Epilepsy & Behavior 7 (2005) 578–601

www.elsevier.com/locate/yebeh

Review

Absolute pitch: Music and beyond

David A. Ross a,*, John C. Gore b, Lawrence E. Marks c
a
Department of Diagnostic Radiology, Yale School of Medicine, Box 208043, New Haven, CT 06520, USA
b
Department of Radiology and Radiological Sciences, Vanderbilt University Medical Center, 1161 21st Avenue South, R-1032 MCN,
Nashville, TN 37232, USA
c
John B. Pierce Laboratory and Departments of Epidemiology/Public Health and Psychology, Yale School of Medicine,
290 Congress Avenue, New Haven, CT 06519, USA

Received 26 May 2005; accepted 27 May 2005

Available online 15 August 2005

Abstract

‘‘Perfect pitch,’’ known in the scientific literature as ‘‘absolute pitch’’ (AP), is a rare phenomenon that has fascinated musicians
and scientists alike for over a century. There has been a great deal of conflict in the literature between advocates of the two main
theories on the etiology of AP: some believe that AP is learned early in life through intensive musical training, whereas others believe
AP to be largely innate. Both theories are alike, however, in considering AP to be exclusively a musical phenomenon. We propose a
paradigm shift by presenting here a new model of AP, one that is predicated on two principles: (1) that AP may be relatively inde-
pendent of musical experience; and (2) that there are different types of AP, each of which can be ascribed to discrete neurobiological
mechanisms. We also review data from a diverse series of experiments that were designed to test explicitly both the predictions of our
model and a series of historical myths about AP. In each case, the data strongly support our model. We conclude with a general
discussion on the nature of AP, the relevance of these findings for other areas of research, and future directions of study.
Ó 2005 Elsevier Inc. All rights reserved.

Keywords: Absolute pitch; Perfect pitch; Music; Preattentive processing

1. Introduction We propose a paradigm shift: rather than assuming

The ability to identify the musical pitch of auditory
stimuli without the use of a reference pitch is a rare skill,
commonly known as ‘‘perfect pitch’’ or, in the scientific
literature, as ‘‘absolute pitch’’ (AP). Because of the
potential musical advantages it may endow, this skill
has enormous sociopolitical significance. Within the sci-
entific community the etiology of this trait has been
vociferously debated for over a century: on one side of
the argument are those who believe that it is learned ear-
ly in life, and on the other, those who believe it is innate.
However, advocates of both these theories share the
underlying assumption that AP is strictly a musical
phenomenon.
*
Corresponding author.
E-mail address: david.a.ross@yale.edu (D.A. Ross).
that AP is strictly a musical phenomenon we explore a
broader view of pitch processing, which sees AP as
reflecting capacities that can far exceed the perception
of music per se. To see AP in this way requires abandon-
ing the traditional definition of AP (i.e., as the ability to
name notes) in favor of a more inclusive criterion. Our
rationale for doing this was originally based on observa-
tions of heterogeneity among subjects who claimed to
have ‘‘perfect pitch’’: we observed some individuals
whose performance in naming notes appears to be spe-
cific to music, and who apparently developed this skill
through particular kinds of early experience; other indi-
viduals with AP, however, evince a capacity to encode
frequency information automatically, not only for music
but for a variety of other acoustical signals. These latter
individuals may name notes with great facility if or when
they are exposed to musical training, but their ability to

