phase In the life
the seme object. 12. 31
volumes of mater1aJ (e.g .. t11e human span during which abnorrnaJ early
bod)'). 111 convergence The ablllty of the two experience can alter normal neuronal
eyes to tum Inward, onen used In crder development. Crttlcal periods are pro-
conductive hearing loss Hear- to place the two Images of a feature In
ing bss caused by problems with the posed fcr the deveopment of l:Jnooular
the world on corresponding locations vision and development of a first human
bcnes of the middle ear. 191 In ths two retinal Images (typlcaJly on 6]
cone A photoreceptor specialized for the fovea of eacl1 eye). Converger>0e
daylight vision, fine >Asual aoulty, and reduoes the disparity of that feature to cross...adaptatlon The redt..dla"l
color. 121 zem tor rearly zero). [6] n detection of one odorant following
exposure to another odcrant. Cross-
cone monochromat An individual cornea The transparent 'window" adaptation Is presumed to occur
with only one cone type. Cone tnono- Into the eyebaJI. [2] because the components of 1110 odo!S
chromats are truly color-blind. [5] correspondence problem 1. In (Of the odora'1ts) In question share one
cone of confusion A region of posi- blnooular >Aslon. the prot>em of figur- or more olfactory receptcrs for their
tions In space where all sounds pro- ing out which bit of the Image In ths left transduction. but the order In which
duoa tile seme time and lsvel Qntenslty) eye should be matched with which bit odorants are presentOO also plays a
dlfferenoes (ITDs and 11..Ds). 11 OJ In the right eye. ll1e problem Is par11cu- role. [14]
cone-opponent cell A cell type- larly vexing wl10n t11e Images consist cross-modality matching The
foun:J In the retina, lateral genlculate of thousands of similar features, like ability to match the Intensities of sensa-
nucleus, and >Asual cortex-that. In dots In rardcm dot stereograms. 2. In tions that come frcm different sens.cry
effect. subtracts one type of cone Input motion detection. the problem faced by modaJltles. ll11s ability allows insight
frcrn another. 151 the motion detection system of knowing Into sensory dlff€fences. Fcr example,
v.tilct1 feature In frame 2 corresponds to a listener might adjust the brightness of
congenital prosopagnosia A form a partla.1lar feature In frame 1. [6, 8] a light until It matches the loudness of a
of 'face blindness" apparently present tone. 11 , 15]
frcm birth, as opposed to corresponding retinal points Two
prosopagnosla," which would typically images of an cbject In the crossed disparity ll1e sign of dls-
be the result of an Injury to the nervous workJ are saJd to faJI on ccrrespond - panty created by objects in tront of 1110
lng points If these points are the same plane of lxatlon ll1e term
system. 141
dlstanoe from 1110 fovea In both eyes. crossed Is used because Images of
conjunction search Search for a 1he two foveas are also corresponding objects bcated in tront of the horopter
target defined by tl1e preser>0e of two points. [6] appear to be displaced to the left 1n the
a more attributes (e.g .. a red, vertical right eye, and to the right In the len eye.
target among red horlzmtal and blue cortical magnification The amount
vertical dlstractors). 17) of cortical area (usually specified In mil- 161
limeters) devoted to a specmc region cue A stimulus tl1at might Indicate
continuity constraint In stereop- (e.g .. 1 degree) In the >Asual letd. 131 where (or what) a subsequent stimulus
sis, the ot:servation that. except at the will be. Cues can be valid (gMng correct
edges of objects. neighboring points In cranial nerves Twetve pairs of
nerves (one for each side of the badly) information). lnvalk:j or neu-
the world lie at similar distances from tral (uninformative). 171
the 'llewer. ll11s Is one of several con- that originate In the brain stern and
straints that have been proposed as reach sense organs and muscles cultural relativism In sensation and
throu;ih openings In the skull. 11 . 15] perception, the Idea that basic percep-
tual expenences (e.g., colcr perception)

GLOSSARY 499
may be determined In part by the cul- dllferent frequencies that arrive at each efferent commands lnforma·
tural environment. 151 ear from different locations In space tlon flowing outward from the central
cycle Fa a grating. a pair consisting (azimuth and eJevatlon). 1101 nervcus system to thl penphery. A
of one dark bar and one bright bar. 131 distractor In v1sual search1 arr;1 stim- ocmrnon example is motor cornrnands
ulus other than t11e target. 171 that regLdate muscle contraction. The
cycles per degree The number of ccpy of such motcr commands Is often
pairs of dark and bright bars (cycles of a divergence The ablllty of thl two callee! an "efferent copy." See alsoeffer·
grating) per degree of visual angle. 11 . 31 eyes to turn outward . often used In ent fiber. 11 21
Cyclopean Referring to stimuli th91 crder to place the two Images of a fea·
ture In the workJ on correspond ing loca- efferent fiber A neuron that carries
are defined by t>nocular dlspanty alma. Information frcm the central nervous
Named after the one-eyed Cyclops of tions In the two retinal Images
on tl1e fovea of each eye). DIVergence system to the periphery. Compare affer-
Homer's Ooyssey. 161 ent fiber. 19, 12)
reduoos the disparity of that feature to
cytochrome oxldase (CO) An zero (or nearly zero). 161 egocenter The center of a referen:::e
enzyme used to reveal tl1e regular array frame used to represent locat1ons rela·
of "CO bbbs. " which are spaced abcul dizziness A commonly usOO lay term
twe to the body. [1 3]
0.5 rnllllmeter apart In the pnrnary visual that nonspeclfioally Indicates any forrn
cortex. 13] of peroolved spatial disonentatlon. with electroencephalography (EEG)
cr without Instability. 11 2] At•ctJil<m ttiat uslrg ll]81JY ")ec-
D doctrine of specific nerve energies trodes on the scalp. measures electrloal
decay The part of a oound during A doctrine. fonnulated by Johannes aotlvty from populations of many neu-
which amplitude decreases (oflSeU. 11 0] MOiier. stating that the nature of a een- rone In the bfaln. 111
decibel (dB) A unit of measure for satk:fl depands on which sensay fibers emmetropla The oondltlm n whlcl1
the physical Intensity of scund. Deci- are stimulated. not on hoW fi bers are there Is no refractive error. because the
bels define the dltterenoo between two stimulated. 111 refractive power of the '1f9 Is perfectly
sounds as the ratio betlNeen two souM dorsal column-medial lemnlsc al matched to thl length of the eyeball. 121
pressures. Each 10:1 sound pressure (DCML) pathway The route from end stopping The process by which
ratio equals 20 dB. and a 100:1 ratio the spinal cord to the brain that carrlee a cell In the cortex first lrcreases Its
equals 40 dB. 191 sQnals fran skin, muscles. tendons, firing rate as the bar length Increases
dermis The Inner o f two major lay - and Joints. 11 3) to fill up Its receptive field. and then
ers of skin, consisting of nutritive and dorsal horn A region at the rear of decreases Its firing rate as the bar is
oonnectrve tissues, within which He the the spinal cord that reoowes Inputs from lengthened further. 131
mechanoreooptors. 11 31 receptors In the skin. 11 31 endogenous cue In directing atten-
deuteranope An Individual who suf- double dissociation The phenom· tion. an endogencc1s cue s loooted In
fers from color blindness that Is due to enrn in which one of tvvo functions.- (endo) or near the o..irrent locatlcn of
the absence of M-coneG. 151 such as hearing andsl;Jht, or first- and attention 171
dlchopUc Referring to the presen- second· order motion - can be dam· endogenous opiate A chernlcal
taticn of two different stimuli, cne to aged wtthoot 11arm to the ot11er. and released by the body that the
each eye. Dltterent from binocular vice versa. 14, BJ release or uptake of raurotransmltters
presentation, which oo..jd lrNdve both double-opponent cell A cell type. necessary to transm it pain sensatlrns
eyes k:oklng at a single stimulus. 161 found ln tl1e visual cortex, In which one to t11e brain. 11 31
diffuse bipolar cell A bipolar retinal region is excited by one cone type . endogenous spatial attention
cell processes are spread out to comblnatk:.n of cones. or color and A fcrm of top-dcmn (knowledge-dnven)
reoolve Input from multiple oones. 121 inhibited by the opponent cones or ocntrol o f spatial attentla1 In which
color (e.g .. R+.G-J. Another adjacent attention Is voluntarily directed toward
diopter (0) A unit of measur.,nent region would be Inhibited by thl fi rst the site where the observer anticipates
of the optic power of a lens. It Is equal Input and excited by the sooond a stimulus wtll occur. [1 3]
to tl1'l reciprocal of the focal length. In In this example. R-/G+). 151
meters. A 2-dlopter lens w111 bring paral· ensemble statistics The average
lei rays of light Into focus at Y, meter (50 dualism The Idea that the mlr-.:t has and distribution of properties ll l<e orl.,1-
crn). 121 an e.xlstenoe separate frorn the mat€flal tation cr color CNef a set o f objects or
"""Id of the body. 111 over a region In a scene. [7]
dlplopla Double vision. If >Asble in
both eyes, stimuli falling outside of duplex In reference to the retina, entorhinal cortex A pt"fy1qJenetl·
Panum's fuslonal area will appear dlp- consisting of two parts: thl rods and caly old cortlca region that provides
loplc. [6] cones, which operate under different the major sensory assoc;latlon Input into
conditions. 121 the hlppocarnpus. The entomlnal cortex
direction Tl1e line one moves along also receives direct projections from
or faces. with reference to the pdnt or E olfactay regions. 1141
region one Is m0\1ng toward or facing. ear canal The canal that corducts
11 21 entry-level category For an ob]oct,
sound vibrations from the pinna to the
tympanlc membrane and prevents the label that comes to m nd mC<Jt
directional transfer tunctlon (DTF) quickly when we Identify it (e.g., •bird").
A measure th9:t describes how the damage to the 1)1npanlc mernbrar1'l. 191
At the subordinate leva. the object
pinna. ear canal. head. and torso eccentricity The distance between might be more specifically named (e.g.,
the Intensity of sounds with thl retinal Image and the fovea. 121 -eagle"): at the superordinate level, It

500 GLCJ5SARY Amma Appa
mlgl1t be m°'e generally named (e.g., extrastrlate body area (EBA) expansion Is one aspect of optic now.
"animal). (4) A reglcn of extrastr1ate visual cortex In (8)
epidermis The outer of two major hUmans that Is specftC<llly and reliably foliate papillae Folds of tissue con -
la'/ ers of skin. (1 3) activated by Images of the body other taining taste buds. Foliate papillae are
than the face. (4)
equal-loudness curve A graph located on the rear of the ton;;iue lateral
plotting sound pressure level (dB SPL) extrastriate cortex The re.glen of to the clrcumvallate papillae, where the
ag alnst the frequency for \'"1lch a lis- oortex bordering the pn rrary \olsual tongue attaches to the mouth. (1 5)
tener perceives constant loudness. [9} oortex and containing multiple areas formant A resorarce of the vocal
lnvowed In visual processing. (4) tract. Ferments are specified by t1101r
equilibrium In reference to the ves-
tibular system, our vesti bular sense F center frequency and are denoted by
comprised of spatial ortffltatlcn pEfoep- familiar size A depth C'-" based on Integers that Increase wlt11 relatr1e fre-
tlon-encornpasslng our perception knowedge of the typical size of objects q...,ncy. (11)
of linear tn)tlon. motlcn. and Ilka humarn or pennies. (6) Fourier analysls A mathematical
tilt - combined wltl1 refiexlve vestibular feature Integration theor; Anne procedure by which any signal can be
responses Ilka posture. vestlbulo-auto - Trelsman's theory ofvlsual attention. separated into cx:-:rnp;::nent slne wtwes
nomlc renexes, and vestlbulo--ocular w l1lch holds that a llmlted set of basic at different frequencies. Combining
refiexes. (1 2) featuroo can be procesood In parallel these sine waves will reproduce the
esotropla Strabismus ln which cne preattentlVely. but that other properties, original signal. (1 , 12)
eye deviates Inward. (6] Including the correct binding of features fovea A smaJ I pit, nearthe centEf of
Euclidean RefEfrlr>;J to the geom - to objects. require attention. (7) the macula, that rontalns the highest
et ry of the world , so named In honor feature sec'"\rch search fe< a target ccncentratlon of ccnes. and no rods. It
Is the portion of the retina that produces
of Euclid. the ardent Greek geometer defined by a single attribute, such as a
the highest visual acuity and serves as
of the third century BCE. In Euclidean salient color or onentatlon. (7)
geometry, parallel li nes remain parallel the point of flxatlm. 121
Fechner's law A principle describing
as they are extended In space. objects the relatlaishlp t.etween stimulus and frame of reference The COCfdlnate
maintain the same size ard shape as resulting sensation that says the mag- systEfn used to define locatlms In
they move around ln space. the internal nitude of subject!\"' sensation Increases spaoe. (1 3]
angles of a triangle always add to 100 to the loganthm of the tree fusion The technique o1con-
degrees. and oo forth. (6] stimulus Intensity. 11 J verg Ing (crossing) 0< d werglng the eyes
event-related potential (ERP) In order to view a stereogram w1tl10ut a
feed-forward process A process
A measure of electrical actMty from a
that carrtoo out a computation (e.g., stereoooope. 161
subpopulatloo of neurcns In response obje:::t recognition) one neural step after frequency Fa sound, the number
to particular stimuli that requires averag- anottl€f. without need for feedback of times per sooond that a pattern of
ing 1rnr11 EEG recordings. (1) from a later stage to an earlier stage. (4) pressure chan;;ie repeats. Freciuency is
exogenous cue In atten- flg..1re-ground asskJ;nment The peroelved as pitch. (9)
tion. an exogenous cue Is located out process of determining that scme functional magnetic resonance
(exo) at the desired final location of reglms of an Image belong to a fore- Imaging (IMRQ A variant of mag-
attentlm. (7) ground object (figure) and other regions netic resonarce !magln;J that makes
exogenous spatial attention are part of the backg round !ground). (41 It possible to measure localized pat-
A form of bottom-up (st lmulus-drtven) fillform papillae Small structures terns of actwlty In the brain. Activated
spatial attention In 'v\'hlch attention Is or the tor>;Jue that provide most of the neurons provoke increased blood ftow,
refiex!Vely (lnvduntanly) d irected toward bumpy appearance. Flllfonn papillae which oan be quantified by measuring
the site at which a stimulus has abruptly have no taste !Unction. (1 5) changes in the response o1 oxygen-
appeared. (1 3] ated and deoXl'!Jenated blood to strong
filter An acoustic. electrical. elec- magnetic fields. (1)
exotropla Strablsmus In which c:ne tronic, or optic device. lnstrurnent 1 com-
eye deviates o utward. (6) puter prc.Qram, or neuron that allows fundamental frequency The low-
explorator; procedure A sterec- the passage of some range of parame- est-frequency canr.:onent of a ocmplex
pe<lodlc sound. (9. 10)
fyped hand movement pattern used to ters (e. g., crlentatlons, frequencies) and
towh cbjects in order to perceive their blocks t he passage of others. (2 , 3) fundus The back layer of the retl na-
properties: each procedure Is best for first-order motion The motion of what the eye doctor seoo through an
determining one (or more) object prop- an object that Is denned by changes In ophthalmosccpe. (2)
erties. (1 3] lumlnance. (8] fungiform papillae Mushroom-
extinctton In visual attention. the flavor The combination of true taste shaped structures (maximum diameter
lnablllty to perceive a stimulus to one (sweet . salty, sour. bltt€f) and retronasal 1 mllllmeteO that are distributed most
skJe of the point of fixation (e.g .. to the densely on the edges of the tongue,
olfactlon. (1 5]
nghtj In the presenoe of another stl!rn- especially the tip. Taste bl.ds (an aver-
lus. typically In a comparable position focus of expansion The point In the age of six per papilla) are burled In the
In the other \olsuaJ field (e.g., on the Ian centef of the horizon from which. when sulface. (1 5)
skJe). (7) we're In motion (e.g .. drMng on the
highway), all points In the p€<Spectlve fusiform face area (FFA) A region
Image seem to erranate. The focus of of extrastrlate visual cortex In humans

