BIO3420.2010.6 Sensory Systems 15 and 18oct10 2

Sensory Systems
Chapter 6; 15-18.10.2010
Sensory Receptors
Range from single cells to complex sense organs
Types of receptors
Chemoreceptors, mechanoreceptors, photoreceptors,
electroreceptors, magnetoreceptors, thermoreceptors
All receptors transduce incoming stimuli into

changes in membrane potential
Receptor protein detects stimulus
Opening or closing of ion channel
Change in membrane potential
Signal sent to integrating center (central nervous
system)
Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
Sensory Receptors
Figure 6.1
Graded Potentials
Generator potential
Sensory receptor is also the primary afferent neuron
Change in membrane potential spreads along
membrane
Receptor potential
Sensory receptor is separate from the afferent
neuron
Change in membrane potential triggers release of
neurotransmitter
Graded Potentials
Figure 6.2
Classification of Sensory Receptors

Based on stimulus location
Telereceptors
Detect distant stimuli
For example, vision and hearing
Exteroceptors
Detect stimuli on the outside of the body
For example, pressure and temperature
Interoceptors
Detect stimuli inside the body
For example, blood pressure and blood oxygen
Tells little or nothing about how receptors work


Based on type of stimulus (stimulus modality) the
receptors detect
Chemoreceptors
Chemicals
For example, smell and taste
Mechanoreceptors
Pressure and movement
For example, touch, hearing, balance, blood pressure
Photoreceptors
Light
For example, vision

Electroreceptors
Electrical fields
Magnetoreceptors
Magnetic fields
Thermoreceptors
Temperature
Sensitivity to Multiple Modalities

Adequate stimulus
Preferred (most sensitive) stimulus modality
Many receptors can be excited by other stimuli, if

sufficiently strong
For example, pressure on eyelid perceive light
Polymodal receptors
Sensitive to more than one stimulus modality
For example, nociceptors; polymodal receptors for
multiple types of pain
Stimulus Encoding
All stimuli are ultimately converted into action
potentials in a primary afferent neuron
How can organisms differentiate among stimuli or
detect the strength of the signal?
Sensory receptors and sensory neurons must
encode four types of information
Stimulus modality
Stimulus location
Stimulus intensity
Stimulus duration
Stimulus Modality and Location

Receptor location encodes stimulus modality and
location
Integrating center interprets modality and location
Modality
Theory of labeled lines
Discrete pathway from sensory cell to integrating center
Polymodal receptors are exceptions

Encode modality via temporal patterns of APs
Location
Theory of labeled lines
Receptive Field and Location of Stimulus

Receptive field
Region of the sensory surface that causes a response
when stimulated
Smaller receptive field allows more precise location of
the stimulus (i.e., greater acuity)
Improved ability to localize stimuli by

Using more than one sensory receptor cell
Lateral inhibition
Signals from neurons at the center of the receptive
field inhibit neurons on the periphery
Receptive Field and Location of Stimulus
Figure 6.3
Dynamic Range
Sensory neurons code stimulus intensity by
changes in action potential frequency
For example, strong stimuli high frequency
Dynamic range
Range of stimulus intensities over which a receptor
exhibits an increased response
Threshold of detection
Weakest stimulus that produces a response in a receptor
50% of the time
Saturation
Top of the dynamic range (maximal response)
Dynamic Range
Figure 6.4a
Dynamic Range and Discrimination

Trade-off between dynamic range and
discrimination
Large dynamic range
Large change in stimulus causes a small change in AP
frequency
Large dynamic range
Poor sensory discrimination
Narrow dynamic range

Small change in stimulus causes a large change in AP
frequency
Small dynamic range
Good sensory discrimination
Dynamic Range and Discrimination
Figure 6.4b
Range Fractionation
Range fractionation
Groups of receptors work together to increase
dynamic range without decreasing sensory
discrimination
Figure 6.4c
Encoding Logarithmically
Encode a wide range of stimulus intensities using a
single receptor cell
Good discrimination at certain intensities
Poor discrimination at other intensities
Figure 6.4d
Tonic and Phasic Receptors