1525-5050/$ - see front matter Ó 2005 Elsevier Inc. All rights reserved.
doi:10.1016/j.yebeh.2005.05.019
 D.A. Ross et al. / Epilepsy & Behavior 7 (2005) 578–601
encode the frequency of auditory stimuli transcends mu-
sic per se. Because we believe these groups differ mecha-
nistically, we eschew the ambiguous label of ‘‘AP’’ in
favor of names that directly reflect the relevant neuro-
biological processes: we refer to the former group as
possessing heightened tonal memory (HTM) and the lat-
ter as possessing the ability to perceptually encode
(APE) the frequency of auditory stimuli.
In this article we review historical perspectives on AP,
discuss their relative strengths and weaknesses, and then
present a new theory that may reconcile previous points
of conflict in the literature. In doing this, we take a phe-
nomenon that has been considered predominantly from
the cognitive and behavioral domains and connect it to
the burgeoning neuroscientific literature on low-level
mechanisms of auditory perception. We then review
data from a series of experiments that were designed
to test both the explicit predictions of our model and a
series of historical myths and dogmas in the field. Fore-
most among these is the assumption that AP is funda-
mentally a musical phenomenon.
2. Cultural and historical perspectives on absolute pitch
Most individuals perceive melodic sequences without
being able to identify absolutely the notes involved. For
example, most people recognize the opening bar of Bee-
thovenÕs Fifth Symphony but are unaware that the first
note is a ÔG.Õ Thus, in most people the dominant mode
of pitch perception entails recognizing the relations
among notes, rather than the exact frequency of each
individual note. When refined further, this skill—the
ability to identify or produce musical intervals accurate-
ly—is referred to as relative pitch (RP).
Nevertheless, a small subset of the population is also
capable of quickly and accurately labeling the absolute
frequency of tonal stimuli (without the use of a reference
pitch). In musical settings, this skill is generally referred
to as perfect pitch, while in the scientific community it is
more frequently referred to as absolute pitch (AP). Given
how rare and how distinct this phenotype is relative to
the ‘‘normal’’ mode of auditory perception (the preva-
lence of AP is generally estimated to be between 1/
5000 and 1/10,000 [1–3]), it is not surprising that AP
has fascinated musicians (and audiences) for years and
has been explored in the scientific literature for well over
a century (cf. [4]). This interest has been further piqued
by frequent suggestions (of varying degrees of legitima-
cy) that possessing AP may confer superior ability in
some musical domains. One obvious advantage is that
it may be relatively easy for AP possessors to transcribe
or remember a piece of music because they can identify
directly each of the notes that are played. Another is
that it may be easier for AP possessors to sight-sing dif-
ficult passages by producing each note individually
579
rather than trying to tune challenging intervals. More
speculative is the claim that AP may be a prerequisite
of musical genius and that possessors have an increased
ability to know whether a particular note is properly
tuned.
Conversely, AP may also be limiting in several impor-
tant ways. For one, strict adherence to the absolute fre-
quencies of tones as found in the equal tempered scale
could hinder a possessorÕs ability to remain in tune with
a choir that drifts flat or to maintain proper intonation
on intervals that differ from equal temperament. More
dramatically, there are reports that a significant percent-
age of AP musicians find that their sense of pitch shifts
as they grow older [1,5], a change that can be completely
debilitating for a professional musician.
2.1. The etiology of absolute pitch
Although numerous studies have explored potential
anatomical and physiological bases that could account
for differences between AP possessors and nonposses-
sors (e.g., possessors tend to have increased asymmetry
between right and left planum temporales [6–8] and
may have decreased P300 responses to tones [9–12]),
the etiology of the trait remains unclear. Early models
of AP suggested that ‘‘true’’ AP is a genetically deter-
mined trait [13,14], and several recent articles have pro-
vided convincing evidence that AP has a genetic
component [1,15–17]. Other models of absolute pitch
advocate the position that AP is the result of early learn-
ing experiences [18–21]. While this debate may seem
largely academic, it is also of great social significance;
a multimillion dollar industry exists solely to train musi-
cians to develop AP.
2.1.1. The early learning theory
According to the early learning theory (ELT), AP
possessors were exposed to a form of training that
caused them, through repeated presentation, to form
strong associative links between specific pitches and
their appropriate musical labels. Thus, an individualÕs
AP should depend on the nature and extent of his or
her musical training (i.e., the age at which the person
began training and the extent to which training empha-
sized the association of pitch names with absolute pitch-
es), and individuals with AP should be more facile at
naming more familiar stimuli [e.g., notes played on their
primary instrument, more commonly encountered notes
(such as white vs black), or within the range of their
instrument]. Critically, according to this theory, the abil-
ity to recognize tones is indistinguishable from the abil-
ity to label them. As one advocate wrote: ‘‘An important
and related point is that AP is not an unusual ability in
the domain of Ôpitch perception,’’ despite the fact that
Baharloo et al. repeatedly refer to the ability this way.
AP is a skill in labeling (a form of long-term memory
580 D.A. Ross et al. / Epilepsy & Behavior 7 (2005) 578–601
and categorization/classification behavior, involving
self-referencing) and has nothing to do with pitch per-
ception per se’’ [22, p. 257].
2.1.2. The innate model
In contrast, advocates of the innate model have argued
that AP may be better defined as an ability to transfer the
initial sensory trace of a stimulus into a more stable, long-
term representation. This representation may, but need
not, involve a musical label [23–25]. Thus, the ability to
recognize and to encode a long-term representation of a
tone may be independent of a subjectÕs ability to label it.
Importantly, this skill may not depend on subjectsÕ musi-
cal experience, though their ability to apply musical labels
to tones would obviously require knowledge of musical
nomenclature. This subtle point is elegantly captured by
Corliss [26, p. 1738], herself an AP possessor: ‘‘There is
certainly some element of memory in absolute pitch, even
if it is just the association of the gamut with certain arbi-
trary pitch standards. In that sense, it cannot be said to be
entirely innate. It seems to me that some of the perception
processes that allow the gamut to be recognized in detail
may be innate abilities.’’ Given this nuance, one might
reasonably argue that a more perspicuous approach
would be to identify AP possessors based on the underly-
ing capacity to recognize pitch chroma1 rather than on the
culturally acquired, if not arbitrary, ability to name notes.
2.2. Weaknesses of historical models
Historically, the ability to name musical notes has al-
ways been regarded as the sine qua non of AP, and we
have followed the tradition in so describing it above
(see the beginning of Section 2). Unfortunately, several
major methodological flaws arise from this definition
that may inhibit researchersÕ ability to meaningfully
study the phenomenon. First, the use of a note naming
paradigm creates a screening bias such that only musi-
cians of Western training can take the test; thus, neither
nonmusicians nor musicians of non-Western training
would know the names of the notes and could not be
included in a standard protocol. More significantly,
the exclusive use of these tests has entrenched in the
community the assumption that AP is fundamentally a
musical phenomenon. Accordingly, the possibility that
AP may be better explained by basic mechanistic differ-
ences in auditory perception has largely been ignored (if
not outright rejected). Finally, by considering note nam-
ing as an end in itself, one may be led to believe that AP
is a simple, binary, trait. To the contrary, note naming is
1
Chroma is typically defined as the frequency of a stimulus
independent of its octave, an attribute particularly important for
music. For example, despite the fact that ‘‘C5’’ has an overall
frequency twice that of ‘‘C4,’’ they both have the same chroma and
perform identical functions in most musical contexts.
a complex behavioral task that may be performed using
different strategies, strategies that in turn may reflect dif-
ferent underlying neurobiological mechanisms. Consis-
tent with this possibility are several reports of
heterogeneity within cohorts of individuals who name
musical notes with facility [1,3,4,19,20,27,28]. Few inves-
tigations, however, have attempted to explicitly address
this heterogeneity, and none has attempted to distin-
guish subtypes of AP mechanistically.
2.3. Reconciling different models of AP
In our own laboratory, we have observed different
subtypes of AP. There are some individuals who name
notes effortlessly and automatically. In our mind, these
are the individuals who possess ‘‘true’’ or ‘‘genuine’’
AP as originally described by Bachem [27]. Elsewhere,
we have argued that this group may be defined by the
unique ability to encode meaningful representations of
stimulus frequency, and we developed a paradigm inde-
pendent of music to test this hypothesis. With this par-
adigm, we showed not only that AP possessors differ
from a group of nonpossessors with comparable musical
experience, but that ‘‘true AP’’ may exist in: (a) musi-
cians without early musical training [29], (b) adult non-
musicians [30], and (c) children who have not yet had
sufficient musical experience to learn the names of all
the musical notes [31].
In contrast, for other AP musicians the ability to name
notes appears to involve active calculation based on com-
parison of target tones to a memorized template. As such,
performance tends to improve the more closely the target
stimuli match items in a subjectÕs memorized template
(where key parameters typically include timbre, height,
and chroma). For reasons that become evident below,
we believe this qualitative difference between the two clas-
ses of AP possessors is one of type rather than degree.
Accordingly, we consider the latter type of subjects as pos-
sessors of ersatz AP, distinct from the group described
above. Rather than being specific to note naming tasks,
the difference between true and ersatz AP possessors
was apparent when subjects were tested using a range of
musically independent paradigms [29].
To some extent, then, the argument over which model
(early learning vs innate) is ‘‘correct’’ may be moot: each
model may accurately describe some possessors of
‘‘AP.’’ It is plausible that the ELT accurately describes
possessors of ersatz AP, whereas the innate model
describes the more limited group of true AP possessors.
2.3.1. A useful analogy
The difference between these subtypes of AP resem-
bles the distinction that has been made between ‘‘preat-
tentive’’ and ‘‘automatic’’ processing of stimuli in the
visual field [32]. TreismanÕs description of preattentive
processing closely matches the precategorical definition
 D.A. Ross et al. / Epilepsy & Behavior 7 (2005) 578–601
of AP: ‘‘Preattentive processing does not require extend-
ed practice. It depends on mechanisms that are either in-
nate or acquired early and outside the laboratory. Its
functions are also different—not to make skilled perfor-
mance autonomous, but rather to meet the needs of the
preliminary stage of visual coding.’’
In contrast, Treisman describes ‘‘automatic’’ process-
ing [33] in the following manner: ‘‘automatization results
from the accumulation across repeated trials of specific
memory traces, increasing the probability of direct
retrieval of the correct response to the evoking stimulus
and bypassing any rules or decision processes.’’ She later
summarizes: ‘‘We tentatively conclude that automatiza-
tion . . . speeds up processing at a later stage, after feature
integration has been achieved, rather than creating new,
preattentively detectable features. The effects seem to
depend on the formation of new and very specific associ-
ations between features, their locations, and the required
responses.’’ These are almost verbatim descriptions of the
early learning theory for absolute pitch recognition.
Qualitatively, we have observed that pitch/chroma is
a salient aspect of auditory stimuli only in true AP pos-
sessors. For everyone else the primary mechanism of
pitch perception is interpreting the intervalic relation
between pitches. By analogy, we propose that pitch chro-
ma may be a feature of auditory perception only in true
AP possessors. In the visual domain ‘‘feature’’ process-
ing is thought to reflect low-level mechanisms used to
encode stimuli. Similarly, we suggest that true AP may
result from a fundamental property of the way the audi-
tory system encodes stimulus frequency.
In contrast, possessors of ersatz AP may have an
increased ability to memorize and retrieve specific con-
junctions of basic auditory features (e.g., timbre, height,
and intensity). However, as in non-AP-possessors (NAP
individuals), chroma is not a basic feature of perception
and the initial mechanisms used to encode and translate
auditory stimuli are similar (though it is plausible that
subjects have differences at higher points in the auditory
pathway). Importantly, the ability to recognize auditory
‘‘objects’’ will depend strongly on previous exposure to
similar objects.
While this model represents a reasonable qualitative
description of AP, we now turn to the critical question
of whether it is consistent with known mechanisms of
auditory perception.
2.3.2. Models attributing pitch perception to low-level
processes
It was originally thought that musical pitch is encoded
primarily as a spectral (or tonotopic) signal.2 By this mod-
el, an incoming frequency stimulus creates a traveling
wave on the basilar membrane and, because of the tension
2
For detailed descriptions of spectral and temporal models see [34].
581
gradient along the organ, the place of maximum ampli-
tude corresponds to the fundamental frequency. This spa-
tial analysis, in turn, leads to increased firing within a
particular characteristic frequency (CF) band in the audi-
tory nerve, such that different pitches may be encoded via
different CF bands. These tonotopic representations are
carried up to the cortex, though they do not convey pre-
cise absolute pitch information in most individuals. To
a certain extent, this pathway undoubtedly affects pitch
perception. However, there are a number of limitations
in the descriptive capacity of this model [35].
More recently, it has been suggested that musical
pitch may be encoded not by CF bands per se, but by
the temporal firing pattern of the inner hair cells. Across
the entire length of the basilar membrane, all of these
cells fire in a pattern that is phase locked to the incoming
stimulus. For example, for a stimulus of 100 Hz, while a
place code may exist leading to increased firing within a
particular CF band, the entire basilar membrane
vibrates at 100 Hz and all hair cells will fire in phase
with this signal at exactly 100 Hz. Thus, the same timing
information exists in all CF bands. The pitch of a stim-
ulus is then extracted via an autocorrelative process that
measures the time intervals between successive spikes.
This process may be conceptualized by looking at a his-
togram of all-order intervals between spikes in the audi-
tory nerve. The highest peak on the plot corresponds to
the inverse of the stimulus pitch and the relative height
of this peak (compared to the rest of the distribution)
may indicate pitch salience [35].3
Strong evidence has accrued in support of this model,
which has excellent predictive power for a wide range of
perceptual phenomena [35]. However, there is also a
central paradox: if the model proposes that there exists
an exact low-level representation of the pitch of all fre-
quency stimuli (as in the autocorrelogram), then why
are most people unable to access this representation?
Unfortunately, few researchers have addressed this
dilemma. One notable exception is a theory proposed
by Cariani [36] in which a series of operations are per-
formed on the output of the autocorrelogram, including:
echoic memory (via recurrent timing nets with delay
loops of various lengths); relative pitch processing of
vertical intervals4 (via feedforward timing nets that use
3
To be clear, it is unlikely that either model is exclusively correct:
current debate is focused on establishing the relative roles of tonotopic
and periodotopic signals across a wide range of perceptual processes
and stimulus types. It should also be noted that most ‘‘temporal’’
models incorporate spectral information by including CF bands as an
additional input factor. The details of this debate are tangential to our
purposes here. In the present discussion, we are interested simply in the
extent to which musical pitch processing makes use of temporal codes
(with or without spectral information).
4
A vertical interval is the difference in pitch between two simulta-
neously presented tones, as opposed to a horizontal interval, when the
tones are presented sequentially.
582 D.A. Ross et al. / Epilepsy & Behavior 7 (2005) 578–601
coincidence detectors to compare interval distributions
of two inputs); and relative pitch processing of horizon-
tal intervals (via a combination of the two previous
mechanisms). Of note, however, all of these operations
take place exclusively in the temporal domain. Thus,
the representation of absolute pitch would exist solely
as an intermediate point in the overall pathway.
2.4. A new theory of absolute pitch perception
The idea of a central subsystem with an exact repre-
sentation of the absolute pitch of stimuli is extremely
compelling. Indeed, it could provide a parsimonious
explanation of the differences between AP and NAP
individuals: What if true AP possessors, possessors of
APE, are defined by the unique ability to access this
low-level representation? In a simple version of such a
model, some individuals would have an additional syn-
apse that translates the output of the autocorrelo-
gram—a temporal signal—into a place code for pitch
(e.g., if a peak at 10 ms synapses onto a cell correspond-
ing to 100 Hz) that is then carried up to the cortex as an
additional stream of auditory information.5
Consequently, we propose the following theory to ac-
count for differences between true absolute pitch and
NAP individuals (as schematically depicted in Fig. 1).
In normal individuals the basic features of auditory per-
ception include: loudness, height (via the tonotopic path-
way), relative pitch, and timbre processing6 (the last two
via the periodotopic pathway). In AP possessors, the
auditory system also contains a mechanism that trans-
lates the most frequently occurring spike interval into a
specific place code for pitch. Thus, in this group, the
absolute pitch of stimuli is coded as an additional basic
feature of auditory perception. Performance by subjects
in this group should match the predictions of the innate
theory: recognition of stimulus frequency would be
immediate and independent of prior experience or any
other stimulus attributes. It is interesting to note that this
simple difference nicely accounts for BachemÕs suggestion
that NAP individuals perform pitch memory tasks using
height only (thus suggesting that the maximum resolu-
tion of the tonotopic pathway is five to nine half-steps),
while AP possessors encode both height and chroma.
We have described another subset of AP possessors
as having an increased ability to memorize and recall
conjoined auditory ‘‘features.’’ These individuals would
recognize a target stimulus by comparing the incoming
signal with the evoked memory of previous, similar sig-
nals. This comparison would depend on the differential
5
N.B. This is not the first model to suggest that AP possessors may
differ from nonpossessors in the character or function of cells in a
brainstem nucleus [37].
6
Which may be viewed in large part as a derivative of vertical RP
processing.
use of the secondary processing systems that are present
in normal individuals. One system likely to be particu-
larly important is that used for interpreting timbre. As
illustrated in Fig. 1 this second groupÕs ability to process
stimulus frequency would depend largely on periodotop-
ic mechanisms. Other researchers, however [38], have
argued that timbre may be processed primarily as a spec-
tral representation. If this were true, then pitch percep-
tion by this subset of AP possessors would be based
primarily on tonotopic information and relatively inde-
pendent of periodotopic coding. A third possibility is
that the extraction of pitch information takes place at
a higher level in the pathway: given that stimulus recog-
nition derives from the processing of conjunctions of
features, it is entirely possible that the critical mecha-
nism occurs at a point where temporal and spectral
streams have already been reintegrated. Regardless of
the precise location in the pathway, though, all of these
possibilities hold in common the essential aspect of our
theory: that ersatz AP possessors have an increased abil-
ity to extract AP information from systems that are nor-
mally used to process other basic auditory features.
Furthermore, because repeated exposure to specific con-
junctions of features will increase the probability of rec-
ognizing the same conjunction on future presentation,
performance by this group should be closely tied to
the intensity and nature of training.
Given this scheme, it is reasonable to believe that
such a learned skill may be present to a varying degree
throughout the population. At its simplest, this type of
process could explain why normal individuals tend to re-
call with surprising accuracy the frequency of the first
note of popular songs [39–41]. Similarly, it is well known
that instrumentalists may form a strong tonal memory
of the frequency to which they tune their instrument
(e.g., a violinist may recall the frequency of A440). In a
more extreme case, this could also explain why a highly
trained pianist may learn to recognize any note played
on a piano but have greater difficulty recognizing notes
with unfamiliar timbre. Much of the literature in sup-
port of the early learning model of AP could be inter-
preted easily within this conceptual framework. It
remains to be seen whether some individuals are predis-
posed, by virtue of innate neurophysiological mecha-
nisms, to being better at such feature integration or
whether it is purely the result of practice.
2.4.1. New terminology
For lay people, the term perfect pitch has come to
have different meanings for virtually everyone who uses
it. Similarly, in the scientific literature, many researchers
have acknowledged that ‘‘AP’’ possessors do not form a
single homogeneous group. This ambiguity poses a
major obstacle to research in the field: it may be virtual-
ly impossible to know what types of subjects were in-
volved in any particular study of ‘‘AP.’’ Ultimately,
 D.A. Ross et al. / Epilepsy & Behavior 7 (2005) 578–601 583