GLOSSARY 501
that Is specllc ally and rellallly actr/ated Integrate Input from all the earlier pro- horopter The locatlm of objects
by human faces. (4. 7] jections, and are thought to be the whose Images lie on corresfX.'f"'ldlrQ
basis of specll c odaant k:Jentmcatlon. points. The surface of zero disparity. (6]
G (14j
G protein-coupled receptor
hyperalgesia An increased or
graV1ty A force that attracts a body heightened response to a normally
(GPCR) Any of the dass of receptors
that are present en the surface of olfac- toward the center of the Earth. (12] painful stimulus. (1 31
tory sensay neurons. All GPCRs are guided search search '1 v.llich hypercolumn A 1-ml Illmeter block
charactE!flzed by a commcn structural attentlrn can be restricted to a subset of striate cortex containing two sets of
feature of seven membrane-spanning of possible Items on the basis of lnfa- cdumns. each covering wary possible
a-helices. (141 matlon about the target Item's basic orientation (0-1BOdegrees). with one
ganglion cell A retinal cell that feaMes (e.g., Its color). 171 set preferring lnpLrt from the left eye and
receives vSual lnfotmatlcn from photo- gustatlon The sense of taste. (141 or-e set preferring Input from the right
receptors via two lnten-nedlate neuron eye. 131
types plpciar cells and amacrlne cells) H hyperopia Farsk;;:lhtedness, a com-
am transmits Information to the brain hair cell Any c<JI that has stereocll la mcn 0Cf1dltlon In which light entering
ard mldbraln. (21 for tranOOuclng mechanK::aJ movement the eye ls focused behind tl1e retina and
In the Inner ear into neural actlvtty sent accommcdatlon Is required In crder to
gate control ttieory A deecnptlon to the brain; some hair cells also recetve
of the pain-transmitting system that see near objects dearly. 121
Inputs from the brain. (9, 12]
Incorporates modulating signals from hyperpolarlzaHon An Ina-ease In
the brain. (1 31 haptic perception Kr<Mitedge of membrane potential such that the inner
the wortd that Is derived from sensory membrane surface beccmes more
geon In Blederman's recog nltlon-by- receptors In skin. muscles, tendons.
oomponents moda , any of the •geo- negatl\i'e than the outer membrane sur-
and joints, usually lnvolvtng actwe faoe (2]
metric Ions" o ut of v.lllch perneptual expjor,.Um. (1 31
objects are built. (41
haptic virtual environment A syn-
Gestalt grouping rules A set of thetic world that may be experienced Illuminant The light that Illuminates a
rules describing which elements In an haptlcally by operation of an aectrome- surface. (51
Image will appear to group together. dhanlcal device that da"9rs faoes to Illusory conjunction An erronea.is
The on\j naJ list was assembled by the hand of the user. (1 31 ccmbffiatlon of two features In a visual
members of the Gestalt scto::>I of sC&le-for exan1ple. seeing a red x
thought. 141 hanmonlc spectrum The spectrum
of a complex sound In which energy Is when the display 0Cf1talns red letters
Gestalt In German. literal tt "form." at lnte;ier multlptes of the fundamental and Xs but no red Xs. (71
In reference to peroeptlm, a schcd of frequency. (91 Illusory contour A contour that S
thought stressing that the perceptual perceived even though nothing changes
w1101e oould be greeter than the appar- hellcotrema The opening that con-
nects the 1>fnperlc and vestibular from cne side of lt to the other In an
ent sum of the parts. (41 Image. [41
canals at t11e apex of the cochlea. (91
gestation Fetal development during Image A picture or II keness. (2(
pregnancy. (141 hertz (Hz) A unit of measure for fre-
quency. One hertz e:iuals me cycle per Imbalance Lack of balance:
glabrous In reference to skin, second. (91 unsteadiness: nearty falling war. (1 21
hair. (1 31
heterodi mer A chain of two mol- inattentional blindness A fallure to
global superiority effect The fim-
ecules that are differmt fran each notice - or at least to report- a s1lmulus
lng In various e:xpenments that the other. (1 51 that WOLrld be easily reportable If It were
prcpertles of the 1.vhole object take pre- attemed. 171
oedenoe over the properties of parts of heuristic A mental shortcut. (4 I
1he ct>ject. (4 I high-spontaneous ftber An aLKll- incus The mid::lle of the three ossi-
tory nerve fiber that has a high rate cles. connecting the rnalleus and the
glomerulus Any or 1he spherlC<ll stapes. (91
oonglomerates containing the lnCO'l'llng (nue than 30 spikes per se0Cf1d) of
axons of the olfactcry sensory neurons. spontanoo.is firtng ; hlgh -spmtanoo.is Inferior colliculus A mldbraln
Each OS N omverges onto 1wo glcm- fibers Increase their flrlrg rate In nucleus In the aLK!ltory pathway. (91
erull (one moolal, one lateral). (141 respmse to relatlvay low levels of
sound. (91
lnferotemporal on cortex Part of
good continuation A Gestalt group- the cerebral cortex In the tower pcftlon
ing ruk> stating that two elements will homologous regions Brain reglais of the temporal lobe, Important In object
tend to group together If they seem to that appear to have the same furx:tbn recognition. (41
lie on the sarre ocruour. (4] In different species. (41 Inhibition of return The raatl>/e dif-
graded potential An electnc al homunculus A maplike represoota- ficulty In getting attention (or the eyes)
potential that can vary continuously In tlon of regions of t he bcdy In the brain. to move back to a recently attended (or
amplltude. (2] (1 31 flxatoo) location. (7]
granule cells Like mltral cells, gran- horizontal eel I A specialized retinal Inner ear A hollow cavity in the tem-
ule oells are at the deepest level of cell that ocruacts both protoreceptor poral bone of the skull. and the s1ruc-
the dfactorybulb. Theyoompnse an and blp<>ar oells. (21 tures within this cavtty: the cochlea ard
extensive network of lnhlbltay neurors.

502 GLOSSARY
the semicircular canals of the vestlbu8' kinesthetic Referring to perception also referred to as .. translation.n Linear
system canals. (9J lnvoMng senOCf)' mecl1a11oreoeptors In acceleration. linear vaoctty. and linear
Inner segment The part of a pho- musdes, tendcns, and joints. (13] displacement all mathematically repre-
toreceptor that lies between the cuter koniocellular cell A neuron located sent linear motion. ( 12J
segment and the cell nlldeus. (2J between tl10 magnocellular and parvo- linear moUon Translational motion
insular cortex The primary cortical cellular layers of the lateral genk:ulate like the predominant movement of a
processing area fa taste -the part of nucleus. This layer is kn:Min as the train car a bobbtehead doll. (1 2J
the oor1ex that first receives taste Infer· konlocellular layer. (2, 3J linear perspective A depth cue
rnatlon. Aloo calW JnsuJa or gustatOI}' koniocellular Referring to cells In the based on the fact that lines that are
COlllj'C.(1 5J korJocelluar layer of the lateral genlcu- parallel in the three-d lmenslonaJ wcrld
lnteraural level difference (ILO) late nuciaus of the thalamus. Konlo w111 appear to converge in a two-dimen-
The dlference In level Ontenslfy) from the Greek for "dust" referring to sional Image. (6J
between a scund arrMng at one ear the appearance o f the cells. (5] lordosls The position that females
versus the other. (1 OJ L of some species (e.g .. pigs and rats)
lnteraural time difference (ITC) L-cone A cone that Is preferentially need to assume In order to be lmpreg-
lhe dlference In tme betWeen a sound senstlve to long wavelengtl-e: colloqui- rated. It Involves the Inward curving of
arrMng at one ear versus the other. (1 OJ ally (but not entirely acourateM known the spinal OOurnn and expceure of the
as a "red cone.· (5J genitals. (14]
interocular transfer The transf€f of
an effoct (such as adaptation) from one labeled lines A t11eay of sens()!)' loudness lhe psychobgloal aspect
6'f0 to the o ther. (8] coding In which each nerve lber carrloo of sound rel9ted to perceived Intensity
(amplttude). (9J
inverse.square law A pri nciple a particular stimulus quality. (1 3, 15J
statlrg that as distance from a ocurce lateral genlculate nucleus low-spontaneous nber An auditory
Increases, k1ter.slty decreases faster (LGN) A structure In the thalam us, nerve fiber that has a low rate than
such t11at deaease In Intensity Is equal part of the mldbraln, that receives Input 10 spikes per seo::;nd) of spontaneous
to the distance squared. This general from the retina! ganglion cells and has llrlng: low·spontaneous fi bers require
law also to optlos and other Input and output connections to t11e relatively Intense sound befae they ,.;11
fom1s of energy. (Kl] visual cortex. (3, 5J nre at higher ratoo. (9J
lpsllateral Referring to the same side lateral Inhibition Antagonistic neural luminance-defined object An
objoct that Is delineated by differences
of the body(or brain). (3. 14J lnteractkln between adjacent regions of
In rellected light. (8J
lpslleslonal fteld ll1e -Asual lleld on trie retina 1
21
the same side as a brain lesion. [7] lateral superior ol Ive (LSO) A relay M
Iris lhe odcred part of the "f", con- station In the braJn stem where Inputs M ganglion cell A ganglion cen
sisting of a muscular diaphragm sur- from OOth ears contribute to detection resemblirg a nttle umbrella that receives
rounding the p upil and regulating the o f the lnteraural level difference. (1 OJ exc itatory Input from diffuse bipolar cells
light entering the fI'{e by expanding and learned taste aversion The avold- and feeds the magnocellular layer o f ths
contracting t11e pupil. (2J anoe of a ro\1€1 flavor after It has been lateral genlculate nucleus. (2]
Isol ntenslty curve A map plotting paired with gast rtc Illness. The smell , A cone that Is preferentlalty
the fln ng rate of an audltcry nerve lber not the taste, of the substance Is key sensitive to mlddle wavelengths; c ol-
against varyirg frequencies at varyin;;i for the tearnoo aversion respcnse in loculally !Put not accuratelY)
humans. (14] known as a wgreen cone." (5]
lntensnles. (9J
lens The lens Inside the 6'f0 that macula In referen:;e to vision. the
J enables the charging of focus. (2J oertral part o f the retina thal has a hlgl1
Just noticeable difference (JND) lesion In reference to neurophyslol- ccncentratlon of ccnes. Jn referenoo to
or difference threshold The small- the vestibular system, any of the spe-
est detectable difference between t'#O ogy, 1. (n) A region of damaged brain.
cl9Jlzed detectcrs of linear
stlmuU, or the minimum charr,Je In a 2. M To destroy a section of the brain.
and gravity fa.md In each otolith crgan.
stimulus that enables It to be ocrroctly (4J
(2, 12J
Judged as different from a reference limbic system The encornpassk1g
stimulus. (1J group of neural stn.ctures that Includes magnetic resonance imaging
trie olfactory cortex, the amygdala, the (MRI) An Imaging technolo;iy that
juxtaglomerular neurons The first uses the responses o f atoms to strong
layer o f cells surrwnding the glomerull. hlppocarnpus, the plrlfonn cortex, and
t11e entcrhlnal ocrtex. lhe llmblc system magnetic fields to form Images of struc-
They are a mixture of excltatcry and turoo like the brain. The method can be
anc
Inhibitory oells respcnd to a wide
ls Involved in many aspects of emotion
and memory. Olfactlon Is unique a mong adapted to measure activity In the brain,
range of odcrants. The selectMr1 of as well (see tuncaona/ magneac rooo-
neurons to specific cdorants Inc reases tl1e senses for Its direct connection to
na1JCe lmagltl(fl. (1]
In a gradient trom the surfaoe of t11e the llmblcsystem. (14J
dfactory bulb to the deeper layers. (14J linear acceleration The rate of magnetoencephalography (MEG)
change of llnear veloclty. Mathemati- A technique, simi lar to eloct roencepha-
K cally, the Integral of linear acoeleratlon Is lography, that measuroo c hangoo In
klnesthesla Perception of the pool- linear velocity, and the Integral of linear magnetic activity across p::pulatlons of
tlon and movement of our limbs In vaocny ls linear displacement, Whloh Is many neurons In the brain. (1J
space. (1 2, 13J

GLOSSARY 503
magnitude estimation A psycho- melody A sequence of notes cr middle ear An air-filled c ham·
physical method In w hb h the partlc l· chords percetved as a eJngle coherent ber containing the mlddle bones , or
pant assigns values accordin;;J to per- structure. [11] osslcles. The rni:Jdle ear =rveys and
r.elvoo magnitudes o f the stimuli. [1 J Merkel cell neurite complex ampllles vlbratbn trom the t)<npanlc
magnocellular layer Either of the A specialized nerve ending associated membrare to the oval window. [9]
bottom Mo neuron<:ontalnlng layers of w ith slowly adapting fibers that middle temporal area (MT) An
the lateral genlc ulate nudeus. the cells have small receptive fields. [1 3] area o f the brain thought to be Impor-
of which are physically larger than those metamers Different mixtures of tant In the perception of motion. [8]
In the top four layers. [3] wavelengths that lock Identical. More midget bl polar eel I A small bipolar
main olfactory bulb (MOB) The any pair of stimuli that are oeHIn the central retina that reoeives
owactory bulb: the b lueberry-slzoo perceived as Identical In spite of ptr/sl· Input from a single cone. [2]
extension of the brain just above the cal differences. [5] mitral cell The deepest layer of neu·
nose: the first reglai o f the brain where method of adjustment A method rons In the olfactcry bulb. Each mltral
smells are processed. In humans we o f limits In which the subject controls cell respcnds only to a fWJ spedflc
refer si mply to "oWactory bulb(s)"; In anl·
the change In the stimulus. [1 J odorants. [141
rnals vllth accessory olfactory bulbs, we
distinguish between •main" and method of constant stimuli monocular depth cue A depth oue
oory." [141 A psychophysical method In w hic h that Is avallable even when the wald Is
many stimuli, raitjng from raretyto \lle'M>d with one ffie alone. [6]
malleus O'le o f the three osslcles. almost always perceivable (or rarely to
The malleus receives \llbratlon the monocular With me eye. [6]
almost always perceivably different fraTI
tympanlc membrane and Is attached to a reference stimulus). are presented monosodlum glutamate (MSG)
the k'lc us. [9] cne at a time. Participants respond to The sodium salt o f glutamlc ackl Ian
masking Using a second sound . each presentation: •yestro.n "same/dif- amino acid). [1 5]
quently nose, to make the detection of ferent." and so on. [1 I motion aftereffect (MAE) The lllu·
another sound more dlffloult. [9] method of limits A psychophysical slon of motion of a stationary object
materialism The klea that the only method In which the particular dlmen-. t11at occurs after prolonged exposure to
t1111-.;i that exists Is matter. and that all sm of a stlrnulLJS. or t110 dlfferenoe a moving object. [8J
things. lnolu::Jlng the mind ard con · bet'Neen two stimuli. is varied Incre- motion parallax An Important depth
oclousness. are the results of Interaction mentally unt II the participant responds cue that Is based on head movement.
between bits of matter. [1] differently. [1] The geometric lnformatlai obtained
mathematical Integration Corn · metrical depth cue A depth c ue from an eye In tv10 different positions at
an lntet1ral-one of the two rm.Jn th:.1t provides quantitative infamatlon two different times Is similar to the lnfor-
operations In caJoulLs other. the about dlstan::e In the third dimension. matlcn 1i"orn two eyes ln different pooi-
operation , Is dlfferentlatlor1). [6] tlons In the head at the same time. [6]
Veloclty Is the Integral of aoceleratlon. mlcrosaccade An Involuntary, small. muscle spindle A sensory receptor
Change of position Is the Integral of 'jer]Sll$e e:l'j 11110'l91119nt, [8] IOCated In a muscle that senses Its ten ·
velocity. I121 slon. [13]
mlcrovllll Slender projections of the
mechanoreceptor A senrory cell membrane on the tips of some myopia Nearslghtooness. a ocmrnon
receptor that responds to mechani- taste bu::J oels that extend Into the taste coodltlon In which light entering the eye
cal stlmulatlai (pressure, vibration, or Is focusoo In tront of the retina and dis·
pore. [1 51
movement). [1 2. 13] tant objects cannot be seen sharply. [2]
mid-spontaneous fiber An audl·
medial geniculate nucleus The tay nerw fiber that has a medium rate N
part of the thalamus that relays audi - (10-30 spikes per secaid) of spontane·
tory signals to the temporal cortex and narve template theory The pro·
cus Iring . The c haracteristics ofmld- posal that the visual system recognizes
receives input from the auditory cortex. fibers are Intermediate
[9] objects by matc hing the neural repre·
between low- and hlgh-spontanoow sentatbn o f the Image with a stored
medial superior olive (MSO) A ftbers. [9] representation of the same "shape" In
relay station in the brain stem where middle (mldlevel) vision A the brain. [4]
Inputs from both ears contribute to defined stage of \llsual processing that nasal dominance The asymmetry
detection oftl1e lnteraural time differ- comes after basic features have been c haracterizing the intake of air by the
ence. [1 0] extractoo from the Image Qow-level. a two r'K:€trlls1 •..vhlch correspords to dif-
Meissner corpuscle A specialized early1 vision) and before cbject reo:ignl- fering sensitivity to odaants between
netve ending associated 1"1th fast- llon and soene understanding Qi>;Jh- the two nostrl B. Nasal dominance alter-
adapting iFA Ii lbers that have small level \llslon). ]4] nates nostrils throughout t11e day, but
receptive ftelds. [1 3] middle canal One of three luld-filloo there Is no proolctablllty about when the
melanopsin A photoplgment, found passages In the occhlee. The middle nostnlsalternate. [14]
In a class of photoreoeptlve retinal gan- canal Is sandwiched bet.-i the tym· Necker cube An outline that Is per-
glion cells, that Is sensHive to ambient panb and vestibular canals and ccn- ceptual ly bl-stable. Unlike lhe situation
llgnt. [2. 5] talns the cochlear partltlai. Also calloo with most stimuli. two Interpretations
sc,,Ja meda. [9] continually battle for perceptual doml ·
nance. [4]