Two classes of receptors encode stimulus duration:
Phasic
Produce APs at the beginning or end of the stimulus
Encode change in stimulus, but not stimulus duration
Tonic
Produce APs as long as the stimulus continues
Encode duration of stimulus
Receptor adaptation AP frequency decreases if
stimulus intensity is maintained at the same level
Tonic and Phasic Receptors
Figure 6.5
Chemoreception
Most cells can sense chemical stimuli
Animals have many types of chemoreceptors
Olfaction (smell)
Detection of chemicals in air
Gustation (taste)
Detection of chemicals emitted from food
Olfaction and gustation are distinguished by structural

criteria
Performed by different sense organs
Different signal transduction mechanisms
Processed in different integrating centers
The Olfactory System

Evolved independently in vertebrates and insects
Vertebrate olfactory system
Can distinguish thousands of odorants

Located in the roof of the nasal cavity
Mucus layer to moisten olfactory epithelium
Odorant binding proteins
Allow lipophilic odorants to dissolve in mucus
Receptor cells are bipolar neurons with cilia

Odorant receptor proteins are located in the cilia
The Olfactory System
Figure 6.6
Odorant Receptors
Each olfactory neuron expresses only one odorant
receptor protein
There are 1000s of different receptor proteins
Each receptor can recognize more than one odorant

Each odorant can stimulate more than one receptor
Odorant receptor is linked to G-protein
Odorant binding causes formation of cAMP
Opening of ion channels
Depolarization
Odorant Receptors
Figure 6.7
Pheromones
Vomeronasal organ
Detects pheromones
Chemical signals between animals
Structurally and molecularly distinct from the

olfactory epithelium
Located in base of nasal cavity in mammals
Located in palate in reptiles
Receptor is linked to G-protein
Activates phospholipase C transduction system
Depolarization
Pheromones
Figure 6.8
Invertebrate Olfactory Systems

Located in many parts of the body
Most near the head
Primarily on antennae in Arthropods

Sensilla
Hair-like projections of cuticle
Sensilla contain odorant receptor neurons
Odorant receptor is linked to G-protein
Odorant binding causes formation of cAMP
Depolarization
Invertebrate Olfactory Systems
Figure 6.9
The Gustatory System

Five classes of tastants
Salty
Sweet
Bitter
Sour
Umami (savory or meaty)
Sweet, umami, and salty indicate carbohydrates,

proteins, and ions
Bitter and sour indicate potentially toxic substances
Taste Buds in Vertebrates

Taste receptors are epithelial cells that release
neurotransmitter
vertebrate taste receptors are not neurons
Each taste receptor expresses more than one kind of

taste receptor protein
Taste receptor cells clustered in groups
On tongue, soft palate, larynx, and esophagus
On external surface of the body in some fish
Vertebrate Taste Bud
Figure 6.10
Taste Receptor Transduction Pathways
Figure 6.11a,b
Taste Receptor Transduction Pathways
Figure 6.1c,d
Taste in Invertebrates
Located on sensilla
Inside and outside the mouth
Along the wing margin
Ends of the legs
Receptors
Bipolar sensory neurons
G-protein coupled
Express only a single receptor protein
Differences between vertebrates and invertebrates

suggest independent evolution
Mechanoreceptors
Transform mechanical stimuli into electrical signals
All organisms (and most cells) sense and respond
to mechanical stimuli
Two main types of mechanoreceptor proteins:
ENaC
Epithelial sodium channels
TRP channels
Transient receptor potential channels
Channels are linked to extracellular matrix

Mechanical stimuli alter channel permeability
Mechanosensory Protein Receptors
Figure 6.12
Touch and Pressure

Three classes of receptors
Baroreceptors
Interoceptors detect pressure changes
Tactile receptors
Exteroceptors detect touch, pressure, and vibration
Proprioceptors
Monitor the position of the body
Vertebrate Tactile Receptors