Fig. 1. Schematic representation illustrating proposed AP pathway (dotted lines).

the cause of this ambiguity has been the use of a diag-
nostic criterion that has both poor specificity and poor
sensitivity. The model that we present suggests the exis-
tence of two discrete mechanisms, either of which could
endow an individual with the ability to name notes and,
thereby, to pass a ‘‘classic’’ AP test.
For the sake of clarity, we prefer to identify groups
on the basis of the proposed underlying mechanism.
Thus, we define APE as the unique ability to perceptual-
ly encode durable representations of stimulus frequency
at a precategorical level, independent of a targetÕs tim-
bre, chroma, or height. This definition corresponds
closely to other groupsÕ definition of ‘‘genuine’’ AP
[27]. In contrast, we define individuals who recognize
target stimuli by comparing them with their memory
of specific auditory events as possessors of heightened
tonal memory (HTM). This definition corresponds (at
least broadly) to that used by advocates of the early
learning theory (e.g., Miyazaki, Levitin, and Takeuchi
and Hulse). By these definitions, APE possessors are
characterized by the way they encode stimulus frequen-
cy, whereas HTM possessors are characterized primarily
by an increased ability to retrieve specific complex audi-
tory memories. While the ability to name musical tones
584 D.A. Ross et al. / Epilepsy & Behavior 7 (2005) 578–601
may be an easy index for identifying ‘‘genuine’’ AP pos-
sessors, we do not believe that this skill is fundamental
to the condition. Rather, we argue that APE is a basic,
psychoacoustic capacity that, in conjunction with musi-
cal training, would make it relatively easy to associate
labels (either from the Western musical scale or other-
wise) with frequency stimuli. Thus, a double dissociation
may be possible between the ability to name notes and
‘‘genuine’’ AP: as illustrated by subject R.M. [28],
APE possessors without musical training may be inca-
pable of naming notes; conversely, expert musicians
may develop a refined sense of HTM that makes it easy
for them to recognize notes in a conventional paradigm.
Importantly, the distinction that we propose between
APE and HTM is easily testable. Foremost, in groups
matched for musical experience, our theory predicts that
performance by HTM musicians should be affected by
the many parameters that define the target tone. Specif-
ically, they should be more accurate at recognizing or
identifying tones with more familiar timbre (e.g., piano
vs sine), chroma (e.g., white keys vs black keys), height
(e.g., within the range of their primary instrument), or
intonation. All of these effects have been reported in
support of the early learning theory [20,42–45]; by our
model, they reflect the limitations of HTM. We predict
that a small subset of the conventionally defined popu-
lation of ‘‘perfect pitch’’ possessors should be minimally
affected by these factors. We regard this subset as the
possessors of APE.
Critically, we may also formulate an explicit null
hypothesis for our model: namely, that the distinction
we have drawn between APE and HTM is artificial.
According to this alternate hypothesis, APE and HTM
possessors are drawn from opposite ends of the same
continuous spectrum, and both result from the same
underlying mechanism (which could be consistent with
either the early learning or innate model). The most
obvious feature that one might use to define such a spec-
trum would be the ability to name notes.
The following experiments were designed to test our
new theory using a series of paradigms that would be
relatively independent of subjectsÕ musical experience
and note naming ability.7 In doing so, we hoped: (1)
to clarify the extent to which AP may be considered a
musical phenomenon versus a generalized perceptual
phenomenon; (2) to identify the preattentive ‘‘features’’
that define ‘‘normal’’ auditory perception; and (3) to
7
Subjects were classified as APE, HTM, or NAP possessors on the
basis of their performance on a series of separate paradigms, with the
most important skill being the ability to encode and reproduce
stimulus frequency accurately after a prohibitive amount of tonal
interference across a diverse set of timbres (including MF, iterated
ripped noises, sine, and piano tones). Full details of this method may
be found in [31]. In total, 17 subjects qualified as APE possessors and
30 qualified as HTM possessors.
clarify the relative importance of tonotopic and periodo-
topic mechanisms in normal and absolute pitch
processing.
The second major goal was to address directly some
of the vast folklore about AP that has accumulated over
time. Not all myths are mythical, but all are untested.
The present experiments were designed to determine
which of the common claims about AP are fictional
and which have a legitimate scientific basis.
3. Encoding and reproducing HugginsÕ pitch stimuli
As described above, early models of pitch perception
emphasize a place code on the basilar membrane,
whereas more recent models emphasize temporal codes.
Oddly, this shift in thinking has not been incorporated
into models of AP. With one notable exception [28], vir-
tually all theorists on the topic maintain that AP pos-
sessors encode the pitch of target stimuli via a simple
place code (generally in the context of discussing why
AP possessors may experience shifts in pitch perception
with increasing age). Ironically, though an advocate of a
place code model [27], Bachem himself originally noted
the correspondence between the ability of hair cells to
phase lock to stimuli and the ability of AP possessors
to identify musical pitch consistently [46]; however, to
the best of our knowledge, this observation has largely
been ignored.
The following study [47] was designed to test the
extent to which AP depends on place versus temporal
codes. Data from an earlier study [31] suggest that spec-
tral cues may be important in HTM perception, as evi-
denced by a poorer ability to encode and reproduce
iterated rippled noises (IRNs) and missing fundamental
stimuli relative to spectral stimuli, such as pure tones. In
contrast, APE subjects performed comparably with all
three kinds of stimuli. While performance on IRN stim-
uli poses a reasonable first test of the relative importance
of spectral versus periodotopic encoding, it is not ideal:
though the stimuli are designed to minimize the amount
of spectral information present, nevertheless, with an
increased number of iterations, a mild place code and
overtone series do exist.
For this reason, we sought a paradigm that could test
more rigorously the importance of tonotopic representa-
tions in AP perception. To do so, we adapted our origi-
nal paradigm [29] to use target stimuli that would
preclude absolutely the possibility of subjects using a
place code—HugginsÕ pitches (HPs) [48]. HP is generat-
ed by an interaural phase transition at a particular fre-
quency within a broadband noise. At this frequency,
the interaural phase relationship changes sharply with
increasing frequency, shifting through 360° over a nar-
row bandwidth (typically 3–6%). Above and below the
transition frequency, the noise is identical in each ear.
 D.A. Ross et al. / Epilepsy & Behavior 7 (2005) 578–601
Because each ear is presented with a white noise, the
pitch is dichotic (i.e., it cannot be heard in either ear
alone), so no place code can exist. Listeners generally
hear ‘‘a warbling tone standing out from the noise,
whose pitch can be matched to the center frequency of
the phase transition’’ [49]. Previous data have shown
that HP may be processed as musical pitch [50]. Howev-
er, to the best of our knowledge, neither HP stimuli nor
any other dichotic pitch stimulus has ever been used to
test AP.
To summarize, traditional theories assume that AP
depends on place (spectral) rather than temporal infor-
mation; consequently, these theories would predict that
AP possessors would have difficulty processing HP stim-
uli. Based on our theory, we predict that individuals
with APE encode pitch through temporal processes
(periodicity) and thus will readily process the pitch of
HP stimuli, whereas individuals with HTM, to the
extent that their ability to encode stimulus frequency
depends on spectral processing, may not.
3.1. Summary of findings
Subjects were presented with a target stimulus fol-
lowed by either a silent interval of varying length (2, 8,
or 16 seconds) or an interval of equal duration filled
with a series of interfering tones (1, 31, or 71 interfering
tones). Previously, we showed this to be a powerful
means of distinguishing among APE, HTM, and NAP
possessors [29,30]. The target tones were HP stimuli
from within the range of C5 (523.3 Hz) to B5
(973.2 Hz); to maximize interference, sine tones were
used from across a wider range (from C2, 65.4 Hz, to
B7, 1975.5 Hz). Following this interval, subjects were
asked to reproduce the original target stimulus by
adjusting the knob of a sine function generator. The
generator was always turned to a frequency of
1000 Hz, and subjects were blindfolded to prevent visual
monitoring of their performance.
Data were analyzed by calculating the distance away
in half-steps (hs, semitones) of each response from the
target. Because we were interested only in subjectsÕ abil-
ity to encode chroma, responses were corrected to the
nearest octave. Thus, the possible range of responses
was from
6 to +6 hs. Data were quantified using stan-
dard deviation (r) as a measure of variability and con-
stant error (CE) as a measure of bias. Perfect
performance would be expected to yield a normal distri-
bution at zero (low CE, low r), whereas chance perfor-
mance would yield a uniform distribution from
6 to
+6 hs (low CE, high r).
Data from representative APE and HTM subjects are
shown in Fig. 2 (without interfering tones) and Fig. 3
(with interfering tones). Data from a cohort of 6 APE
and 13 HTM subjects are presented in [47]. There were
no major effects by group or interval length without inter-
585
ference. With interference, the APE group performed
more accurately than the HTM group. There was also a
significant interaction between group and trial type (i.e.,
with and without interference). These data indicate that
APE subjects were less affected than HTM subjects by
the presence of interfering tones. Interestingly, when com-
pared with performance on identically structured para-
digms with different types of target tones (i.e., sine,
piano, missing fundamental, and IRN targets), APE per-
formance was identical, whereas HTM subjects per-
formed significantly better with HP stimuli than they
had with the other types of targets.
These data show that APE and HTM subjects can en-
code durable representations of HP stimuli, at least as
readily as they do for more conventional types of target
stimuli.
3.2. Discussion
Despite recent research emphasizing the importance of
periodotopic information in low-level frequency process-
ing, virtually every model of perfect pitch has maintained
the dogma that AP processing depends directly on the
interpretation of a place code on the basilar membrane.
Were this the case, one would expect AP possessors to
be significantly hindered in their ability to encode pitch
stimuli that cannot be defined by a spectral code. The
present data refute this hypothesis: both APE and HTM
subjects performed at least as well with HP stimuli as with
conventional spectral stimuli. Previous data have shown
that HP stimuli may be used for relative pitch processing
[48]. The present data corroborate the overall importance
of periodotopic signals in musical processing.
These data are consistent with the mechanism we
have proposed for APE subjects, namely, that subjects
may have an increased ability to extract AP information
from the periodotopic pathway. Of greater interest, they
help refine our perspective on pitch processing in the
HTM group. Previously, we suggested that the ability
to extract AP information from other stimulus features
(e.g., timbre) might take place in the tonotopic pathway,
the periodotopic pathway, or at a relatively high stage,
after integration of these streams. The present data
appear to contradict the first of these possibilities. Fur-
ther investigation is required to determine the relative
importance of low-level periodotopic processing and
higher-order integration in HTM pitch recognition.
4. Distinguishing between properly and improperly tuned
stimuli
Part of the mythology of ‘‘perfect pitch’’ is that musi-
cians with this trait may possess a superior ability to
reproduce specific frequencies accurately (e.g., to sing
A4 = 440 Hz) or to recognize tones that deviate from
586 D.A. Ross et al. / Epilepsy & Behavior 7 (2005) 578–601

Fig. 2. Data from representative APE and HTM subjects for HP target stimuli with a silent ISI of 2, 8, or 16 seconds.