504 GLOSSARY
negative afterimage An afterimage neurons by which they demonstrate a olfactory epithelium A secretory
wl1ose p::larlty Is the opposite of the preference. respondlrg sornevA1at more mucous membrane In the hurran nose
erlglnal stimulus. Light stimuli produce rapidly when a stimulus Is presented In whose primary function ls to detect
dark negative atterlmages. Colors are one eye than when it Is presented In the odorants In Inhaled air. Located en both
complementary; fcr exainple, red pro- o ther. [3] sldas of the upper portion of the nasal
duces green, and yellow produces blue. C<Mty and the olfactory cletts. the dfao-
oculomotor nerves The third
(5] palr of cranial nerves, which Innervate all tory eplthellum contains three types of
neglect As a nairol:::glcal symp- the extrinsic muscles of the eye exoept
cells: dfactory sensory na...ircn s, 00.sal
cells, and supporti ng cells. [141
tom . In vsual attention: (1) The Inability the lateral rectua and the superior
to attend or respond to stimuli in the oblique muscles. and which lnnavate olfactory sensory neuron (OSN)
contraleslonal visual field the the elevator muscle of the upper eyelid , One o f three cell types-the main
lett leld after nght parietal damage). (2) the cl llary muscle. and the sphl nct€f one- In the OSNs
lgncrlng half of the body or half of an muscle of the pupil. (1] are small neurons located beneath a
ctJject. (7] odor hedonics The liklng dimension mucous layer In the epithelium . The
neural plasticity The ablllty of neural of OOor perception. typically measure::l c llla en the OSN dend ntes contain the
circuits to undergo changes In fun:tlCf"l by ratings of an odors 's perceWed receptor sites fcr odorant rrdea.1les.
[14]
er erganlzatlen as a result of previous pleasantness. famil iarity, and Intensity.
activity. (1 3] (14] olfactory tract The burdle of axons
neurolmaglng A set o f methods that odor The translatlcn of a chemical of the mltral and tufted oells w ithin the
generate Images of the strLcture and/ stimulus Into a smell sensation. For olfactory bulb that sends odor lnferma-
er function o f the brain. In many cases, example, 'The c alke has a c llC<XJlate tion to the p rlmarydfactory cortex. (14]
these met11ods allow us to examine the odor." (14] olfactory white T he olfactory
brain In living, behaving humans. (1] odorant receptor (OR) The region equivalent of •white noise" or the color
neurotransmitter A chemical sub- on the cilia of dfactory sensory neurais white. VVhen at leiast 30 cdcrants of
eq ual Intensity that span oWactory phys-
stance used tn neuronal ccmmunlcat1on where oderant moeculoo bind. [14]
lochembal and psychological (percep-
at synapses. (1 I od!>ran A rml©.Jle Hlat 11> defined tual) space are mixed, they produce a
neutral point The point at w hbh an by Its physlcochenioal c t...-actenstlcs. resultant odor percept that Is the same
cpponent color mechanism Is generat - which Is capable of belrg translated by as every other mixture of 30 odaants
Ing no signal. If redi)reen and blue- the neM>US system nto tile percep- meeting the span and equivalent Inten-
yellow mechanisms are at their neutral tion of a smell. Fcr example, •vou were sity c riteria. even t hough the vancus
p::l nts. a stimulus v,111 appear achro - glveri the odorant methyl sallcylate to rTixtures do not share any common
matic. (The black-white prooess has no smeU, which has the odorof wtnter- odorants. [14]
neutral point.) [5] green mint." (14]
ON bipolar cell A bipolar oall that
noclceptor A sensory receptor OFF bipolar cell A bipolar oell that responds to an Increase In light cap-
that responds to pal nful Input. such as responds to a docrease In light captured by the ocnes. [2]
extreme heat or pressure. (1 3] tured by the c ones. (2]
ON -center cell Aoell that depolar-
nonaccldental feature A feature of o FF-center cell A oell that depolar- izes In response to an Increase In light
an object that Is no t dependent on the izes ln response to a decrease In light Intensity In Its receptive-field oenter. 121
exact (or accidental) viewing po,;tlon o f Intensity kl Its receptlve-flekJ center. [2]
the observer. (4] opponent color theory The theory
olfaction The sense of smell. [14] that perception of color Is based on
nonmetrical depth cue A depth olfactory (I) nerves The first pair of the output of three mechanisms, each
cue that provides Information about cranial nerves. The axcns of the olfac· of them resultltlg rrom an oppcnency
the depth order (relative depth) but not tory sensory neurons bundle together between tv,io cdas: red·green. blue·
depth mag nitude (e.g., his nose Is In after passing thrcugh the c nbnfam yellCMI, and blac k-white . [5]
front of hs faoe). [6] ptate to form the dfactory nerve. w hich optic (II) nerves The second pair
nontaster (of PTC/PROP) An conducts imputses from the olfactory of cranial nerves, w hklh arse rrorn the
\'idual born with two recessive alleles for eplthella In the nose to the olfac tcry retina and carry visual Information to the
the TAS2R38 gene and unmle to taste bulb. (i . 14] thalamus and other parts o f the brain.
the cornpa..1nds phenylthiocarbamide [1]
olfactory bulb A b lueberry-sized
and prq:iytthlouracil. [151
extension of the brain just at:ove the optic array The c ollection of light
0 nose. Wl")€(e olfactory Information is rays that Interact wtth objects In the
occlusion A cue to relative depth first processed. There are two dfactory world that are Jn front o1a vielNer. Tarn
erder In w hic h. for example, one object bulbs, me In each brain hemisphere, ooned by J. J. Gibson. (SJ
cbstructs the view of part of anott""er oorrespond lng to the right and left nos-
trUs. [14] optic flow The pattern o f apparent
ctJject. [6] motion of objects In a visual scene pro-
olfactory clett A Mrrow space at duoed by the relatwe motl::fl between
octave The lnte<val be1'Mlen two
the back of the rose Into which air the obserVer and the scene. (6, BJ
scund frequencies having a ratio of 2:1.
(11] ibws and w here the oWactory epithe-
optoklnetic nystagmus (OKN)
lium Is located. [14]
ocular dominance The property A renexlve eye movement In which the
of the reoepllve fields of striate catex

GLOSSARY 505
eyes wlll lnvduntanly track a contlnuaJ Jy oval window The flexible opening to perception The act of gMng mean-
m<Mng object. [81 the cochlea throu;ih vkolch the stapas ing to a detected sensation. 111
orbitofrontal cortex.(OFC) The transmts 'llbratlon to the ftuld lrslde. 191 period For hearing, the time raq ulred
part of the frcntal lcOe of the oortex that p fer a fun wavelergth of an acoustic sine
lies behind the bone (orbit) contaJnirQ P ganglion cell A smaJI ganglion oel waw to pass by a ix;nt In space. 111
the eyes. The OFC Is responsltJe for the that receives excitatory Input from single phantom limb Sensation peroeMld
conscious experlerce of olfactlon, as midget plpolar cells In the central retina from a physically amputated limb of the
well as the Integration of pleasure and and feeds the parvocellular layer of the body. 11 3]
displeasure from food: and It has been lateral genloulate nudeus. 121
retenad to as the second81)' olfactory phase locking Firing of a slngle neu-
conex and the secondaf}' taste cortex. Paclnlan corpuscle A spaclallzed ron at one distinct point In the period
The OFC Is aJso lnvdved In many other nerve ending associated with fast- (cycle) of a sound wave at a given fre-
functions, and It Is critical for asslg n- adaptlrg (FA II) fibers that have large quency. (1ha neuron nood not fire on
lrQ affective value to stimuli-In other receptive !aids. 11 3] every cycle. but each !ring will occur at
words. determining heda1lc meaning. panpsychlsm The Idea that the the same point In the cycle.) 191
[14, 151 mlm exists as a property of all matter- phase A fraction of the cycle of the
organ of Corti A structure on the that Is, that all matter has oorsclous- sine wave described In de;iroos (0° to
basilar membrane of the cochlea that is nees. 111 360") cr radians((}.. to 2o). In hearlrQ,
composed of haJr cel ls and dendrites of Panum•s tuslonal area The region phase can be used to deocribe frac-
auditory nerve fibers. [9] of space, In tront of and behind the tions of a period that relate to time. 11 J
orientation tuning The tendency horopter. within which binocular slrxile phase The relat"9 po,.tlon of a
of neurors In striate cortex to respord Vision Is pos,.ble. 161 gratlrQ. 131
optimally to certain orientations and less papilla Any of mllltlple structures pheromone A ctiemlcal emitted by
to others. [31 that give the tongue Its bumpy appear- one member of a species that triggers a
orthonasal olfactlon Sniffing In and ance. From smalloot to largest. the phy$doglcal or beha'lloral response In
paroeMng odors thrrugh rur nostrils, papilla type5 that contain taste buds anothef member of the same species.
wl1icl1 occurs wl1en we are sm911ng are funglform, foliate, ard cl rcumval - Pherorn::ines are slgn:lls for chem ical
oomathlng that Is C>.Jtslde of us. [14] late: ftlWorm papillae. which do not con- o.:rnrnunlcatloo and do not need to
tain taste buds, are the smallest and have any smell. [141
oscillatory Referring to back-and- moot numerous. 11 5]
forth movement that has a constant phonatlon The prooess through
rrr{thrn . [1 21 parabelt area A retJlon of cortex. which vocal folds are made to vltl"ate
lateral and adjacent to the belt area. wtlen air pushes out of the IUJ'QS. 1111
osslcle Any ol three tiny bones of the where neurons respcnd to more o.:rn-
middle ear: malleus, Ina.is, and stapes. photoacuvatlon Act,'atlon by light.
plex characteristics of sourds. as wall 121
191 as to Input from other senses. (9]
otltls media k1flammailon of the photon A quantum of visible light or
parahlppocampal place area (PPA) other form of e ectrcmagnetlc radiation
rnlddle ear, comrncnty In children as a A regbn of extrastrlate visual ccrtex In
result of Infection. (9] demonstrating both particle and wave
humans that Is spedlcally and reliably
properties. [21
otoconia 1lny calclurn carbcnate activated more by Images of places
stones in the ear that prcMde lnertlaJ than by other stlrmrll. 14 . 7] photoplc Referrlrg to light Intensi-
mass for the otollth organs, enabllng ties that are bright anrugh to stimu-
parallel search A search In which late tlie oore reoeptors and bright
them to sense gravity ard Hnear acoel- multiple stimuli are processed at the
eratlcn. 1121 enotgh to •saturate" the roc1 receptors
same time. 171 (that Is, drive them to their maxmum
otollth organ Etther of two mechani- parallelism A rule for flgure-groun:I responses). 151
cal structures (utrlcle and saccule) In the assignment stating that parallel con-
vestibular system that sense both linear photoreceptor A llgl1t-senslt"9
tours are to belorg to the same recepta in th9 retina. (2]
acceleration and gravity. 11 2] figure. [41
otosclerosls Abnormal growth of pictorial depth cue A cue to dis-
parietal lobe In sach oarebral hemi- tance or depth used by artists to depict
the middle-ear tx:nes that causes hear- sphere, a lobe that lies toward the top
lrQ loos. [91 throo-dlmemlcnal depth In two-dlmen-
of the brain between tile frontal and slonaJ pictures. [61
ototoxic Produ:;ing adverse effects = lpltaJ lobes. [71
on coc:hlear or vestltdar organs or pinna The outer. funnel- like part of
parvocellular layer Any of the top theear. 191
nerves. 191 four neuron-containing layers of the
outer ear The external sound-gath- lateral genlculate nudeus, the cells of pitc h The psychdogicaJ aspect of
erlrxi portion of the ear. oonslstlrQ of which are phy$cally smaJler than those sound related mainly en the fraquenoy
the pinna and the ear canal. 191 In the bottom two layers. 131 of >Abratlon. 19, 111
outer segment The part of a ph:>- parvocellular Referring to oells In t11e place code Tuning of different parts
toreoeptcr that contaJns ph:>toplgment parvocellular layers of the lateraJ genlcu- of the cochlea to different traquendes,
121 late nucleus of the thalarnLS. PaNo In which lnfonmatlon about the particu-
from the Greek fcr •small" referring to lar frequercy of an Incoming sound
the size of the calls. 151 wave Is ooded by the plaoe along the

506 GLOSSARY
cochlear partition that has the greatest probability summauon The and tunher stimulation ts Incapable of
mechanical displacement. [91 Increased detection probability based lncr-ng the firing rate. [91
placebo effect Decreasing pain on the statistical advantage of having rate-lntens tty func tion A graph
sensation when people think they're two (or more) detectors rather than one. p lo ttin g the firtng rate of an auditory
taking an analg esic drug but actually 161 nerve fiber In response to a soum of
are not. [1 31 projective geometry For puiposes ccnstant frequency at lm reasing lnt'en·
polysensory Referring to blending o f stcld)1ng perception of the three- sltles. 191
multiple sen scry systems. [11 dlrnenslonal wond, the geometry that r eactlon11 rno (RT) A rneasure o f the
describes the transformations that time !rem the onset of a s!lrrulus to a
positMsm A philosophic al occur when the three·dlmenslonaJ
arg uing that all we real ly have to go on response. l7J
world Is projected onto a two-dimen-
Is the evlderce of the senses, so the sional sur1ace. For exarr ple, parallel realism A phliosq:ltlcal position
wortd might be rothl ng more than an lines do not oorwerge ln the real 'NCTld. arguing that there Is a real wor1d to
elaborate halluc lnatlon. [61 but they do In the t>IK>-dlmenslonaJ pro- sense. 16]
positron emis sion tomog raphy jection of that world. 161 receive r o perati ng charac t erlsUc
(PET) An Imaging technology that proprto ceptlon Perception medi- (ROC) c urve In studies of signal
enablee us to d elne locations In U10 ated by kinesthetic and Internal recep - detection, the graphb al p lot of the hit
brain where nsurors are especially tors. 1131 rate as a function of the false·alarm
actr.<e by measuring the metabolism of rate. If these are the sarre, pd nts fall
brain csl ls cslng safe radioactive lso- prosopag nosla An Inability to recon the diagonal, Indicating that the
topee. [11 ogn ize faoas. 141 observer canrot tell the dlff€fence
protanope An indivk:l ual win suffers between the presence and aooenoa of
prnatte ntlve stage The process-
from color b lindness that Is due to the the signal. As the observer' s sensttlv-
k1g of a stimulus that occurs before
absence of L-cones. 151 lty lr-.:reases, the curve boNs upward
selective attention Is deployed to that
toward the up per left ocrner. That point
stimu lus. 171 proximity A Gestalt grouping rule
stating that the ta1dency of two fea- represents a pertect abl llty to distinguish
prefrontal cortex A region of the sl;Jnal trom noise (100% hits, 0% false
brain concerned cog nition and tures to group togeth€f w111 Increase as
the distance between t hem decreases. alarms). 111
control. [131
141 receptive f ield The regbn on the
presbyopia Literally 'old slgl1t. " The retina In Which visual stlmull ln1lueroe a
age-related loss of accommoclation, psychoacoustics The study of the
psychological correlates of the physical neuron's flrlng rate. [2]
w hich malkee It to focus on near
objects. [21
dlrnensJcns of acoustics: a branch of receptor adaptation The bloci1"1'1 e
p syd>0physbs. 191 cal pherrrnencn, oo::;urrirQ after OCfi·
primary audttory c ortex (A1) The tlnual exposure to an odorant, whereby
first area w ithin t he temporal lobes of psychophysics The science of
defining quantltatr.<e relat bnshl ps receptors step respcndlrQ to an odor·
the brain respmslble for processing ant and detection ceases. 1141
acoustic lnfcrmatlon. 19] between physical and psyctdoglcal
(subjective) events. 11, 141 receptor potential A change In volt·
primary o lfactory c ortex or plrl- age across the rnem brane of a sensory
pupil The dark, circular opening at
fonn cortex The neural area where receptor cell (In the vestibular system, a
dfactory Information Is l rst prooessed. the canter of the Iris In the we, where
light enters the eye. 121 hair ool) In reepmse to stimulation. 11 21
It ccrnprlses the arnygdala, parahlp-
recognition- by- components
pooampal g;rus, and Interconnected a model Blederman's model of otject
areas: and it Interacts closely with the qualla (si ng. qua/e) In phi losophy, reoognltlcn whic h holds that objects
entorhlnal cortex. 1141 private consdcus experiences of sen· are recognized by the Identities an<I
primary visual cortex (V1). area satlon or perception. 11 . 5] relatlcnshlps of their component parts.
17, or striate cortex TI10 area of 141
the cerebral oortex of the brain that R
receives d irect Inputs trom the laternl random dot stereog ram (RDS) reflect To redirect ocmethlng that
genlculate nudeus, as well as feedback A stereogram made of a large number strt kes a surtaoa - especlally ll;Jht.
(ofien In the thousands) of sound, a heat-usually back toward Its
frcrn other brain areas. 131
placed dots. Randcrn d ot stereograms point of ortgln. 121
primer pheromone A pha"omone contain no monocular cues to depth.
that tnggers a physiolog ical (onen hor- renec tance The percentage of ll;Jht
StlmuH vlslble stereosccplcally In ran· h itting a sur1ace that Is refiected and
rnonal) change among oonspeclncs. dom dct sterecgrarns are Cyclcpean
This effect usually ln'lolves prolonged rot absabed Into the surface. Typically
stimu li. [61 rel ectance Is as a function o f
pheromone exposu re. 1141
ra pid serial visual pres entation wavelength . [5]
principle of unlvarlance The fact (RSVP) An experimental prooedure In
that an Infinite set of different WCNe· reflexive eye moveme nt A move·
wh ich stimuli appear In a stream at one ment of the we that Is autcrnatb and
length-Intensity con1blnatlons can elic n looatlon (typically the point of fixation) lnvduntary. (81
exactly the same response fran a at a rapid rate (typically abcut eight per
s lngkl type o f p ho to receptor. One pt10- second). [71 refract To alter the course of a
toreceptor type cannot make color dls· wave of energy that passes into sane·
c rlmlnatlons based on wavelengt h. 15] rate saturation The point at which a thing from anoth€f medium, as water
nerve fi ber Is firing as rapidly as