Widely dispersed in skin
Receptor structure
Free nerves endings
Nerve endings enclosed in accessory structures
(e.g., Pacinian corpuscle)
Vertebrate Tactile Receptors
Figure 6.13
Vertebrate Proprioceptors
Monitor the position of the body
Three major groups
Muscle spindles
Located in skeletal muscles
Monitor muscle length
Golgi tendon organs

Located in tendons
Monitor tendon tension
Joint capsule receptors

Located in capsules that enclose joints
Monitor pressure, tension, and movement
Insect Tactile Receptors

Two common types of sensilla
Trichoid
Hairlike projection of cuticle
Bipolar sensory neuron
TRP channel
Campaniform
Dome-shaped bulge of cuticle
Bipolar sensory neuron
Insect Tactile Receptors
Figure 6.14
Insect Proprioceptors
Scolopidia
Bipolar neuron and complex accessory cell
(scolopale)
Can be isolated or grouped into chordotonal organs
Most function in proprioception
Can be modified into tympanal organs for sound
detection
Scolopidia
Figure 6.15
Equilibrium and Hearing

Utilize mechanoreceptors
Equilibrium (balance)
Detect position of the body relative to gravity
Hearing
Detect and interpret sound waves
Vertebrates
Ear is responsible for equilibrium and hearing
Invertebrates
Organs for equilibrium are different from organs of
hearing
Statocysts
Organ of equilibrium in invertebrates
Hollow, fluid filled cavities lined with
mechanosensory neurons
Statocysts contain statoliths
Dense particles of calcium carbonate
Movement of statoliths stimulate mechanoreceptors
Statocysts
Figure 6.16a.b
Insect Hearing
Strong vibrations sensed by trichoid sensilla
Weak vibrations and sounds are detected by
chordotonal organs
Clusters of scolopidia
Located on leg
Mechanosensitive ion channels
Tympanal organs
Thin layer of cuticle (tympanum) overlays
chordotonal organ
Vertebrate Hair cells

Mechanoreceptor for hearing and balance
Modified epithelial cells (not neurons)
Cilia on apical surface
Kinocilium (a true cilium)
Stereocilia (microvilli)
Tips of stereocilia are connected by proteins (tip
links)
Mechanosensitive ion channels in stereocilia

Movement of stereocilia change in permeability

Change in release of neurotransmitter from hair cell
Hair cells
Figure 6.17
Signal Transduction in Hair Cells
Figure 6.18
Fish and Amphibian Hair Cells

Hair cells detect body position and movement
Neuromast
Hair cells and cupula
Stereocilia embedded in gelatinous cap
Detect movement of water
Lateral line system

Array of neuromasts within pits or tubes running
along the side of the body
Fish Neuromast
Figure 6.19
Vertebrate Ears
Function in both equilibrium and hearing
Outer ear
Not in all vertebrates
Pinna
Auditory canal
Middle ear
Not in all vertebrates
Interconnected bones in air-filled cavity
Inner ear
Present in all vertebrates
Series of fluid-filled membranous sacs and canals
Contains mechanoreceptors (hair cells)
Mammalian Ear
Figure 6.20
Inner Ear: Vestibular Apparatus and Cochlea

Vestibular apparatus detects movements
Three semi-circular canals with enlarged region at one
end (ampulla)
Two sacklike swellings (utricle and saccule)
Lagena
Extension of saccule
Extended in birds and mammals into a cochlear duct
or cochlea for hearing
Hair cells present in vestibular apparatus and

lagena (cochlea)
Vertebrate Inner Ears
Figure 6.21
Vestibular Apparatus
Mechanoreceptors of the inner ear
Macula
Present in utricle and saccule
Mineralized otoliths suspended in a gelatinous matrix
Stereocilia of hair cells embedded in matrix
>100,000 hair cells
Detect linear acceleration and tilting of head
Cristae
Present in ampullae of semicircular canals
Gelatinous matrix (cupula) lacks otoliths
Stereocilia of hair cells embedded in matrix
Detect angular acceleration (turning) of head
Vestibular Apparatus
Figure 6.22a.b
Maculae Detect Linear Acceleration

and Tilting
Figure 6.23
Cristae Detect Angular Acceleration
Figure 6.24
Sound Detection by Inner Ear