Fig. 3. Data from representative APE and HTM subjects for HP target stimuli for ISIs with 1, 31, and 71 interfering tones.
 D.A. Ross et al. / Epilepsy & Behavior 7 (2005) 578–601
proper tuning (e.g., to recognize if an instrument is
slightly out of tune). In a sense, these purported skills
are mirror images of each other in that both effectively
ask how finely AP possessors internally represent pitch,
the former in the context of reproduction, the latter in
the context of encoding and recognition.
Different models of AP yield different predictions as
to how well possessors should be able to perform on
these tasks. According to the early learning theory (or
our own definition of HTM), certain individuals,
through repeated exposure to musical stimuli, form a
template of specific tones that they consider to be ‘‘in
tune.’’ In this case, performance on the task should cor-
relate with both the extent and the specific nature of mu-
sical experience; i.e., the more experience one has with
an equal tempered instrument the better that individual
should be. Thus, one would expect to see a spectrum in
performance, with highly trained keyboardists doing
best and less trained vocalists doing relatively worse.
Alternatively (and consistent with our definition of
APE), different pitches may possess an inherent salience,
independent of a personÕs musical experience. By this
rationale, a minimum amount of training should confer
to these individuals a sense of which specific frequencies
are ‘‘correct,’’ and the ability to distinguish in-tune and
out-of-tune stimuli should be relatively independent of
training (at least beyond a certain threshold).
Although these issues have been studied indirectly in
several ways, e.g., how small a difference can AP pos-
sessors remember after a delay with interference [23]
and how do AP possessors categorize notes that are mis-
tuned across a spectrum [20,22,51], to the best of our
knowledge, no study has explicitly tested the ability to
distinguish between properly and improperly tuned
stimulus frequencies. In fact, the only explicit reference
of which we are aware is by Levitin, who wrote:
Recent articles . . . have demonstrated confusion about
the nature of AP by using the term ‘‘Perfect Pitch’’ inter-
changeably with the term ‘‘Absolute Pitch.’’ The unfor-
tunate implication of the term ‘‘Perfect Pitch’’ is that
possessors of the ability have some sort of super resolu-
tion in their pitch perception, and that they can tell
whether a tone is perfectly in tune or not. In fact, there
is nothing ‘‘perfect’’ about absolute pitch. APers are no
better at tone discrimination than other individuals, and
they are no more accurate at noticing deviations from
perfect intonation. What they are better at is labeling
tones along the unidimensional continuum of frequency,
but there is some ‘‘slop’’ or ‘‘hysteresis’’ in their category
boundaries. [22, pp. 256–257]
The goal of the following study [52] was to com-
pare the ability of APE, HTM, and NAP musicians
to distinguish between notes tuned to the equal tem-
perament (ET) scale with A4 = 440 Hz and notes that
deviated either 25 or 50 cents from this scale. Effec-
587
tively, we asked: Can musicians with ‘‘perfect pitch’’
distinguish accurately between in-tune and out-of-tune
notes?
The experiment was conducted using a confidence
rating paradigm from signal detection theory. This
paradigm offers an important benefit: it ameliorates at
least some of the arbitrary nature of the task (i.e.,
knowledge of A4 = 440 Hz). Because of the way it is
structured, the paradigm allows the possibility of doing
either significantly better or significantly worse than
chance. For example, if a given AP musician usually
played an instrument that was chronically flat, she or
he might perceive the stimuli that were 50 cents off as
being correct and the others as mistuned. Rather than
performing at chance, data from such an individual
would yield a receiver operating characteristic (ROC,
in signal detection theory) that fell below chance perfor-
mance (thus measures of sensitivity, such as d 0 , would be
negative, and the area under the ROC, p(A), <0.5).
Thus, rather than asking how well subjects recognize
the precise frequency values of the ET scale, the para-
digm tests how well subjects can distinguish among three
different sets of stimuli, each of which is internally con-
sistent and slightly different from the others.
4.1. Summary of findings
Subjects were presented with 100 pure sine tones rang-
ing from C3 (130.8 Hz) to B4 (493.9 Hz). Thirty-four of
the tones were tuned to the ET scale with A4 = 440 Hz,
33 differed by 25 cents, and 33 differed by 50 cents. For
each tone, subjects were required to rate their confidence
that the tone came from the ET on a scale ranging from
1 = completely confident that it did not come from the
ET scale to 6 = completely certain that it did.
Mistuned (MT) stimuli were defined as the targets, and
separate ROC curves were generated for the 25- and 50-
cent stimuli. Data were quantified by calculating p(A),
the area under the curve, for each subject, and, to account
for the possibility of ‘‘mistuned’’ subjects (who would have
negative ROC curves), the average of the positive distribu-
tion (APD = the absolute difference between p(A) and 0.5
for each subject) was also calculated for each group.
Fig. 4 shows p(A) values for representative APE,
HTM, and NAP subjects. Data from 17 APE, 29
HTM, and 8 NAP control musicians are presented in de-
tail in [52]. APE performance was significantly better
than chance for both conditions. HTM performance
exceeded chance for the 25-cent stimuli and there was
a trend toward above-chance performance for the 50-
cent stimuli. The NAP group did not differ from chance
for either set of stimuli. Statistical comparison showed
that APE subjects were better able than HTM subjects
to distinguish between ET and MT tones and that
they were more sensitive to the difference between
25- and 50-cent stimuli.
588 D.A. Ross et al. / Epilepsy & Behavior 7 (2005) 578–601
Fig. 4. p(A) data for representative APE, HTM, and NAP subjects for target stimuli mistuned by 25 or 50 cents.
Separate analyses were conducted to test a number of
alternate explanations of our data. These analyses
showed that performance on this task did not correlate
with: type of instrument, years experience, or the age
of onset of training. Further, neither APE nor HTM
subjects were significantly affected by target chroma
(i.e., they did not perform more accurately for common
notes—C, E, G—than uncommon notes—D#, F#, G#),
and neither groupÕs performance correlated with either
the ability to name notes or the ability to reproduce
A4 = 440 Hz accurately.8
4.2. Discussion
According to popular mythology, musicians with
‘‘perfect pitch’’ are highly adept at distinguishing
between properly tuned and mistuned stimuli. This
belief, however, has been largely anecdotal. The present
study is the first to assess this skill explicitly and the data
show a clear empirical basis for the myth.
While at face value these results appear to reflect dif-
ferences in musical skill, this is not the case. All of the
subjects who participated had sufficient musical experi-
ence to familiarize themselves with the ‘‘correct’’ value
of each note. Yet despite their extensive training, the
NAP controls were incapable of distinguishing between
properly and improperly tuned stimuli and the HTM
subjects were only marginally better. The lack of corre-
8
These data were collected after completion of one of the previous
reproduction tasks. After the last response, the sine function generator
was reset to 1000 Hz and subjects were asked to tune it to match A440.
They were not given any feedback on the task and there was no
immediate musical context to assist them.
lation with any other musical parameters—note naming
ability, memory for A4 = 440 Hz, or overall experience —
similarly confirms the nonmusical nature of the task.
Instead, these data support the fundamental assertion
of our model: Stimulus chroma is a salient feature of
perception for APE subjects only. In this group, differ-
ences in pitch of as little as 25 cents are meaningfully
encoded. For these individuals our task entailed identi-
fying the pitch of each stimulus within a continuous
spectrum and judging the distance to the nearest known
reference point. Because the 50-cent stimuli lie further
from such references, they were recognized with greater
confidence and accuracy.
While the HTM subjects performed mildly better than
chance, they were relatively insensitive to subtle differenc-
es in pitch. These data suggest that HTM subjects per-
formed the task using a more basic strategy: each target
was compared directly to the memory of a ‘‘correct’’ tone
and the judgment was made as a simple same/difference
response. Thus, while these individuals have been able
to establish in their minds several fixed points of reference,
these references may be relatively broad and their utility is
limited. While they may enable subjects to assign labels to
musical tones, at a perceptual level, fine-grained differenc-
es in pitch are not meaningfully encoded. These data sup-
port the assertion that HTM is fundamentally a skill of
retrieval and comparison rather than a basic property of
perceptual encoding.
5. Does AP confer superior auditory memory?
Another facet of the mythology of ‘‘perfect pitch’’ is
that possessors of this trait may have an increased mem-
 D.A. Ross et al. / Epilepsy & Behavior 7 (2005) 578–601
ory for musical stimuli. This belief may be fueled, in
large part, by the performance of certain savant musi-
cians [53–57]. One famous example appears in the story
of a trip by Mozart to the Vatican during which he
reputedly heard a single performance of AllegriÕs Mise-
rere and transcribed the entire piece from memory.
While the historical accuracy of this anecdote may be
difficult to verify, the myth remains popular and with
it the question: Are AP possessors predisposed to having
better musical memories?
As discussed above, there is an appreciable body of
literature demonstrating that AP possessors have
decreased decay rates for individual tones. In a situation
where one group has the ability to name the target item
and the other does not, such an effect is not particularly
surprising. Perhaps for this reason, few investigators
have explored differences between AP and NAP individ-
uals in remembering sequences of tones or other extend-
ed musical stimuli. The myth remains untested.
We have presented a model of AP in which APE pos-
sessors differ from ‘‘normal’’ individuals, and from pos-
sessors of HTM, by the presence of an additional stream
of auditory information that may parallel the ordinary
RP processing pathway. In this scenario, APE individu-
als possess an extra channel carrying information about
stimulus frequency to the cortex. As such, one might
reasonably expect that they should have an increased
capacity to remember musical stimuli; a greater overall
amount of information about auditory stimuli is trans-
mitted to the cortex. This might also account for subjec-
tive perceptual differences described by AP and NAP
musicians. In our own informal surveys, the vast major-
ity of NAP musicians reported that when listening to a
complex piece of music, they can attend to only a single
aspect of the piece (e.g., in a symphony they could focus
on the flute, the violins, the chord progression, or some
other harmonic element, but not more than one of these
items at a time). In contrast, most of our APE musicians
report that it is relatively easy for them to keep track of
several streams at once. This ability is closely related to
the concept of preattentive processing (see below).
Unfortunately, we are unaware of any systematic efforts
to study differences in musical memory between AP and
NAP musicians. The goal of the present study was to
compare APE, HTM, and NAP musicians on a series
of memory tasks and to determine the extent to which
a subjectÕs memory depends on his or her ability to name
the stimulus in a conventional manner.
5.1. Memory for sequences of individual piano tones
Subjects were presented with sequences of one,
four, or seven individual piano tones followed by a
probe tone. Tones were taken from the range of G3
(196 Hz) to F#4 (370 Hz). Subjects rated their confi-
dence on a scale from 1 to 6 as to whether the probe
589
had been part of the preceding sequence. Approxi-
mately half of the time, the probe tone matched a tone
from the sequence; when it did, the target was always
presented in either the second or third position of the
sequence. When the probe did not match a tone from
the sequence, it always differed by a half-step from a
tone in one of the target positions. For each subject
and each condition, a ROC was again generated from
the confidence ratings, area under each ROC, p(A),
was calculated, and each groupÕs performance was
then compared with chance (p(A) = 0.5) using means
and SE.
Data from representative APE, HTM, and NAP sub-
jects are shown in Fig. 5. Group data for 12 APE, 17
HTM, and 9 NAP subjects are presented in [58]. The
data show a clear ordering: APE subjects performed
best, followed by HTM and then NAP subjects.
One might speculate that performance on the memo-
ry task depends directly either on subjectsÕ musical expe-
rience or on their ability to name notes. However,
extended analyses showed no correlation between per-
formance on this task and either of these parameters,
neither for within-group analysis nor for the combined
cohort. Thus, the ability to perform this task accurately
appears to reflect inherent group differences in the per-
ceived salience of pitch stimuli.
5.2. Memory for sequences of complex piano chords
In the first part of the experiment, stimuli were indi-
vidual piano notes. Consequently, APE and HTM sub-
jects might have had an advantage in that they could
label each tone and remember the appropriate label
(rather than having to rely on difficult relative pitch cal-
culations). The goal of the present experiment was to
present subjects with stimuli that would elude conven-
tional notation and thus eliminate any bias acting in
favor of individuals who can name notes. To this end,
piano chords were carefully designed so that they could
not be named according to any standard musical
conventions. Each chord consisted of six notes: the root,
the major third and fifth one octave above, and a
doubling of the root three octaves above the bass. The
other two notes were ‘‘blue’’ notes and consisted of
either the sixth or seventh below and either the ninth
or eleventh above the third/fifth cluster. Chords were
then constructed using every permutation of flatting
and sharping the accidentals (i.e., flat or natural sixth;
flat or natural seventh; flat, natural, or sharp ninth; nat-
ural or sharp eleventh). Thus, there were 20 possible
permutations for each chord root, giving a total of
240 possible chords. A sample chord would be: G2 –E3 –
B3 –D4 –A#4 –G5 (root–sixth–third–fifth–sharp ninth–root).
Approximately half of the time, the probe chord matched
a chord from the sequence. When the probe differed, it
differed only by a half-step in one of the two ‘‘blue’’ notes
590 D.A. Ross et al. / Epilepsy & Behavior 7 (2005) 578–601

Fig. 5. p(A) data for representative APE, HTM, and NAP subjects for sequences of 1, 4, and 7 stimuli.

of the chord; the root and the harmonically important
notes were always identical. Thus, an incorrect probe
for the sample above might be G2 –Eb3 –B3 –D4 –A# 4 –G5
(root–flat sixth–third–fifth–sharp ninth–root). Sequences
were randomly generated such that no chord root repeat-
ed within a sequence, and for each chord the specific con-
figuration of ‘‘blue’’ notes was randomly determined. All
other parameters were the same as described in the previ-
ous section.
Data for representative APE, NAP, and HTM sub-
jects are shown in Fig. 5. Data from the full cohort of
subjects are presented in [58]. Performance for all three
 D.A. Ross et al. / Epilepsy & Behavior 7 (2005) 578–601
groups differed from chance for the one chord condition;
for the four-chord condition, only the APE and HTM
performances were significantly better than chance; for
the seven chord condition, none of the groups differed
significantly from chance.
As above, there were no significant correlations
between musical experience or note naming ability and
p(A) for any group at any sequence length.
5.3. Discussion
In the first experiment there were clear differences
among APE, HTM, and NAP groups. One might
speculate that APE and HTM subjects performed bet-
ter because they could label the component tones of
each sequence. This undoubtedly played some role in
their improved performance compared with the NAP
controls. However, data from the regression analysis
of p(A) and note naming ability should temper this
simple conclusion. Within the group of individuals
with ‘‘perfect pitch,’’ there was no correlation between
these two variables. This is somewhat less surprising
when one considers the speed of the sequences: the
interstimulus interval of 800 ms is approximately as
fast as our best note namers (cf. [31]) and consider-
ably faster than the majority of HTM subjects
[20,44]. Instead, these data suggest that performance
on the paradigm may reflect fundamental differences
in the perceived salience of stimulus frequency. They
also support the widespread belief that individuals
with ‘‘perfect pitch’’ have better memories for simple
musical stimuli.
The second part of the experiment was designed to
test whether APE and HTM possessors have a superi-
or memory for musical stimuli even when it is not
possible to label them using any conventional nomen-
clature. To meet this end, the chords were designed to
be foreign: it is highly unlikely that a subject had ever
heard any chord like them and conventional names
for them do not exist. In an isolated context perhaps
it would have been possible for highly trained musi-
cians to determine the ‘‘quality’’ of the chord. Howev-
er, in a random context and with a total of only
800 ms per chord, this is implausible. As a qualitative
check, subjects were asked, after the completion of the
last task, if they had ‘‘figured out’’ how we had
designed the stimuli; while many subjects appreciated
that the changes always took place in the inside voic-
es, none was able to describe accurately the formula
we had used.
Despite the imposing difficulty of the task, both the
APE and HTM groups performed significantly better
than chance in the four-chord condition. As before,
performance on this task did not correlate with either
years of experience or the ability to name notes. These
data support the hypothesis that AP possessors are
591
disposed to having an increased ability to remember
musical stimuli. For the APE group, this effect closely
matches our initial prediction. For the HTM group,
we were surprised to find that they performed almost
as well as the APE musicians. Perhaps subjects with
HTM performed well because the stimuli comprised
chords with many notes presented simultaneously in
what might be called a ‘‘timbral mass.’’ If HTM pos-
sessors are defined by their ability to extract pitch
from complex timbral cues, particularly given that
the stimuli were all played on a piano, it is plausible
that they were able to transfer some degree of abso-
lute identification skills to the identification of the
pitch complexes.
In conclusion, these data show clear differences in
the ability of APE, HTM, and NAP individuals to
remember short musical sequences. Performance on
these tasks did not correlate with subjectsÕ ability to
name notes. Impressively, both APE and HTM sub-
jects performed significantly better than chance with
sequences of four chords that were specifically
designed to be unnamable. These data are the first test
of overall musical memory skills in AP possessors and
give credence to the popular belief that people with
‘‘perfect pitch’’ have an inherently greater ability to
remember musical stimuli. These findings are consis-
tent with the assertion of our model that stimulus fre-
quency may be a more salient feature of perception
for APE and, to a lesser extent, HTM possessors.
6. Preattentive processing of pitch and timbre
As discussed above (Section 2.3.1), a significant
body of research has explored the concept of preatten-
tive processing in the visual domain. As initially
described, ‘‘features’’ of visual perception are thought
to reflect the basic stimulus attributes encoded by the
visual system [32]. Despite major progress in this field,
surprisingly little research has been dedicated to
exploring this phenomenon in other domains, such
as the auditory. From this perspective, the question
remains open: What are the basic features of audition?
Obvious candidates include: loudness, height, timbre,
and, of greatest interest to the present research, pitch.
For pitch, several studies have shown preattentive pro-
cessing in tasks requiring response to ‘‘oddball’’ stimuli,
which differ saliently from others in a sequence [59–61].
While these data suggest that pitch differences may be pro-
cessed preattentively, they do not address the question of
whether pitch, per se, may be considered a basic feature of
perception. Given that the absolute frequency of a stimu-
lus has little (if any) salience to most individuals, it would
be counterintuitive to qualify it as such. Perhaps, as dis-
cussed above (and originally hinted at by Bachem), height
may be a feature of perception in normal individuals,
592 D.A. Ross et al. / Epilepsy & Behavior 7 (2005) 578–601
whereas some basic difference in signal processing makes
chroma an additional feature of perception in AP
possessors.
One reason that research into preattentive process-
ing in audition has proceeded so slowly may be the
difficulty of adapting classic paradigms to the auditory
domain. For example, one standard test of preatten-
tive processing is to measure search time for a target
in a visual field: if the target is preattentively pro-
cessed, RT should be largely independent of the num-
ber of distractors. The obvious auditory analog of this
task would be to translate visual space into time: thus,
one might present a target and then ask subjects to
signal when that target was presented during a
sequence of interfering tones. However, several prob-
lems arise. Foremost, a memory component is neces-
sarily introduced into any task spread out over time.
Second, a paradox emerges from the way the target
is initially defined: if subjects are asked to search for
an item by name (e.g., respond to each ÔAÕ), then
the task could be performed only by experienced
musicians with AP; if, however, the desired target is
presented at the start of each sequence, then it be-
comes impossible to prevent the task from being per-
formed on the basis of relative pitch. Thus, with this
type of paradigm, there is no obvious way of testing
the extent to which normal individuals preattentively
process the absolute pitch of stimuli.
An alternative paradigm of preattentive visual pro-
cessing requires subjects to focus on a central image
and then evaluates their ability to process peripheral
stimuli. According to the theory, only preattentive fea-
tures (e.g., color, orientation) can be perceived in this
manner. For example, a peripherally presented red ver-
tical line could be perceived either as red or as vertically
oriented, but not both. To do so would require the
assembly of features into a complex object, which can
only happen with overt attention. This type of paradigm
is more easily adaptable to the auditory domain and
allows us to ask the more explicit question: To what ex-
tent do individuals encode representations of stimulus
frequency (as opposed to differences in pitch) when they
are not attending to the target?
The following experiments were designed to test the
extent to which APE, HTM, and NAP individuals pre-
attentively process the pitch and timbre of auditory
stimuli. Our model predicts that APE possessors should
be very good at this task, whereas NAP individuals
should be relatively poor. To the extent that HTM sub-
jects ‘‘automatically’’ process the targets (i.e., recognize
them because of their similarity to items in their memo-
rized templates) they could outperform the NAP con-
trols; however, this form of processing should be both
inefficient and inaccurate as compared with true preat-
tentive processing. For timbre, we did not expect to find
any differences between groups.
6.1. Preattentive processing of pitch or timbre sequences
Eleven APE, 21 HTM, and 13 NAP controls partici-
pated in the following experiment.9 On each trial, sub-
jects were instructed to remember a series of six
images.10 Coincident with each image a tone was pre-
sented. The tonal sequences varied in either pitch or tim-
bre (e.g., six different pitches played on a piano or six
different instruments playing middle ÔCÕ). Following
the sequence two separate probe images were presented.
For each one, subjects were required to respond yes/no
as to whether that image had been part of the sequence.
Following the probe images a probe tone was presented
and subjects were required to rate their confidence (1–6
scale) as to whether the tone had been part of the pre-
ceding sequence. Probe tones were part of the sequence
approximately half of the time; when they were, the tar-
get was always presented in either the second or third
position of the sequence. Subjects were explicitly
instructed that their first priority was to remember the
visual stimuli accurately and that their data would be
thrown out if they failed to do so.
Data were first analyzed for visual accuracy and any
trial in which a subject responded incorrectly to a probe
image was excluded.11 Visual accuracy (VA) was defined
as the proportion of trials in which a subject responded
correctly to both probe images (i.e., chance VA = 0.25).
Data were then separately analyzed for timbre and pitch
sequences, and ROC curves were generated from the
confidence rating data. Data were quantified by calcu-
lating p(A)pitch and p(A)timbre for each subject. ANOVAs
were conducted with these data as the dependent
variable.
APE, HTM, and NAP subjects had VAs
(mean ± SE) of 0.64 ± 0.03, 0.65 ± 0.02, and
0.67 ± 0.03. ANOVA with VA as the dependent vari-
able showed no effect of group [F (2, 42) = 0.58; ns].
9
APE: 6 male; average age of 20.6 ± 0.7 years; average of 14.5 ± 0.7
years of experience playing a musical instrument, HTM: 5 male;
average age of 20.6 ± 0.6 years; average of 14.3 ± 1.0 years of musical
experience, NAP: 7 male; average age of 19.4 ± 0.3; average of
10.1 ± 0.7 years of experience on their primary instrument.
10
Images were presented for a duration of 1500 ms with an
interstimulus interval (ISI) of 250 ms. Stimulus tones were presented
for 600 ms and were generated using EasyBeat MIDI software. Tones
were selected from a set that consisted of each note of the ET scale
from C4 (261.6 Hz) to B5 (987.8 Hz) played on each of 20 diverse
timbres. Six blocks consisted of 20 trials each and for each block a
different category of images was used (including beetles, green
minerals, trees, geometric shapes/colors, warblers, and stamps).
Subjects were encouraged to rest as necessary between blocks.
Completion of all six blocks typically required between 45 minutes
and an hour and subjects performed two complete cycles. Identical
Psyscope scripts were used for all subjects.
11
One AP subject was colorblind and had a particularly difficult time
performing the visual tasks. For this subject, data were included for all
trials, regardless of his visual accuracy (though he was given the same
instructions as the other subjects).
 D.A. Ross et al. / Epilepsy & Behavior 7 (2005) 578–601