GLOSSARY 507
does to light entenr>;J It from the air. 2. behind the palate Into the nose. such scotoma Tile blind spot In the mid-
To measure the degree of refractlcn In a odor sensations are perceived as origi- dle of the visual l eld. 121
lms or "fe. 121 nating from the mouth . even ttJough the scotoplc Referring to light Intensities
Reissner's membrane A thin actual ocntact of odorant and receptcr that are bright enough to stimulate the
sheath of tissue separating the vestibu- occurs al the cifactory mucosa 1151 rod receptors but too dim to stimulate
lar and middle canals In the cochlea. [91 rhodopsln The visual pigment found the oone receptcrs. 151
relatabillty The degree to which two In roos. 121 second- order motion The motion
line segments appear to be part of the rod monochromat An lrdti.r1dual of an object tllat Is defined by changes
same contour. 141 wrth no cones of any type. In addition In contrast or texture, but not by lumi-
related color A color. such as to being trulycobr-blind, rod mono- nance. 181
brown or gray, that is seen only in chrornats are badly visually Impaired In sele ctive attention The form of
relation to other colors. For example. brtght light. 151 attention Involved when processing Is
a "gray" patd1 In complete darkness rod A photcreceptcr specialized for restricted to a subset of the possible
appears white. 151 night vision. 121 stimuli. 17]
relative height As a depth cue. round window A soft area of tissue semi cl rcular canal A11y of three
the observation that objects at differ- at the base of the tympan;, carol that torddal tubes In the vootlbular system
ent distances from the \'!ewer on the releases excess pressure remaining tllat sense angular motion. 11 21
ground plane V.0 11 form Images at differ- flcrn extremely Intense sounds. 191 sensaUon The ability to detect a
ent heights In the retinal Image. Objects Ruftlnl ending A specialized nerve stimulus and. perhaps, to turn that
farther <>Nay V.Oli be seen as higher In ending associated V.Ot11 sl°""Y adaptlr>;J detection Into a private experlenoe. [1 J
the mage. [61 (SA Ill fibers tllat have large receptive sense of angular motion The per-
relatlve metncal depth cue fields. 1131 ceptual modality that senses rotation.
A depth cue that could specify, fcr
example. that object A Is tv.1ce as far s 11 21
away as object B wlt11out provldlr>;J S- cone A cone that fs preferentlalt>,t sense of linear motion The per-
lnfcrmatlcn about the absdute dlstarce sensitive to short wavelengths: cclloqul- ooptual modality that senses transtat k::n .
to elth€r A or B. 161 ally (but not entirely aocurateM krown 11 2]
as a 'blue cone." 151 sense of tilt The perceptual modal-
relative size A comparison of size
between Items without kn°""ng the saccade A fype of f!Y0 movement, ity that senses head lrcllnatlon with
absolute size of either one. 161 made both Vd untanly and lnvcluntarlly, respect to gravity. 11 21
In which the "fel3 rapidly change fixation sensitivity 1. The ability to respond
releaser pheromone A pherc:mcne from one ob)act or locatlon to another.
that triggers an Immediate behavioral to transmitted slgras. 2. In signal
181 detectloo thecry, a value th:tt defines
response among conspeclfics. [14]
saccadlc suppre ssion The the ease 1._th whk::h an observer can
response enhancement An effect reduction of vlsual sensitivity that tell the difference between the presence
of attention on the response of a neu- occ:urs wha"l we make saocadlc eye and absence of a stimulus or the differ-
ron In which the neuron responding to movements. Sacca.die suppression ence betV\fSen stimulus 1 and stimulus
an attended stimulus gives a bigger
el!mlnates th9 smear from retln:tl Image 2. 11 , 21
response. 171 motion durtng an "fe movement. [81 sensorineural hearing loss Hear-
retina A Ught-sensltlve membrane ln saccule One of the t>.m otolith ing loss due to defects In t11e cochlea cr
the back of the eye that contalrs roos agans. A saclike structure that con - auditory nerve. 191
and cones, which re:eive an Image tains the saooutar macula AJso called
from the lens and send It to the brain sensory exafference Change In
througl1 the optic nerve. 121
S8Cct//U8 ' I121 afference caused by external stlmull.
salience The vlvK1noos of a stimulus For the vestibular system, vestlWar
reUnltls plgmentosa (RP) Apro- afference evoked by passive head
relative tQ 'ts neig hbors. [7]
gresolve degeneration di th<i motion would )'leld sensory exafference.
that affects night vision and peripheral salty of the fOLr basic tastes; t11e
taste quality produced by the cations Compare senscvy reefference. 11 21
vlsbn. RP commonly rurs In farnllles
and can be caused by defects h a of sallts (e.g., the sodium In oodlurn sensory Integration The process
number of dnerent genes ttJat have chloride produoes the ealfy taste). of combining different sensory
reoontly been ldentlned. 121 Some cations also proooce other taste Typically. com bining several signals
qualities (e.g., potassium tastes bltt€f yietls more accurate and/or more pre-
re1ronasal olfaction PercelvlrYJ as wen as ealty). The purest salty taste cise Information than can be obtained
odors tllrough your mouth while breath- is prod uced by sodium chlonde (NaCl), from individual sensory signals. This is
lr>;J and chewing . This occurs when we common table salt. I151 rlOi the mat11ematlcal proooss of Inte-
are smelling somethi ng that Is Inside our gration learned In calculus (e .g. 1 t11e
mouth an::l ls v11hat gives us the e.xperl- scatter To disperse something -
such as light-In an Irregular fashion. 12] Integral of aocel8'atlon Is velocity). 1121
enoe of flavor. 1141
scene-based guidance lnfcrma- sensory reafference Change In
retronasal olfactory sensation afferenoe caused by self-generated
The sersation of an odor that Is per- tlon In our urderstandlrg of scenes that
helJOS us find speci e objects In scenes actMty. For the vestibular system. ves-
oelved when chewlr>;J and swallow- tibular afferenoe evcked by an active
lr>;J force an odorant In the mcuth up (e.g .. objects do not float In air. fauoets
are near sinks). [7] -generated head motion would )'leld

508 GLOSSARY
sensory reafferenoo. Compare sensory sine wave or pure tone The wave- spatial orientation A soose ro1-
exafference. [12] fam for wl1icl1 varlatlcn as a tunctlcn of slstlng of three Interacting modalities:
serial self-termlnaUng search time Is a sine f]..!nctlon. [9] peroeptlm of linear motion , argular
A search trom Item to Item, ending sine wave grating A grating with a motion. and tin . [1 2]
when a target Is found. [7] sinusoidal rumlnarce profile as shown in spatial-frequency channel
set size The number of !terns In a Figure 3.4b. [3] A pattern analyzer, Implemented by an
Visual display. 171 single-opponent cell Another way ensemble of cortical neurons, In 'Aitllch
to refer to OCfle-opp::nent cells. In order each set of neurons Is tuned to a limited
shape.-pattem theory lhe current range of spatial trequendes. [3]
dcmlnant biochemical theory for how to differentiate them frcm double-oppo-
chemicals ccme to be perceived as nent cells. [5] specific anosmla 1he Inability to
sped! c odors. Shape-pattern theory sinusoidal Referring to any oocllla- smell one specific compound amid oth·
conten::Js that different scents- as tlon. SL.Ch as a so.Jrd wwe CT rotaticnal erwlse ramal smell peroeptlcn. [141
a functic:n of the flt between odorarit motion. v.ih::t>e waveform Is that of a specific hungers theory The idea
shape to OR shape- activate differ- sine curve. The period of a sinusoidal that delclency of a given nutrient pro-
ent arrays of dfactory receptcrs In osci llation Is the time that rt takes for dL.Ces aavin-J {a si:;eciflc hunger) for that
the olfactory eplthelia. ll'1ese varklus one fUll back-and-forth cycle of the nutnent. Curt Richter ! rst proposed this
arrays produce spedl c !nng patterns motion to occur. The frequency of a theory and demonstrated that cra\lings
of neurons In the olfactory bulb, which sinusoidal osclllatl:::n Is de! ned as the for salty or for sweet are associated wnh
then determine the partlrular scent we numeral 1 dM::fed by the period. [1 2] de!clemloo In thooo substanoes. How-
parceve. [141 smooth pursuit A of volu ntaty """· the idea prwed '"'ong for other
sharper tuning An effect of attention movement in which the eyes move nutnents (e.g .. Vitamins). [1 5]
en the response of a neuron In wtilch smoothly to follow a moving obJeot. [8] spectral power clstributlon The
the neurcn responding to an attended physical energy In a light as a !Unction
somatosensation Cdlectlvely,
stlmulus responds mcra predsety. For sensory signals from the skin. muscles. of wavelength. [5]
example, a neuron that responds to tendons, joints, and Internal receptas. spectral reflectance function The
lines VY!th or1entatlons from -20 degrees [1 3] percentage of a partlcular wavelength
to +20 degrees might come to respord that Is reflected trom a surface. [5]
to ± 10-degree llnea. [7] somatosensory area 1 (51) The
primary receVlng area for touch in the spectral sensltlVity Referlng to the
signal detection theory A psy- cortex. [1 3] sensltMty of a ca l or a device to differ-
chcphyslcal theory that quantrnea the ent wavelengths on the electrornagnetlc
reSfXAISe of an observer to the pre- somatosensory area 2 (52) The
secondary receiving area for tou::::h in speotrurn. [5]
sentation of a signal In the presence of
noise. Measures obtained from a series the cortex. [13] spectrogram In souro analysts, a
of presentations are sensitivity Id') and somatotoplc Spatially mapped in three-dimensional display that plots
criterion of the observer. [1] the somatoeensory cortex ln correspon- time on the tulzontal axis, frequency
denoe to spatial events on the sl<ln. [1 3] on the vertical axis, and ampUtude
similarity A Gestalt grouping rule Ontenslty) on a color or gray scale. [11]
statln;i that the tendency of two fea- somatocyplcal Having normal
tures to group together will Increase as spectrum A representation of the
sornatooormtlon. [1 3]
the be™""1 them Increases. relative erl9fgy Qntenslty) present at
[4] sour One of the four basic tastes; the each frequer):))'. [9]
taste quality produced by the hydrogen
simple cell A cortlcal neuron wtose Ion In acids. [1 5] splnothalamic pathway The route
receptive field has defined excit- trom the spinal cord to the brain that
source segregation or auditory carries most of the lnfarnatlon about
atory and Inhibitory regions. [3]
scene analysis Processing an aLdi· skin temperature and pain. [1 3]
simultagnosla An Inability to per- tory soene consisting of multiple scund
ceive more than one object at a time. sources Into separate sound Images. staircase met11od A psychophysical
Slmultagrl061a Is a cor-.sequence of method for determining the ocnc.entra-
[10]
bilateral damage to the parietal lcbes tlon of a stlmulus required for detection
(Balint syndrcme). [7] spatial disorientation Any Impair- at the throoh:::ld lava. ll'1e staircase
ment of spatial or1aitatlcn. Me<e spedft- method ls an example of a method
sine wave A simple, smoothly cally, any Impairment of our sense of ofllmlts. A stimulus (e.g .. ode<ant) Is
changing ooclllatlcn that repeats across linear motion. an;;iular motion. or tilt. [12] presented In an ascerdlng concentra-
space. Hlgl1erlfe)'.1uemyslflllw'""'s tion sequence until detectla1 ls lndl·
spatial frequency The number of
have more osc:llatbns and lo.vef tre- cated. and then the concentration Is
quendes haVe f.......- oscillations CNer a cycles of a grating (e.g.. d811< and bright
bars) per unit of visual angle (usually shifted to a descerding sequence untU
given distance. 1. In hearing, a wave-
form for which as a fLnCtlon of speclfted n cycles per degree). [1 , 3] the response changoo to ...no detec-
tkln." lhls ascending and descen::Jlng
time Is a sine 1Ll'JC11on. Also caled pure spatial layout The description of tlie sequence Is fypicatly repeated several
tone. 2. In \lislon, a pattern for v..tllch structure of a scene (e.g.. erclosed. tlmoo. and the concentratklns at wl1lch
vanatlrn In a property like brightness open. rough, smooth) without referenoe reversals occur are averaged to deter-
CT color as a function of space Is a sine to the Identity of specrnc objects In tile mine the threshold detectlcn level of
function. [11 scene. 171 that odor.mt fcr a gwen ln::JMdual. Also
catled reversestalrcasemeimd. [14]

GLOSSARY 509
stapedius Tha muscle attached subtrac tion method In fl.motion syna ptic termlna I ll1e location
to the stapes: tensing the staped lus magnetic Imag ing. brain activity Is where axons terminate at the synapse
decreases vibration. 191 measured in two conditlcns: one with fcr tranemlsslon of Information by the
stapes One of the three osslcles. and one without the Involvement of the release of a chernlca tranem ltter. 121
cmnected to the lncus on one end, the mental prooess of Interest. SUbtraot!rYJ syncopation Afr>' from a
the two cond itions shows regions of
stapes presses against the oval window regcj1"' rhyttrn. 111 1
brain specf0oal ly activated by that pro-
of tlie coohlea on the other end. 191
cess. 141 T
stereoac uity A measure of the tactile a gnosla The lnablllty to Iden-
smallest blnooular disparity that can subtracttve color mixture A mix·
ture of pigments. If pigments A and B tify objects by touch. 113)
generate a sensation of depth. 16]
mix, some of the light shining on the target The goal of a visual search. 171
stereobllndness An lnablllty to surface will be subtracted by A. and
make use of binocular disparity as a some by B. Only the remainder contrib - tastant M Ystimulus that can be
depth cue. ll11s term Is typically used to utes to the perception of color. 151 tasted. 11 5]
describe lndMduals with vision In both taste Sensal lons evoked by solutions
superior colliculus A structure In
eyes. Some::::ne who has lost one (or In the mouth that oontact receptors on
the mldbraln that Is Important In Initiat -
eyes Is not typlcally described as the tmgue and the roof of the mouth
'stereoblind.• 161 ing and gcndlng eye movements. [8)
that then connect to axons in cranial
stereocl lium Arrf of the hairlike superior olive An early brain stem nerves VII, IX. and x.
11 5)
extensions on the tips of hair cells In the regicn in the auditory pathway wrare
lnpcns frcrn beth ears converge. 191 taste bud A globular cluster of c ells
cochlea t hat. when l exed, Initiate the that has tt1e function of creatln;i neu-
release of neurotransm itters. 191 supertaste r Supertasters are those ral signals oomeyed to the brai n by
stereolsomers lsaners (molecules Individuals who experience the most the taste neives. Some of the cells In
Intense taste sensations: fcr some a taste bud have specialized <ites on
that can exist In different structural
stimuli, they are dramatically mae their apical prqectlons tl1at Interact with
forms) In which the spatial arrange-
ments of the atoms are mirror-I mage Intense than fa 1nedlurn tasters or non· taste stimuli. Some of the cells form
rotations of one another, li ke a right and tasters. SLJpertasters also tend to expe- synapses l'l;th taste nerve llbers. 11 5]
left hand. 114] rience mcre Intense oral burn and oral
touch sensations. A variety of factors taste receptor c ell A cell within
stereopsls The ability to use binoru- may contribute to this heightened per- the taste bud that contains sites on Its
lar disparity as a cue to depth. 161 ception, among the moo.t lrnrx..-tant ls apical projection that can Interact with
the density offunglforrn paplnae. 11 . 15] taste stimuli. ll1ese sites fall Into two
stereoscope A device for simultane- major categories: those Interacting w ith
present Ing one lmag e to one eye supporUng ce ll One of the three charged particles (e.g., sod ium ard
ard another Image to the other eye. types of cells In the olfactory eplthellum. hydrogen Ions), and those Interacting
Stereoocopes can be used to prooent Supportln;i cel ls provide metabolic and with speclllcchemlca structures. 11 51
dlchoptlc sti muli for stereopsls ard bln- physlca support for the d factory sen-
ocu lar rivairy. 16] taster (of PTC/PROP) An Individual
sory neurons. 11 41
born with cne or both dcmlnant alleles
Stevens' powe r law A principle suppression In vl<ion, t110 Inhibition fcr the TAS2R38 gene and at>e to taste
describing the relatlon!'/llp between of an unwanted Image. Suppression the compounds phen;1thlocarbamlde
stimulus and resulting sensation that oocurs in people with and propylthlouracil. PTCIPROP tasters
Sl'tfS the magnitude of subjective sen - strablsmus. 161 who also have a high deo<ity of flmgl -
sation Is prcportlonal to the stimulus fa m papillae are PROP supertasters.
magnitude raised to an exponent. 11 l
surroundedness A rule fa figure-
ground assignment stating that If 11 51
stimulus onset asynchrony (SOA) one region Is entlr<Jy Slirrounded by tau M Information In the cptlc now
The ti me between the onset of one another. It Is llk<Jy that the surrounded that could signal time to co lislon (TTC)
stimulus and the onset of another. 171 region Is the ngure. 141 without the necessity of estimating
strablsmus A misalignment of the sweet One of t he four basic tastes; either absdute dlstanoes cr rates. The
two eyes such that a <i ngle object In the taste quality produced by some ratio of the retinal Image size at any
spece Is Imaged on the fovea of one sugars. such as glucose. fructose. anJ mcrnent to t he rate at which the Image
eye and en a nonfoveal area of the sucrose. These three sugars are par- Is expanding is tau . and TIC Is propa-
other (turned) eye. 13. 6) ticularly blolajcally useful to us, and tlonal to tau. 181
structural description A descrlp· c.ur sweet roceptors are tuned to them. tectorlal membrane A gelati nous
tlon of an object In terms of the rature Some other ccmpounds (e.g., saccha- structure, attached oo one €n d, that
of Its constituent parts am the relation· nn, aspartame) are also sweet. 11 5] extends Into the middle cana of the
ships between those parts. 141 symmetry A rule for flgure-groum ear, fioatlng above inner hair cells and
structuralism A schOO of thou;Jht assig nment stating tl1at symmetncal touching outer hair cells. 191
believing that complex objects or per- reg bis are more likely to be seen as tempo The perceived speed of t he
captions could be un:Jerstood by analy- llgure. 141 presentation of scunds. 1111
sis of the components. 141 synapse The jurciioo between neu- temporal code Tuni ng of different
substantla gelatlnosa A Jellylike ron s that permits Information transfer. parts of the cochlea to different fre-
region of Interconnecting neurcns In the 111 quencies. In whloh Information about
dorsal h:m of the spinal cord. 113] the particular frequency of an incoming