Fish
Sound waves cause otoliths to move
Displacement of cilia on hair cells
Some fish use the swim bladder to amplify sounds
Figure 6.25
Sound Detection by Inner Ear

Terrestrial Vertebrates
Hearing involves the inner, middle, and outer ear
Sound transfers poorly between air and the fluidfilled inner ear
Amplification of sound waves
Pinna acts as a funnel to collect more sound
Middle ear bones increase the amplitude of vibrations
from the tympanic membrane to the oval window
Mammalian Middle and Inner Ear
Figure 6.26a
Mammalian Inner Ear

Specialized for sound detection
Perilymph
Fills vestibular and tympanic ducts
Similar to extracellular fluids (high Na+ and low K+)
Endolymph
Fills cochlear duct
Different from extracellular fluid (high K+ and low
Na+)
Organ of Corti
Hair cells on basilar membrane
Inner and outer rows of hair cells
Stereocilia embedded in tectorial membrane in

cochlear duct (filled with endolymph)
Mammalian Inner Ear
Figure 6.26a,b
Sound Transduction
Sound waves vibrate tympanic membrane
Middle ear bones transmit vibration to oval window
Oval window vibrates
Pressure waves in perilymph of vestibular duct

Basilar membrane vibrates
Stereocilia on the inner hair cells bend
Hair cells depolarize
Hair cells release neurotransmitter (glutamate)
Glutamate excites sensory neuron
Round window serves as a pressure valve

Encoding Sound Frequency

Frequency Detection
Basilar membrane is stiff and narrow at the proximal
end and flexible and wide at distal end
High frequency sound vibrates stiff end
Low frequency sound vibrates flexible end
Specific regions of brain respond to specific

frequencies
Place coding
Encoding Sound Amplitude

and Amplification
Amplitude Detection
Loud sounds cause larger movement of basilar
membrane than quiet sounds
depolarization of inner hair cells
AP frequency
Outer hair cells amplify quiet sounds

Change shape in response to sound
Do not release neurotransmitter

Change in shape increases movement of basilar
membrane
Increased stimulus to inner hair cells
Detecting Sound Location

Brain uses time lags and differences in sound
intensity to detect location of sound
Sound in right ear first
Sound located to the right
Sound louder in right ear

Sound located to the right
Rotation of head helps localize sound
Photoreception
Ability to detect visible light
A small proportion of the electromagnetic spectrum
from ultraviolet to near infrared
Ability to detect this range of wavelengths supports
idea that animals evolved in water
Visible light travels well in water; other wavelengths do
not
Electromagnetic Spectrum
Figure 6.27a,b
Photoreceptors
Range from single light-sensitive cells to complex,
image-forming eyes
Two major types of photoreceptor cells:
Ciliary photoreceptors
Have a single, highly folded cilium
Folds form disks that contain photopigments
Rhabdomeric photoreceptors
Apical surface covered with multiple outfoldings called
microvillar projections
Microvillar projections contain photopigments
Photopigments
Molecules that absorb energy from photons
Phylogeny of Photoreceptors
Figure 6.28
Vertebrate Photoreceptors
Vertebrates have ciliary photoreceptors
Rods
Cones
Both have inner and outer segments

Inner and outer segments connected by a cilium
Outer segment contains photopigments
Inner segment forms synapses with other cells
Vertebrate Photoreceptors
Figure 6.29
Characteristics of Rods and Cones
Table 6.1
Diversity in Rod and Cone Shape