Fig. 6. Average ROC curves for APE, HTM, and NAP subjects for preattentive processing of pitch.
For the pitch sequences, APE, HTM, and NAP
groups performed at (p(A) ± SE) 0.85 ± 0.02,
0.81 ± 0.01, and 0.75 ± 0.02, respectively (see Fig. 6).
All of these values were significantly better than chance
(P < 0.01). For the timbre sequences,12 APE, HTM, and
NAP groups performed at: 0.74 ± 0.03, 0.74 ± 0.02, and
0.76 ± 0.02, respectively (see Fig. 7).
ANOVA showed a significant effect of task
[F (1, 39) = 18.14, P < 0.0001] but not group
[F (2, 39) = 1.30, P > 0.20]. The interaction between task
and group was highly significant (F (2, 39) = 5.17,
P = 0.01). These data show that APE, HTM, and
NAP subjects performed differently on the pitch and
timbre tasks: as predicted, there was a clear order effect
(APE > HTM > NAP) for the pitch task but no group
differences for timbre.13
12
Three subjects (two AP, one HTM) misunderstood the task and
thought that targets need only correctly match the pitch of an item- as
such, they responded ‘‘6’’ to all timbre sequences. Accordingly, data
from these subjects have been excluded from the timbre analysis.
13
Despite the magnitude of this effect, we believe these data may
actually give a falsely low impression of APE performance. In point of
fact, there were two outlier individuals who performed significantly
worse than the rest of the group. One was an individual who
performed poorly in the seven note condition of the previous
experiment and clearly had lower overall memory abilities. At a later
date, we questioned the other outlier about his subpar performance
(relative to his performance on the rest of our tasks) and he reported
that he had not felt well the morning of the experiment. He agreed to
retake the test and was rerun using a new script. On the retest he had a
p(A) of 0.97 for pitch, which was the highest of any subject (compared
with 0.74 during his first session). Another interesting overall indica-
tion of APE performance is that eight of the top nine scores were by
APE possessors.
593
How might we explain the results? The most obvious
explanation is that the APE subjects have an increased
ability to preattentively process pitch. In fact, the data
closely correspond to the two major predictions of our
theory: increased performance on the pitch task for
the APE group and equal performance across groups
for the timbre task. However, several alternate explana-
tions exist.
First, it is possible that the visual task did not fully
maintain subjectsÕ attention; in this case the paradigm
would become a divided attention task rather than one
of preattentive processing, per se, and a trade-off would
be expected between visual and auditory performance.
Regression analysis of p(A) as a function of visual accu-
racy showed this not to be the case for any group at
either task. This explanation also seems less likely given
the nearly identical performance of all three groups on
the visual task.
A second possibility is that performance on the audi-
tory tasks depends directly on subjectsÕ ability to label
the auditory stimuli. Regression analyses of APE and
HTM data showed no correlation, however, between
note naming ability and either p(A)pitch or p(A)timbre.
A third explanation is that APE possessors are some-
how endowed with an enhanced ability to attend to mul-
tiple tasks: thus, rather than reflecting a difference in
preattentive processing, the data could reflect a more
generalized increase in attentional resources for auditory
stimuli. To some extent, this hypothesis is refuted by the
failure to find group differences in the timbre task. Nev-
ertheless, a more rigorous test would be to show that
pitch and timbre perception in the present tasks truly
qualify as preattentive. As described above, a hallmark
594 D.A. Ross et al. / Epilepsy & Behavior 7 (2005) 578–601
Fig. 7. Average ROC curves for APE, HTM, and NAP subjects for preattentive processing of timbre.
of preattentive processing is that subjects may perceive
features individually but are incapable of fusing multiple
features into an object without overt attention. Thus, a
control for the above experiment is to test how well sub-
jects can preattentively process stimuli that vary based
on both pitch and timbre. If the APE group maintained
elevated performance for such an ‘‘object’’ task it would
suggest either: (1) that the paradigm does not test preat-
tentive processing (i.e., that accuracy is proportional to
subjectsÕ ability to divide their attention between visual
and auditory tasks) or (2) that pitch and timbre may
not rightfully be considered as discrete preattentive fea-
tures of auditory perception.
6.2. Preattentive processing of combined pitch/timbre
sequences
Five APE, seven HTM, and five NAP musicians were
recruited from the previous study to take part in the fol-
lowing experiment. One additional NAP subject was
recruited.14
In the previous experiment tonal sequences were
designed to vary in either pitch or timbre. To test how
well subjects can perceive conjoined features, sequences
in the present experiment were constructed so as to vary
in both pitch and timbre simultaneously. Thus, six dif-
ferent pitches were presented, each of which was played
14
Performance of these subjects was generally comparable to that of
the larger groups described in Section 2. APE, HTM, and NAP
subjects had average VAs of 0.69 ± 0.05, 0.58 ± 0.05, and 0.66 ± 0.04;
average p(A)pitch of: 0.89 ± 0.02, 0.85 ± 0.03, and 0.76 ± 0.03; and
average p(A)timbre of 0.81 ± 0.02, 0.78 ± 0.02, and 0.80 ± 0.02.
on a different instrument. For trials where the probe was
present, the target was either in the second or third posi-
tion. When it was not, the second and third tones each
matched one of the parameters. Thus, in all sequences
subjects heard both the pitch and timbre of the probe
during the main sequence; the question they were being
asked was whether they heard the two as a conjoined
object.
In this experiment, subjects performed six blocks of
15 trials each. All other parameters were the same as de-
scribed above.
APE, HTM, and NAP subjects had average VAs of
0.71 ± 0.05, 0.58 ± 0.04, and 0.74 ± 0.07. These values
were similar to their performance in the previous task.
Average ROC curves for the object task are shown
for each group in Fig. 8. As predicted, subjects per-
formed significantly worse in the present condition: the
APE, HTM, and NAP subjects performed at average
p(A) of 0.62 ± 0.03, 0.67 ± 0.04, and 0.66 ± 0.04. An
ANOVA with task (pitch vs object) as a within group
factor showed a significant effect of task
[F (1, 15) = 100.35, P < 0.0001] but not of group
[F (2, 15) = 1.88, P > 0.15]. The interaction between
group and task was significant [F (2, 15) = 6.53,
P < 0.01], reflecting a greater effect of task in the APE
group.
6.3. Discussion
Data from these experiments show that APE subjects
are better than either HTM or NAP subjects at preatten-
tively processing pitch. This finding cannot be accounted
for by differences in note naming ability or musical
 D.A. Ross et al. / Epilepsy & Behavior 7 (2005) 578–601
Fig. 8. Average ROC curves for APE, HTM, and NAP subjects for combined pitch/timbre processing.
experience. Further, poorer performance for the object
task rules out the possibility of baseline differences in
either memory or attention. It remains unclear whether
the APE groupÕs poor performance in the object task
results from a difference in strategy (i.e., they tried to
use pitch and timbre information) or a limited ability
to make use of height cues (perhaps secondary to lack
of use in ordinary life). Nevertheless, taken as a whole,
these data strongly support the hypothesis that the dom-
inant mode of perception in APE subjects is to process
pitch and timbre as separate features that cannot be
fused without overt attention.
Data from the NAP group provide a fascinating
counterpoint. One surprising aspect of their baseline
data was that NAP subjects performed comparably for
pitch and timbre tasks and that performance was signif-
icantly better than chance for both. If only APE possess-
ors are able to preattentively process absolute pitch, why
did a group of control musicians perform so well on this
task?
The NAP groupÕs performance on the pitch task likely
reflects limitations of the paradigm itself. To a certain ex-
tent, it was possible to perform the task on the basis of
height only. For example, if a sequence fell mostly within
the fourth octave and the probe was in the fifth octave,
no knowledge of absolute frequency would be required
to respond correctly. A second factor to be considered
is that the timbre of an instrument is known to vary con-
siderably across its natural range, even within intervals
as small as a major third [45]. This effect is likely to be
particularly prominent given the irregular nature of the
timbres we used. Thus, the combination of cues from
the height and timbre of the stimuli provides a consider-
595
able amount of information about each note: subjects
may have been able to identify many of the tones with
sufficient precision as to be able to perform the task with-
out needing to recognize the pitch of the target, per se.
From this perspective, the equivalence of NAP perfor-
mance for ‘‘pitch’’ and ‘‘timbre’’ tasks is not as surpris-
ing; in point of fact, performance for both tasks may
depend strongly on timbral cues. Similarly, performance
on the object task suggests that NAP subjects may have
used some sort of fused height/timbre representation.
Suffice it to say, future research elaborating the basic fea-
tures of auditory perception in ‘‘normal’’ individuals
would be extremely useful.
Of note, an important historical difficulty with these
types of experiments (including our own) is that many
‘‘pitch’’ tasks are susceptible to subjectsÕ exploitation
of height cues. One possible way to circumvent this con-
found would be to present sequences across a wider fre-
quency range and define ‘‘correct trials’’ as those in
which the chroma of a target was presented in a different
octave (e.g., for a sequence that included C3, a probe of
C4 would be considered a ‘‘hit’’). Such a manipulation
would eliminate height cues and allow one to test more
precisely the extent to which stimulus chroma is pro-
cessed as a feature. Based on our theory, we would pre-
dict even larger effects than seen in the present article:
NAP performance should fall to chance whereas APE
performance should be unaffected.
Performance by the HTM subjects is more difficult to
interpret. Data from the first part of the experiment con-
firm that HTM subjects performed better on the pitch
task than normal controls, though not as well as the
APE subjects. As in the APE group, performance was
596 D.A. Ross et al. / Epilepsy & Behavior 7 (2005) 578–601
markedly diminished in the object task. These data sug-
gest that the difference in HTM processing may be con-
nected to the ability to exploit height/timbre cues.
Unfortunately, the present experiment was not designed
to test the nuance of our modelÕs distinction between
APE and HTM. It would be interesting to explore this
question at greater length in future experiments.
One test would be to compare performance of APE
and HTM subjects for stimuli with common or rare tim-
bres. To the extent that the recognition of pitches by
HTM possessors depends on a targetÕs similarity to a
memorized template, we would expect the group to per-
form better for familiar timbres. Another follow-up
would be to replicate the object experiment with similar
manipulations in timbre. Again, to the extent that HTM
subjects have memorized specific tones (e.g., an A4 on a
violin) they should be better able to recognize these
fused objects. In the interim, however, these data contin-
ue to support the distinction we have drawn between
APE and HTM individuals.
7. Discussion
For more than 100 years, musicians and scientists
alike have been fascinated by the enigmatic phenome-
non of ‘‘perfect pitch.’’ Yet for all of the attention it
has received, many aspects of this condition have re-
mained largely unresolved; conflict persists regarding
its etiology and its musicosocial significance, and, at
the most fundamental level, there is no consensus on
how the condition should be defined, let alone how best
to account for heterogeneity among its possessors.
In this article, we propose a new model of AP that
may help reconcile conflicting historical perspectives.
In doing so, we attempt to reframe a phenomenon that
has been explored almost exclusively from a behavioral
perspective within the context of modern research on
low-level mechanisms of pitch perception. Our new
model arose from the qualitative observation that indi-
viduals who can name notes with great facility constitute
a heterogeneous population. As such, the model subdi-
vides AP possessors into two groups based on proposed
differences in the mechanisms used to encode pitch. We
define APE as the ability to perceptually encode stimu-
lus frequency. We argue that this skill may result from
fundamental differences in the processing of periodotop-
ic pitch. This skill is automatic and independent of any
musical experience. Further, subjectsÕ ability to encode
stimulus frequency is independent of the specific nature
of the target.
In contrast, we define HTM (heightened tonal mem-
ory) possessors as those individuals with an increased
ability to form and evoke memories of specific complex
frequency stimuli. These individuals may be able to
identify the AP of targets by comparing them to a
template of memorized items. The more familiar the tar-
get stimulus, the easier it should be for an HTM posses-
sor to recognize (and encode a long-term memory of) it.
Broadly speaking, our definition of APE corresponds to
the definition of AP described by the innate model and
our definition of HTM corresponds to that of the
ELT. The present experiments were designed to test
both the explicit predictions of our new model, and a
series of historical myths and dogmas in the field. Fore-
most among these is the assumption, held by advocates
of both the early learning and innate models, that AP is
fundamentally a musical phenomenon.
Data from these experiments showed that: (1) APE
and HTM subjects were equally adept at encoding
long-term representations of HP stimuli (relative to their
own baseline), (2) APE subjects were not only more
accurate than HTM subjects at distinguishing between
properly and improperly tuned stimuli, but they were
more sensitive to fine-grained differences in frequency;
(3) There were clear group differences in subjectsÕ ability
to remember simple musical sequences; more impres-
sively, both APE and HTM subjects performed signifi-
cantly better than chance at remembering sequences of
chords that could not be identified using any conven-
tional nomenclature, (4) APE and, to a lesser extent,
HTM possessors have an increased ability to preatten-
tively process pitch but do not differ from NAP controls
in preattentive processing of timbre; follow-up data
from an ‘‘object’’ task support the idea that APE sub-
jects perceive pitch and timbre as distinct features of per-
ception, NAP subjects perceive height and timbre (but
not pitch, per se) as basic auditory features, and HTM
subjects may extract information about AP from height
and/or timbral cues.
7.1. Reevaluating our model: Are there two different types
of ‘‘perfect pitch’’?
One might object that our distinction between APE
and HTM is arbitrary, i.e., that the groups we defined
are simply part of the same continuous spectrum and
result from the same underlying mechanism. In contrast
to this hypothesis, not only did we observe major differ-
ences between the groups, but for each experiment there
was a significant interaction between the main experi-
mental manipulation and AP status (i.e., APE vs
HTM). Additionally, while the ability to name notes
would be an obvious way to define a continuous spec-
trum of AP ability (as has been used by virtually every
other research group), there was no correlation between
performance on our gold standard note naming task and
any of the other output variables measured, either with-
in or across groups. Instead, the only factor that accu-
rately predicted performance was the one that we used
to define the groups in the first place, namely, the ability
to encode accurate long-term representations of
 D.A. Ross et al. / Epilepsy & Behavior 7 (2005) 578–601
stimulus frequency. These data provide overwhelming
support to the legitimacy of our distinction between dif-
ferent types of ‘‘perfect pitch’’ with different underlying
mechanisms.
7.1.1. (Re)defining APE
According to our theory, APE subjects are distin-
guished by a low-level difference in the pathway used
to process periodotopic pitch. Specifically, we argue that
they may be uniquely able to access the low-level repre-
sentation of pitch that is thought to emerge via an auto-
correlative process but is inaccessible in ‘‘normal’’
individuals. This hypothesis may be difficult to prove
in any conventional, noninvasive manner; nevertheless,
the present data support it in several important ways.
First, data from the HP experiment show that APE
subjects are not limited in their ability to encode repre-
sentations of purely periodotopic pitches; these data
confirm that APE subjects are, at the very least, able
to encode pitch via some mechanism in the temporal
pathway. More significantly, data from the other exper-
iments strongly corroborate our qualitative description
of APE. Possessors of APE appear to encode the pitch
of stimuli immediately and absolutely. This process is
independent of either attention (see Section 6) or the
ability to name notes (as evidenced by lack of correla-
tion between performance on the gold standard note
naming test and any other tasks). Further, the represen-
tations of pitch formed by these subjects fall along a
continuous spectrum within which subjects can resolve
differences a fraction of the size of the smallest meaning-
ful unit in Western music (see Section 4).
Further experiments should be aimed at measuring
neurophysiological signals so as to test the proposed
mechanism more directly. Useful methods may include
MEG, EEG, or brainstem imaging with functional
MRI. If our model is correct, one might expect to see
differences between APE and NAP individuals as low
in the auditory pathway as the cochlear nucleus or supe-
rior olive.
7.1.2. (Re)defining HTM
Our model defined HTM as an enhanced ability to
memorize and retrieve specific configurations of audito-
ry features from which subjects could infer AP informa-
tion. The system used to process timbre was
hypothesized to play a prominent role. By this defini-
tion, an underlying difference distinguishing HTM pos-
sessors from NAP controls, if it exists, could lie in
either the spectral or temporal pathway or at a higher
level, after the two streams have been reintegrated.
The present data help refine this original definition.
At a qualitative level, performance by the HTM group
in the intonation task supports the idea that subjects
may encode stimulus frequency by comparing targets
to a memorized template and that this template may
597
be relatively crude and idiosyncratic across subjects.
Second, as discussed with the APE subjects, accurate
performance with the HP stimuli contradicts the pri-
mary importance of any spectral processing mecha-
nism. Other data (including those obtained in what
is dubbed a ‘‘Garner interference paradigm’’ [31]) lend
further support to a mechanistic distinction between
HTM and NAP individuals. Finally, the most reveal-
ing finding for this group may be subjectsÕ perfor-
mance on the preattentive pitch, timbre, and object
tasks: while subjects demonstrated a moderately
increased ability to process pitch preattentively, they
were not as inhibited as the APE subjects in their abil-
ity to process conjoined pitch/timbre objects. This
finding supports the idea that HTM subjects do not
process pitch as an independent feature of audition;
rather, as argued above, their ability to process pitch
appears closely tied to the height and/or timbre of
the stimulus.
As a whole, these data support the idea that HTM
subjects differ from APE and NAP possessors at a rela-
tively high level in the auditory pathway: specifically,
they seem better able to memorize and retrieve complex
stimuli that are integrated across multiple auditory
streams. The greater the number of familiar features
that define a stimulus, the easier it seems to be for them
to recognize it. An interesting quirk of the behavior of
many of our HTM subjects was the importance of
motor memory in trying to recall a specific pitch. For
example, many of the string players would ‘‘finger’’ a
note that they were trying to either name or remember.
Other subjects reported being able to name notes only
while they were holding their instrument in their hands.
These types of motor cues seemed to facilitate subjectsÕ
auditory imagery for tones. These observations not only
support the qualitative description of HTM as depen-
dent primarily on retrieval mechanisms, but they also
suggest that the critical pathways involved may be quite
high in the auditory pathway, perhaps involving cortical
memory systems that can allow for integration both
within and across modalities.
7.1.3. Two groups or three?
It should be emphasized that these data point to a
fundamentally different process used by APE and
HTM possessors to encode pitch. While the former
group is distinguished by a difference in the ability to
encode periodicity pitch, the HTM group is distin-
guished by an increased ability to form and retrieve
memories of specific complex stimuli. As such, a major
unanswered question is whether, in the spirit of John
Watson, we can take any child and provide sufficient
training to induce HTM, or whether some kind of innate
predisposition is necessary. This question is equivalent
to asking whether the difference between HTM and
NAP individuals is one of type or of degree.
598 D.A. Ross et al. / Epilepsy & Behavior 7 (2005) 578–601
It seems likely that the appropriate type of training,
particularly at a young age, may lead to the formation
of a template in virtually any individual. However, some
individuals may be predisposed to either: (1) being more
susceptible to forming templates from specific auditory
events; or (2) being better able to retrieve memories from
their templates. Either of these possibilities would lead
to greater fluidity in naming notes: the former by creat-
ing a more thorough template and the latter by allowing
easier recognition of tones that are similar to, but do not
exactly match, items in a subjectÕs template. At this
point, while it is clear that differences exist between
NAP and HTM groups, the relative roles of innate
and epigenetic factors remain ambiguous.
While our definition of HTM resembles the ELT in
many ways, we must emphasize that it differs in the most
critical one: though the ability of this group to recognize
pitches may be based on a memorized musical template,
we believe that the ability to name notes is a symptom
rather than a cause of the condition. A salient feature
of HTM, considered broadly, is an increased ability to
memorize and retrieve specific complex frequency stim-
uli—a skill that may facilitate note naming if items in
the template are paired with musical labels, but may also
be relatively independent of labeling. Two major factors
have led researchers to overemphasize the importance of
attaching labels to items in a subjectÕs template. First,
classic AP tests screen musicians only, a group in which
musical labels are likely to be more important. More sig-
nificantly, these types of paradigms explicitly ask sub-
jects to apply musical labels to targets; thus, the extent
to which unlabeled representations exist in subjectsÕ tem-
plates has rarely been examined. Given this context,
however, the recent data showing that ‘‘normal’’ indi-
viduals remember the starting pitches of their favorite
songs with disproportionately great accuracy are of par-
ticular interest in that they provide clear evidence for a
dissociation between the ability to memorize/retrieve
specific complex auditory stimuli and the need to overtly
label these representations.
Because most of the present experiments sought to
test differences between APE and NAP subjects, further
experiments that are specifically aimed at characterizing
the HTM group could be illuminating. As discussed
above, MEG, EEG, and fMRI paradigms would be
helpful in clarifying the relative importance of different
neural systems, which could occur anywhere between
brainstem nuclei and parietal association cortex. At a
behavioral level, expanded testing could elucidate the
extent to which note naming ability in HTM subjects
is a direct outgrowth of the tonal memory that has been
observed in NAP controls.
7.1.4. Qualitative observations on APE and HTM
Although subjects were classified as APE or HTM
based on their performance on our reproduction tasks,
ultimately, our model was formulated based on qualita-