510 GLOSSARY
sound WENe Is coded by the timing of syllables, and so on. when In the tip-of- trochtear (IV) nerves The fourth
neural firing as It relates to the penod of the-nose state. llis Is an example of pair of cranial neNes, which Innervate
the sound. [91 how language and olfactcry perneptlm the supertcr oblique muscles of the
temporal Integration The process are deeply disconnected. [141 eyeballs. 111
by w hich a sound at a constant level Is tone chroma A SQ...lnd quality shared tufted cell The next layer of cells
perceved as being lcuder when It Is of by tones that hENe the same octENe after the Juxtagbrnerular neurons. They
greater duration. The term also applles interval. 1111 repom to fewer cdorants than the jux-
to percelVed bnghtness. wl1lch depends tone height A sound qualty cor- taglomerular cells, but more than neu-
on the duration of light. [91 responding to the leve of pitch. Tone rons at the deepest layer of ee ls. [141
tensor tympani The muscle height Is morotonlcally raated to fre- two-point touch threshold The
attachecl to the malleus; tensing the quenc,,,. [111 minimum dStance at y.,tJlch two stlmull
tensor tympani decreases vibration. [91 tonotoplc organization An (e.g., two simultaneous touches) are
texture gradient A depth a.10 based arrangement In which neurons that just perceptible as separate. [1 , 131
on the geometric fact that rtems of the respond to d ifferent freqU9flcles are two-tone suppression A decrease
same size form smaller lrnages 1At1en organized anatomicalti,t In order of fre· In the firing rate of one audltoiy nerve
they are farther CN/d'f. An array of Items quency. 191 fiber due to one tone, when a secom
that change In size smoothly acroos topographical mapping The tone Is presented at the same time. [91
the Image wlll appear to form a surface orderly mapping of the wcrld In the tympanlc canal One of three fiuld -
tilted In depth. [61 lateral genlculate nudeus and the visual illed passages In the cochlea. The
texture segmentation Garvlng an c ortax.131 "t)lmpanlc canal extends from the rcund
Image Into regions of common texture window at the base of the cochlea to
transduce To convert from one
properties. [41 the hellcotrema at the apex. Also called
fain of energy to another (e.g., frcrn
texture-defined object or contrast- light to neural electrical energy, CT from sea/a tympe.nl. 191
defined object k1 object that Is mechanical movement to neural electrl· tympanlc membrane The eardrum;
defined by differences hi contrast. or cal energY). [2, 121 a thin sheet of skin at the end of tl1e
texture, but not by lumlnanoe. [8] transmit To corweysornethlng (e.g .. outEf ear canal. The tympanlc mem-
thermoreceptor A senrory reoeptcr llghn from one place or thing to another. brane In response to SC<Jnd. 191
that signals informatlcn aba.Jt dlanges 121 u
In skin temperature. [1 31 transparent AllO'Mng light to pass umaml The taste sensation pro-
threshold tuning curve A graph throu.Jh with no Interruption, so that duced by monosodlurn glutamate. [1 51
plotting the throoholds of a neuron CT objects on the other side can be cearly uncrossed disparity The sign of
fiber In response to sine waves wit11 ' S<l'J0, 12) dlspanty created by objects b€hlnd the
var>1ng freqe»ncles at the lowes(tnten- triangle test A test In which a pw- plane of fixation (the horopter). The term
slty tl1at will give nse to a response. [91 tldpoot Is gt.fen three odorants to smell, uncrossed ts usecl because Images of
tilt aftereffect The perceptual Hlu- of wtich two are the same and one objects boated behind the horopter will
sbn of tilt. produoad by adaptation to a Is dfferent. The participant Is required to be dlsplaoad to the right In
pattern of a given CTlentatlon. [3, 6] to state "'1ch Is the odd odor out. the right eye, and to the left In the left
TyplcaNy, the crder In which the three 9)'0. [61
tilt To attain a sloped position like that
odorants are given (e.g., same, same,
of the Leanl ng Tower of Pisa. I121 unique hue k1V of four odCTs that
different: different. same. same; same. can be described "1th only a single
timbre The psych::loglcal sensation different. same) Is manipulated and
by which a listener can judge that Nvo cdor term : red, yellow, green, blue.
the test Is re-peated several times for Other colors (e.g .. pufllle or orange) can
sa...mds with the same loudness and greater accuracy. [141
pnch are dissimilar. Timbre quality Is be described as ocmpounds (reddish
conveyed by harmonics and other high trichromatic theory of color vision blue. reddish yellow). 151
frequencies. 19. 10] or trlchromacy The theory thal: the uniqueness constraint In stere-
c ol:::< of any light Is defired In our opsls, the observation that a feature In
time lo collision (TTC) The time system by the relatlonst1lps of three
requi red for a moving object (such as the work:1 Is represented once In
numbers-the a.rt.puts of three recep· each retinal Image. This constraint sim -
a cncket balOto hit a stationary object tor types nCJN known to be the three
(such as a batsman 's head). TIC= dis- plifies the correspondence problem. [61
cones. Also called the Young-Helmholtz
tance/rate. [81 theol)'. [51 unrelated color A cdCT that can be
tip link A tlrry filament that stretches expe1lenced In looatlon. 151
trigeminal (V) nerve The fifth cra-
frcrn the tip of a stereod llum to the side nial rerve, which transmits Information utrlcle One of the two otollth organs.
of Its neighbor. 191 about the "feel" of an odorant (e.g., A sacllke structure that contains the
mint feels CCQl, ci nnamon 1eels warm), utricular macula. Also called utrtcul.Js.
tip· Of· the-nose phenomenon The
Inability to name an odor, even thoogh as well as r:;ain am lrrltatlcn sensations [1 21
rt Is very familiar. Contrary to the tlp-of- (e.g .. ammonia feels burning). 1141 v
the·tongue phenorrar-01, me has no trltanope AA individual who suffers vanishing point The apparent point
lexical access to the name of the odor, from color blindness that Is due to the at which parallet lines receding in depth
sudl as first lettff . rhyme. number of absence of S -cones. 151 converge. 16]

GLOSSARY 511
vectlon An Illusory sense of self- vestibulocochlear (VIII) nerves vitreous humor The transparent
motloo caused by movi ng visual cues The eighth pair of c ranial nerves. w 111c11 fluid that Il ls the 'lltreous c hamber In the
when ooe is not, In fact, actually mov- connect the inner ear with the brain, post9flor pan of the eye. [2]
ing. [12] transmitting lrnputses concerned with vocal trac t The airway above the lar-
velocity The speed and d irection In hearing and spatial orlentatloo. The ves- )'T'O< used for the production of speech.
which scmethlng moves. Mathemati-
tlbulocochlear nerve Is composed o1the The vocal tract Inc ludes the oral tract
cochlear nerve branch and the vesti bu-
cally, velocity Is the Integral of accelera- and nasal tract. [11]
lar nerve branch. [1]
tloo. In words, linear velocity Is distance volley principle The Idea that mul-
divided by time to traverse th9! dis- vibration theory An altemat'"' to tiple neurons can pr0\'1de a temporal
tance: angular velocity Is rotation angle shape-pattern theoty fcr describing oode for fi'equerY;y If each neuroo fl res
divided by time to traverse th9! angle. how ol1actlon works. Vibration theory at a distinct point In the period o f a
[12] proposes that every odorant has a dlf- sound wave but does not fire oo every
ferant 'llbratlonaJ treq uency, and that
vergence A type of eye movem ent perbd. [91
In whk;h the t\•10 eyes move In opposite molecules trat produce tl1e same 'llbra-
tlorBI frequencies vJlll smBI the same. vomeronasal organ (VNO)
directions; for example, both eyes tum A chemical sensing org an at the base
toward the nose (convergence) or away [14]
o f the n asal ce'llty w ith a cur.'<ld tubu-
from the nose (divergence). [8] fl lier circle The location lar shape. The VNO evol\ied to detect
vertigo A sensation of rotation cr o f objects whose images fall on geo- chemicals that cannot ba processoo
splnnlrg. The term Is often used more metrioally corresponding pci nts In the by ORs, such as large ard/cr aqueous
generally to mean any fcrm of dizziness. two retinas. If 111e were sJ rnple, this clrcle rnclecules . the types of rnclecules that
[12] would be the horopter. but me Is not ocnstltute phercmmes. AJso called
si mple. [6] Jacobson's organ. [14]
vestibular canal Ole of three fluid-
v1ewpolnllnvarlance 1. A propeny
filled passages in the cochlea The
of an object that does rot change 'When w
vestlbUlar c anaJ extends trom the oval warmth fiber A sensory nerve
ctJserver vtewpci nt changes. 2. A c oos
window at the base of the cochlea to tiber that fires when skin temperature
o f theories o f ctJject reoognltloo that
the hellcotrema at the apex. Also caJled Increases. [1 3]
scala vestibul. [9] proposes representations of objects
that do rot change when viewpoint wave An osdHatbn that travels
vestibular organs The set of five c hanges. [4] through a medium by transferring
seroo organs -throo semicircular energy fi'orn one partide or point to
canals an::::I two otollth crgars -located vlsual a cuity A measure of the flnest
detall th9! can be resolved by the eyes. another w ithout causing any permanent
In each Inna' ear that sense head displacement of the medium. [2]
rnotlcn and head orientation with [2]
respect to gravity. See also vesYbular visual angle The angle subtended wavelength The distance req uired
system. It 2J by an object at the retina [3] for one full cycle of osclllatlrn for a sine
wave. [1]
vestibular system The set of flve visual crowding The deleterlous
seroo organs -throo semicircular effect of clutter on peripheral object rec- Weber fraction The constant of pro-
canals and two o tollth crgans -located ognmon. [3] port bna/lty In weber's law. [1]
In each Inner ear that sense head vlsual search Search fa a target Weber's law The prlnclple descrlb·
motlcn and head orientation with In a display contai ning distracti ng ele- Ing the relationship betwoon stimulus
respect to gravity and tt1e neural path- ments. [7] and resultlng sensation that says the
ways directly associated vAth these just noticeable d ifference (JND) Is a
sense organs. See also vestibular vlsual- fleld defect A ponion of the ooostant traction of the companson
organs. [1 2] 'llsual field with no vision cr w rth abnor- stimulus. [1]
mal vlslcn. typical ly resulting from dam-
vestlbulo- ocular reflex (VOR) A age to the '1sual nervous system. [7] white noise Noise consisting o f all
shon-latency re!ex that h elps stabilize audible frequencies In equal amounts.
vsbn by ccunterrotatlng t he eyes when vitallsm The Idea that there is a force White n oise In heanng Is analogous to
the vestlbular system senses head In life that Is distinct trorn physlcaJ enti- white light In vision . for w hlc l1 all w ave-
movement. [1 2] tles. [1] leng ths are present. [9]
8230336 Amma App.:1

4
' 8230336 Amma Appa

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lq
8230336 Amma Appa

Index
8230336 AmmJ App.l
Entries \<1.1 ith an ita1ic/ next to the page number indica te that the information will
be found in a figu re. Entties with an italic t next to the page number indiClte that
the information will be fou1'd In a table.
A adaptingstimu.l.i, 146 angular motion, 352--353

A"beta fibe1s Addams, Robert, 237, 238 angulaJ· velocity, 367/
desai pti.on of, 392 00.ditive color mixtmes, 129 anisometropia, 86
dorsal hom, 397/ <Kleno-assodated viral (A VV) vectot's, Aniston, Jennifer, 94
pain and, 402 50 anemia, 145
A-delta fibers, 395 oorial pmpectives, 165-166 ansomia, 431, 432-433
d.esaiption ot 394 afferent fiber s, Tl2 specific, 440
dorsal hom, 397/ afferent signals, 351 ante1for cinguli!.te cortex (ACC), 403
nod<Pption, 406 Afrlcan elephants, 430 aperture. 242-243. 242/
abduams nerve (CN VI), 19,f, 20 afterimages, color contrast. 146 apparent motion, 240-241, 240f
ahsolute mett'ical depth cues, 165 age/ aging aqueous humor, 33
absolute pitch (AP), 32W25 hearing and, 286, 300-301 Ard1il'frtmnl View (Martini)1 166-167,
ahsolute thresholds olfactio n and, 449 167f
50% definition ot, 8 pel'ceptttal learning ai1d, 86 area 17, 65
commonsense, 7t sm ell function and, 433 Aiis10tle, 47, 238, 349-350
d.efiaition of, 7 age-related macular degeneration aromatherapy, 462-464
absotption, 31 (AM D), 50 articulation, 330--332
acceleration, 356 agnosia Sre nlso prosopag nosia asparagus, 440
aocesso1·y olfactory bulb (AOB), 459 description of, 92, 145 aspartame, 479
accidental viewpoint, 105 tactile, 416 ostigmatism, 34, 34/
accommodation tem poral lobe lesio ns ai,d, 95 attack, sow1d, 307--308
aging and, 35f akinetopsia, 257-258 attention
desaiption of, 34, 171 Albeiti, Leon Batista, 166 activity coord ination and, 217
achromatopsia, l 40 Alzheimer's disell8"1, 450 audito1y, 317-318
acoustic cues, multiple, 338-339 amacriru! cells, 37f, 38, 40-43, 47 definition of. 201-202
aC01.LStic reflexes, 268 Tl1e A111lwSimfors (Holbe in), 168, 168/ physiological basis of, 215-219
acoustic startle reflex,. 317 ambiguous figures, 104, process ing of stimuli and,
action potentials, 22- 24 amblyopia, 85, 86 scene p"2'rception and_,201- 234
active Sc:!nsing, 351 A moore, Jolm, 443 single cells and, 2'1 7- 219, 217/
acuity amplitude "spotlight" of, 205
definition of, 55 cochlear roding of, 272- 'Il4 visual disorders, 219-222
touch, 405-410, 409/ o tolith ox!ed, 365 attention-.defid.t hyperactivity disorder
types of, 5St senUcit•ettlar can..Us and, 359-361 (ADHD), 221
Vemier,, 185 sound waves, 262 attentional blink, 212- 213,
visual, 43, 53-<;2, 58/ spatial orientation and, l53 4
adaptation anl pullae. 359 Aube1t, Heimann Rudolf. 369
cognitive habituation, 453--454 amusement park rides, 384-385 audibilty threshold, 282
color contrast, 146 amygdala-hipporampal complex, 434, auditory attention" 317- 318
oont1'ast sensitivity fut1ctions and. 6lf 457 auditory color constan cy, 306-307
to cLuk, 47-50, 4Sf analgesia, 404 auditory distance perceptions, 301- 302
definition of, 77 onainorphic projections, 168, 169,f auditory neurons
to light. 47- 50 amunorphos.is, 168 de&ription of, 269-270
olfactory identification and, 450-453 "AND gate,'' 239 physiology, 274-279
rates ot 391 and.rostadienone, 461 threshold tuning curves, 274
selective, 78f, 79f, 80-81, 81{, 412 androstenone , 440. 446 tuning curves, 2¥
spatial frequency and, 80 Mencephaly, 484-485 auditory scene ana.lysis. 308--314, 309
angular acceleration , l52 auditory stream s"gregation , 310

540 INDEX
auditory syst<>m, 265-281, 280f auditory struct\Jfes, 279-281 cold fibei., 394
autonomic nervous system, 377-378 cortex of, 20/ color-anomalous individuals, ·144
Axel, Richard, 436 cortical visual pathways, 54/ color assimilation, 145, 145/
azimuth, 293 limbic system, 435 color-matching techniques_. 128, 143
m essages from skin, 396-402. 398f color pe1·ception, 123-155
B motion-sensitive areas, 244{ individual diffenmces in, 140-145
Bach, Johann Sebastian, 310, 310{ speech in, 343-346
Bacon, Francis, 47 B1·e,,·ste1·, David, 176 three Sh>ps to, 123-123
balance system, 373-381 Bridgeman, B1'ttce, 178 color s paa!s, 132
Banks, Marty, 167 Brnnellesco, Filippo Di Ser, 166 color v ision
Barlow, Ho1'ace, 92 Buck, Linda_. 436 colo r blindness, l25, 143-144
Barr, TI10mas, 286 Billthoff, Heiruich, 115 genetic differe nces in, 143-145
Barty, Sus an, 178 butanol, 453 l!Sefulness of, 151-155, 152,1. 153f
basal cells,. olfacto1y system,. 429 colors
basicrolorterms, 141-143 c appea1a11ces, 132-134
basic tast<>S, 47'>-480 C fibers. 394, 395, 397f constancy, 147-151
basilar: me1nbmnes, 268 C tactile (CD afferent neurons, 395, 401 14.S, 145J
Bayes, ll'°mas, 110, 186-167 Cajal, Santiago Ram6n y, 21-22, 21/ dlsctiminatlon of, 124-132
Bayesian approach, 110, 186 Ca.lifomia gn:rund. squirrels, 456, 457/ picking of, 133-134
Beauchamp, C.'1y, 476 camouflage, 103-104, 103f relat<>d, 146
Beethoven, Ludwig van. 324 Gimpbell, Fergus, 59, 81 wirelated, 146
Behnnatu1, Marlene, 220 cancer patients colum ns
belt aJ'eas, 281 painful mouths in, 491 CO blots, 76, 77f
benz.lldehyde, 446 taste sensations in, 4i'5 description of, 74-77
Betinmo populations, New Guinea, 142 capsaidn, 20, 491 committees
bifocals, irnrention of, 35 carbon monoxide, 428/ multiple recognition, 116-117
binaural 1·ecording, 300 carrageenai> 469 perceptual, 104
birl.:1tual ri\•a.hy. 443 D-caivone, 441 comparators, 255, 255/
bh1ding problems, 211, 2llf cataracts, 35 complex oolls, 73, 73f
bhlOCU!ar depth cues, 160 cJtegot'ic.al pera!ption, 335-337, 335f complex sowlds
binocular disparity, HlO change blindness, 227-229, 229-233 restomtion of, 315-316
iniormation, 188 frequendos (CF), 274, 276 spect1'ogram, 343/
stereopsis and, 172f Otaudha.ri, Nirnpa, 48i comput«l tomogrnphy (CT), 27, 27/
bu)()CtllM timlry, 190-191, 191/, 192f. cheese, smell of. 456 conditioned food preforences, 486-487
196-197 chernosignals, 459-462, 461 conductive hearing loss, 285
binocular stereopsis, 181-182 chemotherapy, taste sensations and, 475 cone monochroma.ts, 144
binocular summation, 159 chili peppe15, 490--491 cone-opponent cells
binocular vision, 159 chinchillas, 336{ lateral genkulate 131,
development of, 192-19'7 Chlll<?so characters, 208-209, 20Sf 138-140
disruption of, 195 Chino_. Yuzo, 194 retinal, 131
biological motiot\ 248, 249/, 250{ chloride deficiency, 476 See n!so L-cones;
bipolar cells chocolate, 442 5-cones
diffuse, 43 chorda tympatti, 468, 471/, 483 operation of, 40
function of, 43, 47 chords1 musical, 323 retinal, 41/
information, 42--43 clu·omophores. 40 sensitivity, 49
midgot, 43 Chui> Manin, 213, 214f cones of confusion. '297
OFF-<enter, 43. 45{ cilia, olfactory, 429 congenital prosopagnosia, 118
ON-=ter, 43, 45( circadian clocks, 131 conjunction s.ea1'Ches, 210
birds.. recognition of, 116-117 circumvallate papillae, 472 consonants
bitter Mstes, 473, 475, 477-478 Citralva, 446 acoustic differences, '33if
Blaken\01·e, Colin, 77 dtr1.1s 01·de1·s, 442 dental-stop, 340{
Blakeslee. Albett, 480 principle of1 100 English, 332,1
blind spots. 36f coartirulation preceding vowels and, 337f
blood oxygen level-dependent (BOLD) consonant sounds, 337/ voicing, 333
signals, 28, 68 dclinition of. 333-334 constant stimuli methocl, &-9, Sf
blood pressure. influences on, 378/ itwad.ance and, 334 context sensitivity, spetich, 334
Blundell, John, 4BO spectrnl contrast and, 337-338 continuity constraint, 184
bcdy imagos, 400-401 , 400f cochll?a, 268 continuity effects, 314-315
Boltot\ Thaddet1s, 326 anatomy of, 269/, 273f contoW'S, illurory, 98, 98/
BoiulOt de Condillac, E.tiem'le. 3-4, 4f coding, 272-274 contt·alateral, definition of, 64
borders, shaip, 150 in1pla11ts, 28.s, 21fJf, 346 rontraleslon..."'1 fields, 220
boundaries, object_. 109f cochlear nuclei, 279 contrast, 46, 54, 60
Braille patterns, 401, 400 cochlear partitions, 26S contt-ast-defined objects, 246
brain cognJ.tlvedemons, 104 sensitivity
attention and, 215, 215f cognitive habituation, 453-454 development of, 84-85

INDEX 541
SlMtial vision and, 6 lf roding in semicircular ca.na.l.s, 361 European starlings, 311V
contrast sensitivity ftmct:ion.s (CSF), otolith coded, 365-366 event-related potential (ERP), 25-26, 26/
60-61, 60J, 84-BS spatial 01':i.entation and, 353-355 4?vidence, rules of, 98-100
contrast th1'esholds, 60 directional transfer functions (DTFs),
convel'gence, 17L 'mf,300 color vision, 151, 153
com.ea,32 dimirninability (d'), 315 Darwin's theory of, 18
coirespondence, problems with, 182, distances. 368-369 eci.uilibrium and, 351
241-242. 24'\f distractors, 205 howing, 261, 268
corresponding retinal points, 173, 173/ di\'ergency, 171 olfactory, 45&-458
C01'teX divided attention, 202 speech, 328
homologous regions, 93 dizzi.ness. 349. 367 toucl\.398
macaque monkey, 9Qf, 9'3f doctrine of .specific nerve energies, 1S vertebrate, 351
topography of, 67-68 dogs vestibular system, 349, 35'1
visual 90f, 91f olfactory sensoty new·ons. 431 excentt':i.city, 38
cortical magnification, 68 \'ision of, 153 Exner, Sigmund.. 240
letti?t chart ;md, Wf dorsal col wnn-medial lemniscal E?Xogenous cues, 203
perceph1al consequences, 68-70 (DCML) pathway, 398, '!BSf 4?Xogenous spatial attention, 419
cowue1it'fitation, 404 dorsal horn, spinal oord, 397, 397/ exotropia, 196
covert attcintion, 202 double dissociation.. 118, 247 &ploratory procedw·es, 410
c1·anial nerves, 18, 19-20, 19/, 468 double-0pponent c.lls, 139, 139/ @eternal attention, 201
cribriform plate, 20, 431 Dra yna, Dermis, 481 extinction, 220-221
c1i.stae, 359 dreams, olfactory sensations in, 445 extra.striate body area (EBA), 113
c1ite1ia. response levels and, 13 dualism, philMOphy of, 5 @Ctrastriate cortex, 89
crit:ic.i.l. bandwidths, 284-295 duck.rabbits, 105 eye movements, 250-257
critical periods,85, 195-197 Duffy, Valerie, 482 rompensation. 256-257
c1'0.SS-dd..1ptation. 452. 490 duplex, definition of, 3B physiology oi, 253-254
cross-modality matclting, 11, llf, 482 duplex retina,. 49 l'Qflexive 254
crossed dispality, 175, 175/ eyeballs, shape of, 34
cry•stallins.35 E eyes
cues ear canals, 266 accommodation, 171
definition of, 203 ears, anatomy of, 266-277, 267J, 356[ See an.1tomy of, 33/
development of, 204 alw heat'ing blind spots, 36f
from haze, 166/ edges 171
motion, 169-171 figtt1'e-grow1d assignment and,, 107, dive:rge?nce, 171
olfacto1y, 462-464 lfll/ frontal, 15'l
sound locallz,,tion and, 297-298 occlusion, 101 fundus of, 36f
cultutal relativism. 141 recognition of, 97- 101, 9Sf light mpture, 32-40
Cutting, James, 249 sharp borders, 150 muscles, 252f
cycles, 55 effei·ent conu1mnds, 351 optics of, 311
cycles per degree, 17, .59 efferent fibers, 'Zl3
cyclopean, 180 efficiency, feah11-e seal.'ches, 207- 20S F
cytochrome oxidase (CO), 76, 77f egocenter, 417, FA I fibers, 393
(EEG), 24, 25f FA Il fibers, 393, 406, 412
D electronic ea1·s, 28S-289 fudal 337
Dani populations, New Guinea. 142 enunetmpia, 33 facial nerve (CN VIQ. 1'!f, 20
dark adaptation curves, 4Bf emotional touch. 395-396 facial recognition, l17-118, 117/
Davidson, Terry, 480 emotions, music and.1 325 fam.iliru· sizes, 165
Davis, Clara. 485-4S6 end stopping, 73 fanchat't:,
inattentional, 317 endocannab:inoid receptors, 437 Fast, Kathetine, 482
decay. sound, 307-308 endogenous a.ms, 204 fat, dietary, 488
decibels (dBs), 263, 2631 endogenous opiates; 404 fatty stimuli, 476
dental-stop consonants. 340/ endogenous spatial attention, 419 feature integration theory, 211
depth cues, 161, 164J, 167. 186-192 endolymph,. 359 fuatw·9 searches, 2fJ7
dermis, anatomy of, 300 Etu·oth-Cug4?ll, Christina, 62 Fechner. Gustav, S, 5/
Descartes, 72 ensemble statistics, 'lXJf Fechner's, law,6-7, 7f, 11
TI1e Descent of Man (Darwin}. 18 entorhinal rortex, 435 fued.fol'ward process4?s, 95
detection tlu·esholds 446--447 entry.level category, 117 l<>edback processing, 118-119
deuteranope individuals.. 144 environment, sounds in, 264 figu1"e-growid assignm ents, 106-100,
dichoptic, definition of, 1\4 epidemtis, a.n.atom.y of. 390 107/
diets, specific htmgers and, 485-487 equal-loudness curves, 282 figures, gmund atld., 106-108
difference thresholds, 6 equilibl'ium, sense of, 349-386 filifom\ papillae, 471
diffuse b:ipolar cells, 43 Escher, M_ C., 160f filters, defi1tition of, 46, 72
diopters (D), 33 esotmpia, 196 finge1pril1ts, 412-413
dip lopia, 173 Euclidean, desc1iption of. 157 fingers, skin ridges, 412{
direction Euclidean grometry, 15S/ fingertips. touch and. 415/

542 INDEX
firefl y 154 colot' vision and, 143-145 Hayes, John, 482

first-ord er motion, 246 touch sensations and, 40.5-406 haz.es, cu es from , 166f
Fisd wr, Roland, 481 voice and, 329--330, 339 "head directio n cells/ ' 381
Fixi11gMyGm(Ban y), 178 generative models, 111 heaii ng, 261-289
flavo rs geneti cs, color vision, 143- 145 absolute thresholds, 7t
localizing the sens ation of, 468-469 geons, 114-115, 1lSf effects of noise on,. 287/
taste L'ef.SflS, 467-470 Gestalt grouping rules in the environment, 291-319
volatile-<mhancod taste and, 469-470 definition of. 00 function of, 261
Flegl, Katherine, 480 good continuation principle, 100 human t'ange, 263
fl oate1·s, description ot 33 law of common fatl?, 312-313 heali ng lass, 384
Flourens, Jean Pi erre, 37S by onset, 312- 313 conducti\•e, 285
focus o f expansion,. 248 parallelism, 102, l !llf sensori.neural. 286
foliate papillae, 472 proximity, 102, 102f treatment of, 237- 288
food additii,•es, 469 similaiity, 102, 102/, 311 hedonics, o lfactory .. 454--458
formant, 330 symmetty, 102, l !ll/ Heeger, David, 218, 246
Foul'ier, Joseph, 15f Gestalt theory helicon:ema, 268
Follt'ier analysis, 1>-18 d efiniti on ot 00, l OOf Helmholtz, Hermarm Ludwig Ferdi-
description,. 17 p1ind ple of good continuation, 314 nar1d von, 20, 21f , 12S, 256, 489
of m otion. 363 gestation, o lfacto1y system, 455 heptic virtual environments, 423, 423/
sine wave, 62f Gibson, J.]., 247 He1i ng, Ewald, 135
fovea C ilad, Yoov, 4.36 H ering illusions, 189, 190f
functio n of, 38 Gilbertson,. Tim, 488 hertz (Hz), 263
im ages on, 172 Giorgio Maitini, Francesco di, 166 heterodimet10, 479
p1'operties of, 38t glabrous skin, 392 hew·istic, definition of, 108
Fox, Althu1·, 480 global superiority effect, 109 high-spontaneous 6bel's , 277
fram es of reference, 417 glom eruli, 434 Hodgkin, AIan, 23, 'l3f
Franklin, 35 glos.sopharyngeal nerve (CN lX), 19/, 20, Hoffman, Don, 109
Frantz, Robert, 83 471f, 483 Holbein, Hans, 16.I
free fusion, 176 gl ucose, structure of, 478J Holmes, Oliver Wendell, 176
freesla, 441 gl utamate. ne motransmission, 487--488 H olmes ste1·eoscope, 176/
freq_ue:ndes Goddard, George W., 181 homologous regions.. 93, 'l!if
cochlear coding of, 272- 274 good con tinuation, pl'inciple of, 100 homunculus
sound, 2811 283-285 graded p:>tentials, 41 definition of, 399
sOUfld waves, 262 "grandmother cells," 92 sensoiy, 399-400, 399f
Frey, von, 405 granule cells, 435 H ooke, Robert, 57
Flisby, Jorn\ 77, 176 gratings horizontal cells, 42, 47
frontal eye fields (FEF), 257 com pound patterns, S2f horopte1·s , 173
fni ctose, 47!¥" Michelson de£initio11,. 60 HSB color spaoos, 133-134
fru it flies, olfaction in. 441, 446 gravity, m otion sensing and, 353 Hubel, David, 70-76
functional magnetic resonance imaglllg " gl'eebl"'," 116, 116f hue 134f , 135f
(IMRI), 28, 69f C!'een, Dan, 59 hues, unique, 136
fundamental frequencies, 264, 303, 304/ Greene, Michelle, 224 H UGO Gene Nome nclat1.ue Committee,
lundus, 36f, 37 C reer, Charles, 430 477
fungifo m \ papillae C1·oss, Chatlie, 92 Hlunan Cenonw Project, 4'17
chilibumand, 491 gl'ound1flgures and, 106-108 Httr\'ich,. Leo. 136
description of, 471-472 guar gum, 469 Huxley, Andrew, 23, '2:3/
in superta.sters , 482 guid ed searches, 209-210 hyper.acuity_, 185
v ariability in, 472- 473, 472f Guifrntss Wotld Rerords 2005, 59 hyperalge!!ia, 405
fusiform foe< area (FFA), 94-95, 113, 216, gustation, definition of, 427 hype1:coluir1.ns, 74-77. 76f
216f gustatory a>rhlX, 47 4f hyperromplex cells, 73
gustato1y S)'Slt>Ol, hypernpia, 34
G hyperp olarization, 41
C protein-coupled rec('ptors (CPCR), H hypnosis, PET signals and, 403
,..
ganglion cells
4i9, 479f
habituation, cogniti\'e, 453--454
hair cells, 270-272
d epolari zation o f, 357
hypoglossal nerve (CN XII), 19f, 20
hy potheses, m odel a·e.a tion and, 110
fw'lctio n of, 4l-47 description of, 269-270

equiliblium and, 356-JSS, 357/ illuminants, 148f
P, 43, 4v, 47 heruing loss and, 286 problems with, 149
photomicrographs, 37/ hamsters, olfactory system, 459/ illusions
receptive fields, 45f Hmidbook of Physiology (Mitller), l S construction of space and, l BS-100
responses to g mtings, 63 H ansen's disease, 395 waterfall, 237
retinal, 62-64 haptic perception, 41 0-422, 414-415, 418/ illusoty conjunctions, 212, 212f
gate oontrol theo1y , 402 harmonic sow1ds, 265f, 303, 30'lf illuso1yrontours.• 98, 98/, 101f
gender harmonic spectra , 264 images

8230336 Anma Appa INDEX 543
d!?finition of, 32 Kohlel', Wolfgang, 99 colot' signals, 137-138

reconstructions, 82f koniocellularcells, 44, 64 defoos, 144
IMAX movies, 370 koni()(ellular layers, 13 1 photopigments, 152
ilnbalances, d!?finition of. 367 Kuffler, Stephen, 44 photoreceptol'S, 154
inattentional blindness, 231-232 response of, 126, l'llf
incus, 266 L M ganglion cells, 43, 44f, 47
infants. Stt also M\"'boms L..cones.124 Maasai peoples, 431
lea.ming to listen, 339-340 oolor signals, 137- 13B Mach bands, 46
leaming words, 34()..342, 34lf defects, 144 macronutt'ients, 486
olfactiot\ 455 photopigments, 152 macul<1£!, 363
premature, 393 photoroceptors.. 154 magru;>tic resonance imaging (NIRJ), 27,
response to touch_. 393 t'esponse of, 126, 127f 27/
inferior colliculus, 280 labeled lines magMtoencephalography (MEG), 26-27,
infet'OtemporaJ. on cortex, 92, 9'3f desc1iption of. '397 2hf
inhibition of return, 204-205 itch transmission, 40'V magnitude estimation, 9, 1Qf
inne1· ea1·, 268 pain transmission, 404{ magnocollulat· layer, 43, 64
ilu"ter hair cells. 270--m taste newons, 489 main olfactory bulb (MOB), 459
inner segments, photoreceptor, 40 language mal de debarquement syndrome,
insular cortex, 474 basic color torms, 141-143, 142f 383-3114
intensity, sow1d, 262, 281, 282- 283 brain pt'Ocess.ing of, 342-346 malleus, 266
interaural level di.fferences (ILDs) le.unlng to listen, 339-340 mammals, vestibular system, 356-366
desai ption of, 295- 296 learning words, 340-342 mammograms, 1'.lf, 181, 18'1/
frequency diffor!?nces and, '196f tenns for smells. 448 MMgolskee, Robe11, 478, 4B7
physiology of, 296 latetal geniculate nucleus (LGN), 43 marijuana, 437
sound localiz.ation and, dfiatomy, 64f l\farine Biological Association.. 22-23
interaural time differences (ITDs), 292- oolorin, 138-140 masking expetiments, 284
295, 293f, 294f cone-opponent cells, 131-140 mate.rial pel'ception, 111-112
intem..=tl attention, 201 - 202 function of, 64-65 materialism, philosophy of, 5
Interocular phase difference lateral inhibition, 42 mathematica.ll y 368
195/ lateral superior olive (LSO), 296 Maxwell, JamesC!et'k, 128-129
intet'ocular transfor, 245- 246 learned taste a\•erslons, 434-415, 457 McClh1tock, l\la!tha, 460
lntrnub, Helene, '217 Lee, D. N., 250 lt.•kClintock eff ect, 460
inverse-square la\\', 301 foft s ttpedor temporal gynt.s, 344 McCw·k effect, 345
ipsilateral, definition of, 64, 433 lenses, eye, 33. 35 McK.. , john, 192
ipsilesional fields, 221 lentirular printing, 181 mechanoeleehical transduction (MET),
itis. eye, 33 leprosy, 395 271
is.obutylmethm.ypyrazine, 42Sf Lettvin, Jerry.. 92 mechanoreceptors
isoi.ntensity curves, "05, 276f Lei.-in, Dan.. 229 description of, 392
isovaleric add, 441 Udocaine, 468-469 equilibrium and, 356
itch stimuli, 403, 404,f light hu1ction of,. 393t
physics of, 31-32 chatacteristics of, 392
J visible spectrum, 3'lf sensitivity of,
Jahai proples, 448 limbic systern, 4.35 medial geniculate nuclet.LS, 280
James, William, 83, 215 line junctions. 108 medial nucleus of the trapezoid body
Jameson, Dorothea, 136 lineai· acceleration. 353 (MNTB), 296
Japanese quail, 336f line..:u· motion, 352, 353 medial s upel'ior olive (Jl.·150), 294
Jo hansson,. Ctuu1ar, 24&-249 lineat· perspectives, 166, 166f l\<feissner corpuscles, 391f, 392
John•on, K 0 ., 393, 411 346 melanopsin, 40, 132
]olicoem·, Pierre, 116--117 listening melody
jw1ctions, 108 teaming, 339- 340 definition of_. 326
just notia?able diffe1·ences OND), 6 tasks of, 3l4f development of. 327-328
juxtaglomerular JH?tu·ons, 435 Locke, Johi1, 140 memories
''locked-in syndrome," 445 for objects, 226-2'll
K Loew!, Otto, 22, 2'Jf odor-evok!rl, 462-464
Kamba peoples, 4.11 Logotheti.s, Nikos, 92- 93 for scenes, '11.6-2'0
Kanizsa, Gaetano, 98 lo1·dosis, 4€:D of smells, 447, 447/
Kan.izsa figures , 98, 101, 105 loudness Me:niere'.s syndrome, 384
Kellman, Phillip, 108 definition of, 263 mensttual cycles, 461
kinesthesia, 389 sensation of, 7 menthol, action of, 20
kinesthetic, definition of, 351 sound, 2Sl, 282- 283 fi.•1erkel cell neutite complexes_. 392
kinesthetic perception, 396 low--spcnt..'\lleotLS fibers, 277 Merkel cells, 39lf
kinocilia, 359 ltunina.nce-Oefined objects, 246 metamers, 127-128
Kirchnel', Holle, 223 methanol, sb,icture of, 428/
K.it..wka, Akiyo shi, 238 M method of adjustment, 9
Kaffka, Kurt, 99 M-cones, 124 method of limits_. 9, 9f

544 INDEX
methyl salicylate (wintergre-en), 4S8 nasal dominance, 429 odoratlts

meh'i.cal depth cues, 161, 165/ 456, 456f definition of, 427
Meyel'hoi, Wolfga>ig, 477 information in, pleasantness ratings, 455f
mi.crosaccades, 253 sensitivity to, 446
micro\•illi, 472-473 Navon, David. 109 strttctures of, 428/
mid-spontaneous fibel's, 277 Necker cube, 104, 1()1/ odors
middle canal, 268 negative afterimages, 146, 147f defutition of, 427
middle ear, 266-268, 2115-286 neglect, visual field, 219, 220f discrimination of, 446
middle temporal (lvlT) area, 185,. 186, neural network m.odels, 339 id.entifi.cation of, 448-449
n.:>ural plasticity, 402 negative, 452
development of, 257 tll?ttral points, 146 OFF bipolal' cells, 43
middle (mid.level) vision, 97-1 10 tll?Ufoimaging, 5, 24-29 OFF.center bipola.t· cells, 45f 46
mlddle-vision committees, 105 neuronal connections. 21-22 oliactton, 427-465
midget bifOla.r cells, 43 neurons cognitive habituation, 4.Sl-454
Milton, John, 104 excitatory, 22 cross -adaptation, 452
minimum disa'iminable acuity. 58-59 filing of, 22-24, 23_{, 44, 77, 276, 277/, definition of, 427
minimum recognizable acuity, 58 360-361 detection thresholds, 446-448
minimum resolvable acuity, 58 inhibitory, 22 disaimination, 447-448
minimum visual acuity, 57-SS measuring activity of, 24f evolution of, 456-458
mltral cells, 435 ttuling, 185/ individual di.ffet-ences, 449-450
models. formal, 110 nrurotrn.nsmi.ss.ion, 487-488 neurophysiology of, 4l3-43S
Molyneaux, William, 402. tlru.fOh'ansmitters, 22. physiology of, 427-432
monocular depth cues, 'J 60 newboms, retronasal olfactory sen...q,,ations, 467
monocular vision.. 160 Newsome and Pare paradigm , 244{ o !factory affect, 48S
monofilament tests, 405-409 ..·tot\. Isaac, 128 olfactory appatatus, 428-432
monopotassiwn glutamate, 474 Nilsen, Mary Ellen, 400 olfactory bulb, 430, 433
monosodium glutamate (MSC), 487, 488 Nimoy, Leonard, 300 olfactory cleft, 429
mood, music and, 325 Nobel Prizes oaactory epithelium, 429, 429/. 468f
morning sickness, 476 Alan Hodgkin, 23, 23/ oliactory nerve (CN Q, 19{, 20, 433
motion Huxley, 23, 23/ olfactory perception , theoJ'ies of, 439-441
a!tmffects, 245-246 Charles Sherring:ton, 21, 'l:lf olfactory receptors, 436-438, 483
biological, 248 David Hub.I, 70-71 olfactory sensory neurons (OS:Ns)
eye movements, 250-257 Otto Loewi, 22, 22f action potentials, 430
first-order, 246 Torsten Wiesel, 70-71 olfacto1y system, 429, 430/
global detecto1s, 243/ nociceptors. 394-195 protective barrier, 435-4.36
perception of, 'lSl noise olfactory system, development of, 455
1'ightward, di.?tection of, 239/ comprehension and, olfactory h'act, 435
second-order, 246 signal detection and, 12 olfactory white, 444
sine waves, 363 nonacddnetal features, 108 Oliva, Aude, 224-225, 227
visual perceptio1\ 237- 258 norunetrical depth clues, 161 Olney, john, 488
motion alter.fleet (MAE), 238, 238/, nonspoctral rolms, 134-135, 134{ "omnivoreo's dilemma," 483-488
245-246 nontoster (oi PCT / PROP), 480 ON-center bipolar· cells, 43. 45/, 46
motion pru·allax. 169-171, 170/ nose, anatomy of, ON-center retinal ganglion cells.
mouth, anatomy, 3'Ef nystagmus, 254 63/
movies, three-dimensional, 180-181 opacities, 35, 85, 105
Movshon, Tony, 192 0 opponent oolortheory, 135, 135/
116-117
.. objects
contrast-defined, 246
identification of, 228/
optic atl'ays, 247
optic disc. 36
optic flow, 170-171, 247, 248{
muscle spindles, 396, 396f identification through touch, 300 optic nerve (CN IQ, 19,<; 20
mu.sic interactions with, 410-411, 411/ optokinetic nystagmus (OKN), 254
cultural differences, 323-324 luminance-defined, 246 oral pain disorders, 475
emotions and, 325 memory for, 226-227 orbitofrontal rortex (OFC), 45B-459, 471/,
freq\lency range, 3'11/ perception of, S9-120 474{
notes, 321-325 perception problems, 95- 97 activation of, 401
perc.ption of, 321-346 recognition of, 89-120 taste pl'Ocessing, 47'\{, 475
production of, 32ihl28 recognition problems, 95-97, 96J organ of Corti, 269--270
musk compounds, 440 texture-defined, 246 orientation tuning, 71, 71f
myopia, 34 occlusion, 101, 161, 161/ 111e Origin o_fSpecies (Darwin), 18
octaves, 322 01·thonasal olfaction, 427, 490
N ocular dominance, 72, 7.Sf oscillato1y, definition ot 361
Nadoolman, \Volffe, 491 oculomotol' nerve (CN III), 19{, 20 ossicles, 266
naive template theory, 114 odo1· hedonics, 454 otitis media, 2115-286, 483
Nmmfl, or Coriceming t11e lvfe11tul Lift of odor imagery, 445 otoconia, 364
Plants, 5 !'eceptors (OR), 429, 4'?1Jf, 447f otolitl1 Ol'gans, 353, l.-'6, 364-366

INDEX 545
otosclerosis, 286 responses, 124 Q

ototoxl.c. definition of, 286 retinal, 39f, 124 qualia, 140-141
outer ears, 266 sensitivities, 154 quinine, 48lf
outer hair cells, 270-272 phototopic, definition of, 124
outer segments, photoreceptor, 40 pictorial depth cues, 167 R
oval window, 266 pig1nent epithelium, 37/ radiation therapy,, taste sensations and,
ovett attention, 202 pigs, olfacto1y neurons, 430 475
ovulation, 461 pimao, 266, 267/, 297- 301, 298f ra::inbmvs, limits of.134-135
piri£orm COlteX, 435 Ramacha.ndran, Vilayanur, 400-401
p pitch random dot stereograms (RDSs), 179-
P ganglion cells, 43, 44{, 47 absolute, 324-325 180, 179/
Pacini.an cotpuscles, 391f,. 392 definition of, 263, 321 rapid serial visual presentation (RSVP),
pain sound, 2S1, 283-285 212
components of, 402-405 place codes. 272 Rasch, R. A. , 312
lewis of, 402-404 placebo effects, 404-405 rate-intensity functions, 217
moderation of, 404--405 pleasant toudl,. 395-396 t'ate satw·ation, 276
sensitization to, 405 Poggio, Gian, 185 reaction times (RTs), 203
Pandemonium models, 104/ pointillism, 130, 130/ realism, 157
panpsychi.sm. philo"'PhY of, 5 polyrhytluns, 327 receiver operating ch.,'tracteristic (ROC)
Panum ·s fusional ru·eas, 173 polysensory, 20 curve, "14-15, 15/
papillae, 471-472 Ponzo, Ma.tio, 188 receptive fields, 44
parabelt area, 281 Ponzo illusion, 188 properties, 72
parnhippocam]>.'1 place area (PPA), popcorn, 462-463, 463/ shifting of, 218
94-95, l 13, 216, '116f positivism, 157 su'lgle cell, 218
parallel seard1es, 2rfl ommlqn tomo;!r;'!PhY (PET), size of, 391
parallelism 28, 29f striate cortex. 70-74
figure-ground and, 107 Posner, Michael, 203 updating, 256/
Gestalt grouping rule, 102, ·103f Po.sner cueing pMadigrn, 203f receptor adaptation. 451
parietal lobes postural system, 379 receptor potentials, l57
anatomy. 219 Potter, Molly, 227 rerognition-by-components model, 114
Jes.Ions, 21 9 pra:1ttentive stage, 211 reentrant proce.sBing. 118-119
somatosensory area 1 (Sl ), 398, '398/, prefrontal cortex. 403 reflKt, definition of, 32
399f pregnancy. sensations in, 475 reflectance, surfooo, 150
parvocellular layer, 64, 131 pr<Sb)'Opia, 35 reflexive eye mo\·ements, 254
Penfield, \Vllder, 400 Prescott, John, 48i Reissner's membtantt>. 268
pentanol, 452-453 primar)' auditory 001- (Al), 281, 312, relatability, 108
perception 342 relah?d colors, 146
action for, 410-412 primary olfactory cortex.. 435 relative heights, 162, 163/
color, 123-155 primary visual cortex (\11), 65, 89, 471/ relative metri.ml depth cues, 165
definition of, 3 primary visual pathway, 54f relative motion, 107
problems with, 95-97 prim et' ph"1'omone. 460 relative sizes, 162, 162/, 163/, 164,f
sc.aling methods, 9-11 prior piobability, 110 releaser phel'Omones, 460
of spatial orientation, 366-371 probability summation, 159 Remsen. lra,, 479-480
petft.uners, odor image1y in, 445 projective geometry, 161, 161/ Rensink, Ron, 227
perilymph. 359 propa.nol, 453 response enhat,cement,. 218
petiod, sine v1:a\'e, 16 prop1·ioception, 389 restoration effects, 314-315
Pfaffmann, Carl, propyltl1iowacil (PROP), ll, 480-481 retinal prostheses, 50{
phantom limbs, 400-401, 40lf propylthiow·adl (PROP) recoptor, 477 retinas
phantom taste sensations, 475 prosopagnosla, 9S, 118 oone-0pponent cells, 131 , 137-139
phase locking, audito1y nerve, 278, 278/ protanope individuals, 144 description of, 33, 35-36
phases protein molecules, size of, 487 duplex, 49
grating. 62-63 Proustia.n memoties, 462 image movement, 251-252
wave, 16 proximity, Gestalt grouping rule, 102,, images on, 15.1-159, 158[, 17'lf, 187f
phenylethyl alcohol, 442, 454, 455/ in:.fotmation transmission, 130-131
phenylthiocarbamide (PTC), 480-481 prul':iceptors, 403 light focusing onto, 33-35
phetomones, 459-462, 460 pseudogenes, 436 neural circuitty. 49
plionittion, types of, 328-329 psychoacoustics, 281 photomicrographs, 37/
photic sneeZQ reflex, 47 psychophysics, 6, 446 photoreceptors, 124
photoactivation, 41 pupils, eye "rule of tl1uo1b," 39/
photons, 31 desc1'iption of, 33 retinitis pigmentosa (RP),, 50
photopic vision, 42t response to light, 48, 48f retronasal olfaction. 427
photopigments,, regeneration of, 48-49 pure tones, 264 retronasal olfactory sensations, 467
photol'eceptors Reynolds, Jolu1, 218
fwlCtion of, 38 RGBslideis, 133-134
ph-ulcrographs, 37/ rhodopsin, 40, 125

8230336 Anma Appa
546 INDEX
rhythm, mu sic and_, 326-327 functi on of, 36(!f smooth pW'sui t, 251
Richa.rds, \.Vhitm.an,. 109 sensation, definition of, 3 sneeze refl exes, photic,. 47
Richter, Cw·t, 485 sensitivity (.f) Snellen cha11, Slf
Roberson, Debi, 141 levelofresponseand, 13, Sniffin' Sticks, 433
Robso" John, 62. 81 tactile, 405-41 0 sniffing, power of, 445
rocking moveme nts, 379 visual, 43 social insects, 460
rod monoduomats, 144 sensorineural hearing loss, 286 social tou ch, 420
rodents, olfactory neurons, 430 sensory exaffel'e nce, 371 sodium deficiency, 476
rods, 31lf somatosensation , 389
operatio n of,, 40 sensory integration, somatoserulofy area 1 (51), 398-399, 398f,
sensitivity1 49 sensory modalities 399f
roller coaster_, 385 competition between, 419_f, 420-422 somat""""""1'} aroo 2 (52), 399, 399f
Rollers (Kltaoka), 238 integration of, 412/ somatotypical inputs, 397
Roper, Steve, 487 sensory neuroscience, 4--5 sow 1d s
Rosch, FJeanor, 142 sensory roalfurence, 371-372 attack and decayr 307-308
rotation serial self-termina ting sea.rchl?s, 208-209 complex, 264-265, 30>-308, 343f
constant-\•elodty_, 36·t 36"lf set sizes, d esa:iption of, 200 daily e twironments, 264
directions of, 355-356 Seur.u, Georges, 130, 130/ d escription of, 262- 265
perception of, 367- 368 sex differences, in olfaction, 449 familiarity , ...·ith,. 313-314
rotational \'ection, 371/ sexual desire, odot's and, 461-462 harmonic, 265f
round '"'i 1ld.o\\.'S, 268 Shank, Jeffrey, 461 intens.i ty of, 301
Rozin, Paul, 48.3, 486 shape-pattet'll theory, 439-441 localization of, 291-302
Rubin, Edgar, 106 sharper tttnlng, 218 onset of. 308/
Rubi n vase/face figw·e, Shel le n, Herma.t\ Sl pres.su re fluctuations, 262f
Ruffini endi ngs, YJ1f, 392 Shepard, Roger, 277 scaling. 4
Sherrington, Charles, 21, '12f separation of, 'J(1}f
s Shovel!, Sloven, 123 spectral composition of, 301
S-cones, 124 shingles, 403 thresholds, 4
d efects, 144 Shipley, Thomas, 108 wave amplitudes, '02
photoreceptot·s, 154 slgnill detection theoty (SDT). 4, 12-15 sow · tastes, 475, 47/
response of, 126, 127/ aiterion level of response, 13 sow ·ce segregation, 309
SA l fibers, 392, 411 noise and, 12 soy formulas, 476
saocad.es" 253 peroeptual decisions and, 13/ spatial attention, tactile, 418-419
saocadic suppression, 205, 254-255 se:ns itivity m easu1l"s, 13- 14, 14{ spatial disor irotation, 3&l
saochruin_, 479 simi larity, Ges.talt grouping rnle of. 102, spatial freq uencies
saccules, 363 adaptation and, 80, 81/
S<Kks, Oliver, 160 Simons, Dan, 229 description, 17
SAil fibers, 393 simple cells patterns and, 59
salionce. 207 d efinition of, 72- 73 spatiill-li<quency chruu10ls, 81, SJ/
salt hunger, 4S5 flavors of, 72f spatial layouts, 224/
salty taste, 475, 476 tlmi.n g, 73/ global information and, 225
scales, cultmal differences, 324 sine wave grntings, !X>,56f,9'!f, 61-62 spatial ol'ientation
scali ng methods, 9-11 sine \'\>aves description of, 349
scan paths, 253/ cornbinations, 16f perception of, 366-371
scatter, 31 d escription, 15-16, 16/ sensoty mod alities in, 352
scene-based guidance, 210, 211/ sound,264 spatial orfantation cortex, 381-383
scene perception, 201-234 single-opponent cells, 139, 139f spatial vision,. 53-8'7
scents, feel of, 438 Sinha,. Pawanr 86 developme nt of, 83--85
Schade, Otto, 59 sinusoidal motio n, 36'lf speaimint, 441, 442
Sclafani, An tho ny, 488 sizes, cues fromr 162 specifi c ansomia, 440
sclera, ph- n icrograph.s, 37/ skin specifi c- h1.mgers throty, 485-487
sootopic, definition of, 124 anatomy of, 391/ spectrn 1 sound, 264
scotopic vislon, 42t fingel' l'idges, 41 '.Zf spec1tal composition curves, 149
serond-order motion, 246-247, 246/ messages to the brain from , 396-402r spectral contrast. 337-338
serond.aJv taste cortex. See orbitofrontal YJSJ spectral power d istribution, 14S- 149,
oornix(OFC) pei·ception of patterns w ith, 41 5-416 14llf
selective attention, 202, 203 touch reepptors, 389, 39J-J96 spectral reflecta nce fw1ctions, 14Sf
ADHD and, 221 ski n-to-ski n contact, 396 spectral se.nsitivity, 124
binding problem and, 211 sleep, ollaction during, 453-454 spectrill tilt, 307/
Seliridge, Oliver, 104 Small, Dana, 469 spectrograms
semidt'l"ltlar canals smell, sense of, 7t categoricill perception, 335f
anatomy of, 358-363, 351\f sme lls. See also odomnts; odors description of, 331, 331f
description of, 352 ide ntifim tion of, 44S-449 stop consonants, 334{
direction oxling in,. 361 me mory of, 447, 447f speech
dyna.mics of, 361 perception of, 434( brain and, 343- 346

INDEX 547
classification of sotmds, l31-333 symmetry, Gestalt grouping mlQ, 102, contt"aSt, 60

lea.ming to listen, 339-340 Hllf difforence, 6
pe«option of, 321-346 synapses, function of, 22 horuing, 287/
production of, 238-J33, 739f synaptic terminals, 40 olfactory detQction, 446--447
spinal accessory'"""' (CN XI), 19f, 20 syncopation, 3TJ sowld., 4
spinal cord, dorsal horn, 397, 397/ tWO-f"Oi11t touch, 6_, 405-409
spinothalarnk pathway, 398, '398f T velocity, .36Sf
square wave gr a ting, 56, Sf:! tactile agnosia, 416 Thrts Spcte Zamtlmstrn (Sb'auss), 317
staircase method, 447 tactile 1'eceptors, 392-394 tilt
staped.ius, 266 individual differences, 409-410 description of, 352
st-apes, 266 t.'U·gots, description of, 2U5 percoption of, 369-370, 369/
stereo vision, recovering ot 178 Tan-, 115 sense of, 353
ste1wacuHy, 193 TA52R g<ne, 477, 477f tilt aitereflro, 7S-79, 196
stereobli..ndness, 177 tastants, 473-474 tilt tests, vestibular system, 3'79/
ste1wcilia, 359 taste, sense of, 467-492 264-265
desai ption of, 270 absolute thresholds, 7t description of, 304-305, 305{
hair coll, 270-m basic tastes, 475-480 sow1d grouping, 311-312, 311J
photomicrograph. 27lf CNS pr"""'8ing of. 474/ time to collision (ITC}r 250
stereograrns, 175-17B facial expressions and, 484.f tinnitus, 384
filtet'ed version, 184/ flavor versus, 467-470 tip links, 271
random dot, 179-180 health consequences of, 482-483 tip-of-nose phenomenon, 448
stel'eoi.someres, 441 physiology of, 487 Tipper, Steve, 220
stereopsls, 160, 172f ll, 48"1--482 Titcheilel', F.dwcud Bradfot'd, 99
developmontof, 192-197, 193J thresholds, 11 tomatoes_. flavor of, 469-470
on""t of, 193/ taste adaptation, 490 ton• chromas, 322-323
175-178, 175J, 176{ tone heights, 322-323
stereoscopic rort'eSJ.-"\Otldl?tl('Q, 182-184 11ma A tongue, anatomy of, 471]
Stevens. S.S., 10, 481 taste buds tonic water, 477
Stevens' power law, 481 anatomy of, 473J tonotopic organization, 280
desaiption of, 10 desc1'ipti.on of, tons illectomy, 483
meastul:'ments, 11 taste receptor cells, 471, 476{ topographical mapping, 64
stimuli tasters (of PCT/ PROP), 480 Torralba, Antonio, 224-225
constant stimuli method, Sf tastes, 467 touch, descriptors fol', 420
proa>ssing of, 215-217, 217/ tau (•) , 250 touch, sense of, 388-425
receptot' response to, 391 tectotial membranes, 270 absolute thresholds, 7t
stimulus onset (SOA), 204 Te.Iler, Davida, 192 acuity ot, 405-410, 4oW
stop consonants, English, 332f temporal code, 278, 326 interactions, 420-422
strabisnms, 86, 196 temporal intQgration, 283 perreptjon of objects, 413-416
Strauss, Richai·d, 317 -porollobes physiology of, 390-405
striate cortex cellular activity, 91f sensitivity of, 405-410
function, 65-70 language and, 344-346, 344J, 345/ touch 390-396, 395-396
layers, 66 lesions, 90, 95, 118 transduce, desaiption of. 352
receptive fields, 70-74 temporo-parieto-insular cortex. 381 ttansmit, definition of, 32
striolae, 365 tensor tympani, 266 tl'a.nspatent, definitlon of, 32
stripes, fe?t\'.:eption of, 85--86 tetrachronrnts, 153-154 Treisman, Anne, 21 1
structural descrlption, 114 textur..dofined objects, 246 ttiangle test, 447--448
structuralism, 98 texture fields, 163 ttichromacy, 126
substan.tia gelatinosa, 402 texturn gradients, 162 ad\•a ntages, 152
subtractiverolormixttu·es, 129, 129/ texture segmentation, 101-104, 101/ histo1y of the theo1y of, 128-129
sucrose thalamus ttichromatic theo1y of color 1,.-ision, 126
psychophysical functions of, 48lf anatomy, 65 trigemhiai 11Q1'VO (CN \1, 19f, 20, 438,
sttucture of_. 478/ medial gr?rticulate nucleus, 280 438/
supelior collicuJus, 252 taste processing and, 474/ tt'igeminal perception, 449
supe1ior olive. 279 vestibular information and, 381 tt'imethyla.mine.
supedor temporal gyl'tts, 342f thenn oreceptors ttitanope individuals, 144
suPQrtasters, 11 , 481-482 descti.ption of, 394 ttochlear n<?!ve (CN IV), 19{, 20
sttpp:>rting cells, 429 pain and, 402 tufted cells, 435
suppression, 196 thQtmal receptivity. 394/ tttning curves. auditory neul'ons, 24{
sunoundednes.s, 106-108 thickening agents. 469 Tw·i:n,. Luca, 440
sttstained attention, 202 Thorpe, Simon, 223 (n ..•enty) 20/20 vision, 57
sweat glands, 391/ SD-movies, 180-181 two-point touch threshold, 6, 405-409,
sweet tastes, 468, 473, 475, 47&-480 thresholds 407/, 40Sf
Swithets, Susan, 480 absolut<>, 7, 7t t\'Vo--tone suppression, 275, '175/
Sylvian fissure. 281 a udibilty, 282 tympanic canal, 268
auditory nervl? tuning cw·ves, 274 tyrnpanic membt'a:ne. 266

548 INDEX
u 1ww·al pathways, 375/ vootl folds, 329, 330{

umM1i stimuli, 473, 476, 487-4&S l'esponses in the d ark, 376f i,.·ornl tract, 328
uncrossed disparity, vestibulo-spinal reflexes, 380f voicing, 333
llnique hues, 136 vestibulo-spi.nal t'esponses, 378-381 volatile-enhanced taste, 469--470
tul.iquenesscon.straint, 184 vestibttlocochlear nerve (CN Vlll), 19£ volley principle, 278, 279f
univar:iance, principle of, 124-126 20 vomeronasal organ, 459-462
University of Pennsylvania Smell Identi- vibration theo1y of olfaction, 440 vowel sounds, 306, 307f, 332{
fication To.st (UPSID. 433 vibrations
wu-elated colors, 146 detec"tlon of, Wf w
unides, 363 trailSlllission of, 270 warmth fibers, 394
video games, attentional blink in, Wart'et\ R. M., 316
v 213-215 waterfall illU5ion,. 237
V1 area Vieth-Miillel'd1cle, 173 Waterman , Ian, 396
newborns, 194 viewpoi nt invariance, 115 wavelengths
sense of touch and, 401 vigilance tasks, 202 color discrim ination and, 124-132
V2cells, 91 , "2f, 140 viltual reality (VR), 181 color pet'ception and, 123
V4cells, 11 2-113. 112/ vision, 31-51 light, 31
vagus nerve (CN X).19J,20 absolute thresholds. 7t sine waves, 16
Vaina, Lucia, 247 acuity, 43,. 53-62, 58t sound.. 26'lf
Valdez,. Benito Da1a de, SI middle {midle\·el), 97-110 waves
vanillin, 446 sensitivity, 43 light, 31
vanishing points. 167 sow1d localization and, 29'lf phasesof. 16
vection,. definitio n of, 370 spatial.. 53-87 Webe1·, Ernst, 6, 6f
velocity stereopsis. 17'lf Webe1· fractions, 6
definition of, 356 visual angles, 55, SSJ Weber'slaw, 6, 11
tlu·esholds, 368/ visual attention disorders, 219-222 Wells, William Chal'les, 373
velocity storage . .376 visual cortex, color In, 13!i-140 Wertheimer, 99
vergence, 17lf. 253 visual a·owding. 69-70, 70f Whea.tstone,,_ audes, 175
Vemil?t' acuity, 185 visual fields Whi>atstone' • stereo!oopO! 175--176, 'i'l!?f
vertebrates, evolution of, 351 brai n activity and, 215 '"'hite noise, 284
vertigo,349 defocts, 67, 219 Wi...,I, Torsten, 70-76
vestibular canal, 268 human vs. rabbit, 159f Williams, David, 128
vestibular organs, 349 mapping. 68/ wintergreen, 442
vestibular sense visual mapping 65. 67/ words. Ieanli.ng, 340-342
absolute thresholds, 7t i,:isualmotion Wundt, Wlll,.lm, 5, 98-99
acti\•e tll?t\vorks for. 37'2/ computation of, 23S-247
afferent neurons, 372 perception of, 237- 258 x
vestibular system visual pathways, wiring diagram.. 9lf X-rays, limitations of, '2'1, 27/
amusement park rides, 384-385 visual resolution acuity, 55
ascending pathways, 3S2f visual sm.ne, retinal image, 187 y
desctiption of, .349 visual seal'ch ya\•l rotation, 36&-369
manmrnlia.n,. 356-366 binding problem in, 211-212 Yow"lg, ll"lOmas, 128
problems with, 383-334 d es::rtption o f, 205 Yow"'lg-Hehnholtz theoty, 128
sense of equilibrium and,. 349-386 laboratory tasks. 20&f
vestibular thalamocortical p;:i,thways, visual stllbility. 374/ z
381-382 visually-evoked electrical potentials Zellner, Debra, 486
vestibulo-autonomic responses, 377-378 (VEPs), 84-84 Zollner illusions, 189, 190/
vostibulo-ocular reflex (VOR), 350, 350f, vital ism, 20, 21 zonules of Zinn, 34
37'!.-!377, 374f vitamin Bl def:i.cien.des, 485
dynamics of, 371/ vitreous humor, 33


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Brief Contents

CHAPTER 3 Spatial Vi sion: From Spots to Stripes 52

CHAPTER 4 Perceiving and Recognizing Objects 88

CHAPTER 5 Th e Perception of Color 122

CHAPTER 6 Space Perception and Bi nocular Vision 156

CHAPTER 7 Attention and Scene Perception 200

CHAPTER 8 Visual Motion Perception 236

CHAPTER 10 Hearing in the Environment 290

CHAPTER 11 Music and Speech Perception 320

CHAPTER 12 The Vestibular System and Our Sense

CHAPTER 13 Touch 388

CHAPTER 14 Olfaction 426

CHAPTER 15 Taste 466

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JEREMY M- WOLFE is Professor of Ophthalmology and Radiology at Harvard

KEITH A . KLUENDER is Department Hea d, Professor of Speech, language, and

LINDA M. BARTOSHUK is Bus hnell Professor, Department of Food Science and

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ROBE:RTA L. KlATZKY is the Charl es J. Q ueena n, Jr. Professor of Psychology at

OANIE:L M. is a Professor of O tology and Laryn gology at the Harvard

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Introduction 3 Signal Detection Theory 12

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Spatial Vision: Simple and Complex CeDs 72

Visual Acuity: Oh Say, Can You See? 53 Columns and Hypercolumns 74

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The Perception of Color 123 Sensation & Perception in Everyd ay

Space Perception and Binocular Random Dot Stereograms 179

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Attention and Scene Attention and Single Cells 217

Visual Motion Perception 237 Avoiding Imminent Collision: The Tao of

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CHAPTER 9 __ '· Ar;/

The Function of Hearing 261 Basic Operating Characteristics of the

Hearing in the Environment 291 Auditory Scene Analysis 308

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CHAPl'ER 1 1''11 • ; , -y-· r ...

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Touch 389 Haptic Perception 410

Psychophysical Methods for DetectJ'on and

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8230336 Am111a Appa

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Welcome to Our World

Sensation and Perception

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METHOD 1 : THRESHOLDS What is tl'e faintest sound you can hear? H ow

METHOD 2: SCALING-MEASURING PRIVATE EXPERIENCE W hen you

METHOD 3: SIGNAL DETECTION THEORY-MEASURING DIFFICULT DECI-

METHOD 4: SENSORY NEUROSCIENCE Grilled peppers appear on your

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METHOD 5: NEUROIMAGING-AN IMAGE OF THE MIND Suppose you ar-

Thresholds and the Dawn of Psychophysics

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FIGURE 1.5 This illustratbn of Fechner' s law shows

w h e re S is the psychological sen s'1 ti on , which is

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"I hear it:'

"I don't hear it:' J

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