Diverse shapes of rods
and cones among
vertebrates
Shape does not
determine properties
of photoreceptor
Properties of
photoreceptor depend
on its photopigment
Figure 6.30
Photopigments
Photopigments have two covalently bonded parts
Chromophore
Derivative of vitamin A
For example, retinal
Contains carbon-carbon double bonds
Absorption of light converts bond from cis to trans
Opsin
G-protein-coupled receptor protein
Opsin structure determines photopigment characteristics
For example, wavelength of light absorbed
Isomerization of Retinal
Figure 6.31
Phototransduction
Steps in photoreception
Chromophore absorbs energy from photon
Chromophore changes shape
Double bond isomerizes from cis to trans
Activated chromophore dissociates from opsin

Bleaching
Opsin activates G-protein

Formation of second messenger
Ion channels open or close
Phototransduction
Figure 6.32
The Eye
Eyespots
Cells or regions of a cell that contain photosensitive
pigment
For example, protist Euglena
Eyes are complex organs

Detect direction of light
Light-dark contrast
Some can form an image
Types of Eyes
Flat sheet eyes
Some sense of light direction and intensity
Often in larval forms or as accessory eyes in adults
Figure 6.33a
Types of Eyes
Cup-shaped eyes (e.g., Nautilus)
Retinal sheet is folded to form a narrow aperture
Discrimination of light direction and intensity
Light-dark contrast
Image formation
Poor resolution
Figure 6.33b
Types of Eyes
Vesicular Eyes (present in most vertebrates)
Lens in the aperture improves clarity and intensity
Lens refracts light and focuses it onto a single point on
the retina
Image formation
Good resolution
Figure 6.33c
Types of Eyes
Convex Eye (annelids, molluscs, arthropods)
Photoreceptors radiate outward
Convex retina
Figure 6.33d
Compound Eyes of Arthropods

Composed of ommatidia (photoreceptor)
Each ommatidium has its own lens
Images formed in two ways

Apposition compound eyes
Ommatidia operate independently
Each one detects only part of the image
Afferent neurons interconnect to form an image
Superposition compound eyes

Ommatidia work together to form image
Resolving power is increased by reducing size and

increasing the number of ommatidia
Compound Eyes
Figure 6.34
Structure of The Vertebrate Eye

Sclera
White of the eye
Cornea
Transparent layer on anterior
Retina
Layer of photoreceptor cells
Choroid
Pigmented layer behind retina
Tapetum
Layer in the choroid of nocturnal animals that reflects
light
Structure of the Vertebrate Eye

Iris
Two layers of pigmented smooth muscle
Pupil
Opening in iris allows light into eye
Lens
Focuses image on retina
Ciliary body
Muscles that change lens shape
Aqueous humor
Fluid in the anterior chamber
Vitreous humor
Gelatinous mass in the posterior chamber
Structure of The Vertebrate Eye
Figure 6.35
Image Formation
Refraction bending of light rays
Cornea and lens focus light on the retina
In terrestrial vertebrates, most of the refraction occurs
between air and cornea
Lens does fine focusing
Lens changes shape to focus on near or far objects

Accommodation
Image Accommodation
Accommodation
Light rays must converge on the retina to produce a
clear image
Focal point
Point at which light waves converge
Focal distance
Distance from a lens to its focal point
Image Accommodation
Distant objects
Light rays are parallel when entering the lens
Ciliary muscles contract
Lens is pulled and becomes thinner
Little refraction of light by lens
Close objects
Light rays are not parallel when entering the lens
Ciliary muscles relax
Lens becomes thicker
More refraction of light by lens
Image Accommodation
Figure 6.36
The Vertebrate Retina

Arranged into several layers
Rods and cones are are in the retina and their outer
segments face backwards
Other cells are in front of rods and cones
Bipolar cells, ganglion cells, horizontal cells, amacrine
cells
Axons of ganglion cells join together to form the optic

nerve
Optic nerve exits the retina at the optic disk (blind
spot)
Vertebrate Retina
Figure 6.37a
The Fovea
Region in center of retina
Overlying bipolar and ganglion cells are pushed to the
side
Contains only cones
Color vision
Provides the sharpest images
Image is focused on the fovea
Cephalopod Eye and Retina