tive observations and discussion with subjects. During
the course of testing, all subjects were surveyed with a
standardized set of questions including: When did you
first realize you had ‘‘perfect pitch’’? and Are there
any features which make it easier or harder for you to
recognize tones? For virtually all of our subjects,
responses to these questions gave a clear indication of
whether they possessed APE or HTM.
APE subjects typically could not recall a time when
they did not have AP: frequently, they would respond
to this question by recounting the epiphany in which
they realized not that they had AP, but that everyone
else did not. Familiarity effects in this group were gener-
ally minor; subjects reported being equally able to name
musical or nonmusical stimuli, and few reported any
changes in pitch perception over time. One of the most
remarkable features of these subjects was their qualita-
tive assessment of our paradigm: the majority of them
reported that there was no difference in difficulty
between reproducing a tone following interstimulus
intervals with or without interference.
In contrast, many of the HTM subjects recounted
stories of having learned or taught themselves ‘‘perfect
pitch’’ or how it had slowly evolved over time. Virtually
all of them described a set of parameters that facilitated
note identification and many claimed to have ‘‘perfect
pitch’’ only for their primary instrument or only for
some notes. A typical story would be that after having
played piano for 10 years, the subject spontaneously
realized that he or she recognized any note played on
a piano. Then, gradually, he or she gained the ability
to name notes on other familiar instruments. These sub-
jects typically found it quite difficult to remember a tone
after extended interference.
7.2. On the mythology of ‘‘perfect pitch’’
7.2.1. Myth 1: AP is a musical phenomenon
Perhaps because it is such a striking and valuable skill
in a musical setting, AP has historically been defined
strictly as the ability to name notes. As such, advocates
of both early learning and innate models have taken for
granted that the skill is limited to the musical domain. In
fact, to the best of our knowledge, no one has previously
attempted to define AP within the broader context of
auditory perception, nor has anyone devised a paradigm
that could test for AP independent of a subjectÕs musical
experience. Recently, we showed not only that the latter
of these goals was feasible, but that there exist individu-
als who cannot name notes accurately but who neverthe-
less possess AP. These data show that AP may be
independent of a subjectsÕ musical experience. The pres-
ent data extend these findings.
We show that APE and HTM possessors differ from
NAP musicians across a wide range of auditory tasks,
 D.A. Ross et al. / Epilepsy & Behavior 7 (2005) 578–601
some of which may be considered ‘‘musical,’’ others of
which cannot. None, however, relates directly to the
classic AP skill of note naming. The data show that
APE subjects have an increased sensitivity to fine-
grained pitch differences, an increased memory for tonal
stimuli, and an increased ability to preattentively pro-
cess pitch. These differences cannot be explained by note
naming ability. They also do not correlate with musical
experience. Rather, they demonstrate that APE possess-
ors perceive stimulus frequency in a fundamentally dif-
ferent manner from ‘‘normal’’ individuals.
A paradigm shift is in order: it is time to stop defining
AP based on a single, arbitrary, skill and to start explor-
ing the full range of perceptual attributes encompassed
by this phenomenon.
7.2.2. Myth 2: AP possessors identify stimulus frequency
by using a place code on the basilar membrane
Despite a significant body of recent research demon-
strating the relative importance of periodotopic signals
in pitch perception, researchers have persisted in assert-
ing that a simple place code on the basilar membrane is
the critical mechanism used to encode absolute pitch.
This discussion has been carried out largely within the
context of age-related changes in pitch perception in
AP possessors.
The present data show conclusively that AP percep-
tion can be independent of a cochlear place code. While
it is plausible that redundant mechanisms exist (one
spectral and one temporal) it is more parsimonious to
conclude that AP recognition derives from a periodo-
topic mechanism. A worthwhile line of inquiry might
therefore be to inquire how age-related changes might
affect the rate of firing in the cochlear hair cells.
7.2.3. Myth 3: AP may predispose individuals to special
musical skills
Within musical communities, it may often be accept-
ed, either implicitly or explicitly, that having AP may
help make someone a better musician. Obviously, musi-
cianship is multifaceted, and the dominant component
(at least in tonal music) is developing a strong sense of
relative pitch. Nevertheless, AP possessors are frequent-
ly rumored to have increased musical memory or an in-
creased ability to discern in-tune from out-of-tune
stimuli. While commonplace, these suggestions have
been deemed politically incorrect and vehemently reject-
ed by some members of the scientific community (gener-
ally, the same individuals who reject the similarly
politically incorrect idea that AP is innate).
The present data show an empirical basis for both of
these myths. APE possessors were not only able to accu-
rately discern in-tune from out-of-tune stimuli; they
demonstrated a vastly increased memory for simple
musical sequences. Neither of these skills correlates with
either the ability to name notes or with subjectsÕ musical
599
experience. Rather, they appear to reflect a difference in
the perceptual salience of stimulus frequency (as similar-
ly shown in the preattentive processing experiment).
These findings call for an expanded inquiry into other
potential differences between APE and NAP individuals.
Are there other group differences within the auditory
domain? Perhaps of greater interest, do differences
between APE and NAP individuals extend to other cog-
nitive realms (e.g., is the increased memory generaliz-
able?)? A more provocative topic to explore would be
the potential connection between AP and the nebulous
social construct of ‘‘genius.’’ At least among musical
‘‘geniuses’’ there is clearly a disproportionate incidence
of AP—from classical composers (e.g., Mozart and Bee-
thoven), to modern musicians (e.g., Yo Yo Ma, Wynton
Marsalis, and Bobby McFerrin), to special populations
such as autistic musical savants.
A noteworthy aspect of the present studies is that the
research was conducted at Yale University and the vast
majority of subjects were undergraduate students of
Yale College. An interesting aspect of this population
was that while almost all of them were excellent musi-
cians, few intended to pursue music professionally. In
fact, the subjects were quite diverse in their primary
areas of study, including journalism, psychology, and
biomedical engineering. Suffice it to say, while musically
rich, the environment was far from the typical conserva-
tory atmosphere from which most researchers recruit
their subjects.
Nevertheless, within this population the prevalence of
APE was extraordinarily elevated compared with any
conventional estimates (which generally range from
1/5000 to 1/10,000). Currently at Yale College, there
are no fewer than seven individuals with this trait (out
of a student body of approximately 4500), a prevalence
five to ten times greater than the most generous esti-
mates of AP. If we include HTM possessors in this cal-
culation (as published reports almost certainly do), there
are another 16 current undergraduates who were includ-
ed in this study plus at least half a dozen more ‘‘perfect
pitch’’ possessors of whom we are aware but have not
yet tested. Thus, in total, no less than 0.5%—and possi-
bly up to 1%—of Yale undergraduates possess some
form of ‘‘perfect pitch.’’ While the admissions commit-
tee may look favorably on applicants with musical expe-
rience, this proportion is still surprising and is worthy of
further consideration in its own right.
7.3. General implications
7.3.1. Models of auditory processing
The cochlea encodes stimulus frequency along two
orthogonal domains: spectrally, based on the location
of the point of maximum amplitude along the basilar
membrane, and periodotopically, based on the spike
intervals from outer hair cells. Researchers studying
600 D.A. Ross et al. / Epilepsy & Behavior 7 (2005) 578–601
low-level auditory processing tend to emphasize the
importance of periodotopic pathways in pitch percep-
tion. In contrast, researchers studying pitch processing
at the cortical level remain relatively fixated on the
importance of tonotopic representations. At first
glance, it is puzzling why researchers from these two
fields have been unable to reconcile their differing per-
spectives. On closer examination, though, the conflict
may be seen to result from the major irony of ‘‘pitch
processing’’ research: for the vast majority of individ-
uals, ‘‘pitch’’ is a meaningless concept. While precise
low-level representations of pitch may exist, at a cog-
nitive level they are inaccessible: a ÔBbÕ does not sound
any different than a ÔCÕ.
From this perspective, debate about the relative
importance of tonotopic and periodotopic processing
streams, like the debate between the early learning
and innate models of AP, may be moot: each model
appropriately describes a subset of the whole. Simple
‘‘pitch processing’’ tasks (e.g., high/low tasks for two
fixed tones) are likely performed on the basis of height
and processed via tonotopic pathways. More complex
‘‘pitch processing’’ tasks (e.g., roving high/low para-
digms or interval recognition) must be performed on
the basis of relative pitch information, which is likely
processed via periodotopic streams. On one level, this
distinction is supported by the respective methods
used by previous researchers to map tonotopic versus
periodotopic representations in the cortex. From a
theoretical perspective, the functional significance of
this distinction is supported by the fact that tonotopic
representations of pitch follow the Mel Scale (which is
consistent with height but not musical pitch), whereas
periodotopic information is naturally organized in
musically meaningful ways (where the degree of over-
lap between autocorrelograms correlates with the
musical overtone series). These distinctions should
reinforce the importance of eschewing generic descrip-
tors (such as ‘‘pitch processing’’) in favor of more pre-
cise terminology and in carefully specifying the exact
cognitive demands of each behavioral task.
7.3.2. Importance of preattentive processing
The present research was inspired by the observa-
tion that pitch was a salient feature of perception
for only a small subset of the population. The preat-
tentive processing model provides a valuable frame-
work for interpreting this type of observation. The
crux of the model is that behaviorally identifiable
‘‘features’’ of perception may reflect the mechanisms
of low-level stimulus processing. Thus, the presence
of a unique behavioral ‘‘feature’’ in one subset of
the population suggests that a basic neurobiological
difference may define this group.
Unfortunately, preattentive paradigms have been
largely unexplored outside the visual domain. Yet this
type of framework may be ideally suited to audition,
where the dominant features of perception are ill-defined.
Intuitively, one might speculate that auditory features
include height, timbre, and relative pitch intervals. The
present data confirm the importance of height and timbre
in ‘‘normal’’ auditory perception. More significantly, they
confirm the key tenet of our model, namely, that one sub-
set of the population may be distinguished by the percep-
tion of a distinct preattentive feature. These data point to
an innovative application of TreismanÕs methods. While
originally intended to elaborate mechanisms of ‘‘normal’’
perception, they may be equally valuable for identifying
or better characterizing other special populations.
Acknowledgments
This article reports results from a dissertation by
David Ross presented in partial fulfillment for the
Ph.D. at Yale University. The authors thank John
Culling, Ingrid Olson, and Bob Schultz for their collab-
oration on these experiments, and Thomas C. Duffy and
Tom Cantey for important musical insights.
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