Photoreceptors are on the surface of the retina
Project forward
Supporting cells are located between photoreceptor

cells
No other layers of cells associated with photoreceptors
Axons of photoreceptors form optic nerve
Cephalopod Eye and Retina
Figure 6.37b
Signal Processing in the Retina

Rods and cones form different images
Rods
Convergence
Many rods synapse with a single bipolar cell
Many bipolar cells synapse with a single ganglion cell
Ganglion cells has large receptive field

Poor resolution (fuzzy image)
Cones
Each cone synapses with a single bipolar cell
Each bipolar cells connects to a single ganglion cell
Ganglion cell has small receptive field
High resolution
Convergence in the Vertebrate Retina
Figure 6.38a,b

Complex on and off
regions of the receptive
fields of ganglion cells
improve their ability to
detect contrasts between
light and dark
Figure 6.39

On and off regions of the receptive field of
ganglion cells improve contrast of light and dark
Center-surround organization of receptive field
On-center ganglion cells
Stimulated by light in center of receptive field
Inhibited by light in periphery of receptive field
Off-center ganglion cells

Stimulated by dark in center of receptive field
Inhibited by dark in periphery of receptive field
Photoreceptors in center and periphery inhibit each

other by lateral inhibition
Lateral Inhibition in the Retina
Figure 6.40
The Brain Processes the Visual Signal

Action potentials from retina travel to brain
Optic nerves optic chiasm optic tract lateral
geniculate nucleus visual cortex
Binocular vision
Eyes have overlapping visual fields
Binocular zone
Combine and compare information from each eye to

form a three-dimensional image
Depth perception
The Brain Processes the Visual Signal
Figure 6.41
Color Vision
Detecting different wavelengths of visible light
Requires photopigments with different light
sensitivities
Most mammals: see two (dichromatic) colors
Humans: see three (trichromatic) colors
Birds, reptiles and fish: see three, four
(tetrachromatic), or five (pentachromatic) colors
Color Vision
Retina and brain compare output from each type of
receptor and infer the color
Figure 6.42
Thermoreception
Central thermoreceptors
Located in the hypothalamus and monitor internal
temperature
Peripheral thermoreceptors
Monitor environmental temperature
Warm-sensitive
Cold-sensitive
Thermal nociceptors detect painfully hot stimuli
ThermoTRPs
Thermoreceptor proteins
TRP ion channel
Specialized Thermoreception
Specialized organs for detecting heat radiating
objects at a distance
Pit organs
Pit found between the eye and the nostril of pit vipers
Can detect 0.003C changes (humans can detect only
0.5C changes)
Figure 6.43
Magnetoreception
Ability to detect magnetic fields
For example, migratory birds, homing salmon
Neurons in the olfactory epithelium of rainbow trout
contain particles that resemble magnetite
Responds to magnetic field

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Sensory Systems

All receptors transduce incoming stimuli into

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Classification of Sensory Receptors

Tells little or nothing about how receptors work

Classification of Sensory Receptors

Classification of Sensory Receptors

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Sensitivity to Multiple Modalities

Many receptors can be excited by other stimuli, if

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Stimulus Modality and Location

Polymodal receptors are exceptions

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Receptive Field and Location of Stimulus

Improved ability to localize stimuli by

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Receptive Field and Location of Stimulus

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Dynamic Range and Discrimination

Narrow dynamic range

Dynamic Range and Discrimination

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Tonic and Phasic Receptors

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Tonic and Phasic Receptors

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Olfaction and gustation are distinguished by structural

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

The Olfactory System

Can distinguish thousands of odorants

Receptor cells are bipolar neurons with cilia

The Olfactory System

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Each receptor can recognize more than one odorant

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Structurally and molecularly distinct from the

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Invertebrate Olfactory Systems

Primarily on antennae in Arthropods

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Invertebrate Olfactory Systems

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

The Gustatory System

Sweet, umami, and salty indicate carbohydrates,

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Taste Buds in Vertebrates

Each taste receptor expresses more than one kind of

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Vertebrate Taste Bud

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Taste Receptor Transduction Pathways

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Taste Receptor Transduction Pathways

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings