This excerpt from

Language, Brain, and Cognitive Development.

Emmanuel Dupoux, editor.
© 2001 The MIT Press.

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24
The Biological Foundations of Music
Isabelle Peretz

All human societies have music. To our knowledge, this has always been
so. Unlike other widespread human inventions, such as the writing sys-
tems, music was not created by a few individuals and then transmitted
to others. Instead, music seems to have emerged spontaneously in all
forms of human societies. Moreover, this emergence is not recent in hu-
man evolution. Music apparently emerged as early as 40,000 to 80,000
years ago, as suggested by the recent discovery of a bone flute attributed
to the Neanderthals (Kunej and Turk, 2000). Thus, not only is music
ubiquitous to human societies, it is also old in evolutionary terms. Ac-
cordingly, music may pertain to the areas covered by human biology.
Surprisingly, the notion that music might have biological foundations
has only recently gained legitimacy.1 Over the past thirty years, music has
mostly been studied as a cultural artifact, while a growing body of re-
search, well represented by the work of Jacques Mehler, contributed to
the building of the neurobiology of language (following the pioneering
work of Eric Lenneberg, 1967, Biological Foundations of Language). For
most musicologists, each musical system could only be understood in the
context of its specific culture. Moreover, according to most neuroscientists
and psychologists, music served as a convenient window to the general
functioning of the human brain. However, recent evidence suggests that
music might well be distinct from other cognitive functions, in being sub-
served by specialized neural networks under the guidance of innate mecha-
nisms. The goal of this chapter is to briefly review the current evidence
for the premise that music is a part of the human biological endowment.
I begin with the best available evidence, or at least the evidence that
convinces me, that music cannot be reduced to an ephemeral cultural
436 I. Peretz

product. This material relies, in particular, on the detailed examination

of music-specific disorders that can occur either after brain damage at
the adult age or as an acquisition failure. Next, I briefly discuss the issue
of brain localization with special attention to expertise effects, since these
are related and enduring questions. Afterward, I mention some recent and
fundamental discoveries made on infants’ musical abilities and suggest a
few candidates for musical universal principles. Lastly, I conclude with
a few speculations on the biological functions of music.

Specialized Neural Networks for Music Processing

If music is biologically determined, then it is expected to have functional

and neuroanatomical specialization, as do all major cognitive functions,
such as language. Support for the existence of such specialized neural
networks is presently compelling. The major source of evidence is coming
from the functional examination of individuals who happen to suffer
from some brain anomaly and who, as a consequence, exhibit disorders
that selectively either disturb or spare musical abilities. Presently, there is
no evidence for music-specificity derived from the study of normal brains.
However, it will not be long in coming, since the patient-based approach
converges on the notion that music is subserved by neural networks that
are dedicated to its processing. By “dedicated neural structures,” I am
referring to neural devices that process musical information selectively
and exclusively. Support for their existence comes from essentially three
types of neuropsychological findings.
The first type of evidence lies in the classic neuropsychological dissocia-
tions that can be observed after an accidental focal damage in a mature
brain. For example, we have been able to document several cases whose
characteristic symptom was the loss of the ability to recognize and memo-
rize music. These patients retained the ability to recognize and understand
speech, as well as to identify common environmental sounds normally
(Peretz, 1996; Peretz, Belleville, and Fontaine, 1997; Peretz et al., 1994;
replicated by Griffith et al., 1997). The deficit can be remarkably selective.
For example, CN was unable to recognize hummed melodies coming
from familiar songs above chance. Yet, she could perfectly recognize the
lyrics accompanying the melodies that she failed to recognize (e.g., see
Peretz, 1996). Moreover, CN was also able to recognize the voice of
 The Biological Foundations of Music 437

speakers (Peretz et al., 1994) and the intonation of speech (Patel et al.,
1998). The existence of such a specific problem with music alongside
normal functioning of other auditory abilities, including speech compre-
hension, suggests damage to processing components that are both essen-
tial to the normal process of music recognition and specific to the musical
domain. The reverse condition, which corresponds to a selective sparing
of music recognition, has also been reported (e.g., see Godefroy et al.,
1995). In this latter condition, the lesion compromises both speech com-
prehension and recognition of familiar environmental sounds. Such cases,
suggesting isolated sparing of music recognition abilities, complement the
music-specific deficits, and provide the key evidence of double dissocia-
tion. This neuropsychological pattern is the prototypical signature of the
presence of specialized brain circuits.
Further neuropsychological evidence that is indicative of the domain-
specificity of music is apparent in studies of autistic people. The autistic
individual is generally more apt in the area of music in contrast to other
cognitive activities, such as verbal communication (e.g., see Heaton, Her-
melin, and Pring, 1998). Of note is that certain autistic individuals become
musical savants, a term which refers to the observation of high achieve-
ments in musical competence in individuals who are otherwise socially
and mentally handicapped (e.g., see Miller, 1989). The mirror image of
this condition would consist of individuals who are musically totally in-
ept, despite normal exposure to music and normal intelligence. Such indi-
viduals exist and are commonly called tone-deaf (see Grant-Allen, 1878,
for the first report). Affected individuals are born without the essential
wiring elements for developing a normally functioning system for music,
hence experiencing music as noise at the adult age while achieving a high
degree of proficiency in their professional and social life. A well-known
figure who was known to be tone-deaf was Che Guevara (Taibo, 1996).
We have been able to confirm the existence of this music-specific learning
disability in at least twelve other adults (Ayotte et al., in preparation).
The selectivity of their musical failure is striking. All these individuals
match the case of CN (the brain-damaged patient previously described)
with no evidence of brain lesions. Accordingly, these individuals should
be referred to as congenital amusics. These amusic individuals have above-
average language skills, being able to speak several languages without
accent. Moreover, developmental amusics have never been able to sing,
438 I. Peretz

dance, or to recognize music as simple as their national anthem, despite

sterile efforts to learn music during childhood. Their condition is the re-
verse condition of that of the musical savant syndrome, hence illustrating
exceptional isolation of the musical modules in the developing brain.
The third source of evidence that speaks for the existence of neural
networks that are dedicated to music comes from the examination of
epileptic patients. It is well known that in a few individuals, the pathologi-
cal firing of neurons, conductive to the epileptic crisis, will be elicited by
music exclusively. This form of epilepsy is called “musicogenic epilepsy”
(Wieser et al., 1997) and suggests that the epileptogenic tissue lie in a
music-specific neural region. This suggestion is consistent with the con-
tent of the descriptions elicited in vivo by electric stimulation of the brain
of epileptic patients before brain surgery (Penfield and Perot, 1963). Di-
rect electric stimulation of particular areas of the auditory associative
cortex of awake patients often produces highly vivid musical hallucina-
tions. Again, these music-specific experiences support the existence of
neural networks specialized for music.

Localization of the Music-Specific Networks

Taken together, neuropsychological explorations provide compelling evi-

dence for the existence of brain specialization for music. One important
implication of this observation is that music does not look like a parasite
or a byproduct of a more important brain function, such as language.
Although such a conclusion supports the notion that music is a biological
function, it is not sufficient. Brain specialization does not entail prewiring.
Music may simply recruit any free neural space in the infant’s brain and
modify that space to adjust it to its processing needs. These needs may
be computationally complex to satisfy and hence require free and plastic
neural tissue. This opportunistic scenario of brain organization for music
may respond to cultural pressure and not biological requirements. How-
ever, if this were true, a highly variable distribution of the musical net-
works should be observed across individuals. Depending on the moment,
quality, and quantity of exposure, various brain spaces might be mobi-
lized. Thus, if music is a brain “squatter,” localization should vary capri-
ciously across members of the same culture.
 The Biological Foundations of Music 439

A prewired organization must exhibit consistency in localization. The

primary auditory areas are systematically located in the Heschl’s gyri,
buried in the sylvian fissure; this holds for all humans including individu-
als who are born deaf. Similarly, brain regions that are dedicated to music
processing should correspond to a fixed arrangement. That is, brain im-
plementation of music networks should be similar in the vast majority
of humans, nonmusicians and musicians alike. Moreover, this organiza-
tion is not expected to vary as a function of the musical culture consid-
ered. Musical functions are expected to be similarly implemented in the
brain of an isolated Pacific islander, a Chinese opera singer, and a West-
ern fan of rap music. This is a very strong prediction, perfectly suited to
the exploitation of the new brain imagery techniques.
Although clear and straightforward, the demonstration of a similar
brain organization for music in all humans remains elusive. The only con-
sensus that has been reached today concerns only one component of the
music-processing system: the pitch contour extraction mechanism that
involves the superior temporal gyrus and frontal regions on the right side
of the brain (see Peretz, 2000, for a recent review). However, it remains
to be determined if this processing component is music-specific, since the
intonation patterns of speech seem to recruit similar, if not identical,
brain circuitries (Zatorre et al, 1992; Patel et al., 1998). Clearly, what is
needed at the present stage is a grid that allows specification of the pro-
cessing mechanisms that are essential to music appreciation, an ability
shared by all humans. Once these essential ingredients have been identi-
fied, their respective localization could be tracked down in the brain of
musicians and nonmusicians of different musical cultures. The research
agenda involved is dense and will only be briefly sketched in the next
section.

What Is the Content of the Music-Specific Neural Networks?

The common core of musical abilities, which is acquired by all individuals

of the same culture and which forms the essence of the musical compe-
tence acquired by members of other cultures, must be organized around
a few essential processing components that are the core of the brain spe-
cialization for music. It is these highly specialized mechanisms that are
440 I. Peretz

probably embedded in the neural networks specifically associated with

music, and which may be detected in neurological practice. Thus, there
is no need for all musical abilities to have initial specialization. Brain
specialization for a few mechanisms that are essential to the normal devel-
opment of musical skills should suffice.
I am proposing that the two anchorage points of brain specialization
for music are the encoding of pitch along musical scales and the ascribing
of a regular pulse to incoming events. The notion that a special device
exists for pitch processing in music has been developed in previous papers
(Peretz and Morais, 1989, 1993) and will thus not be elaborated on fur-
ther here. Similarly, the notion that regularity might be fundamental to
music appreciation is slowly emerging (e.g., see Drake, 1998), although
its specificity to music is rarely addressed. It is worth mentioning that the
universality of musical scales and of regular pulse faces difficulties in
finding a niche in ethnomusicologist circles (which are more concerned
about understanding each musical system in its context). Yet the plausi-
bility of considering these two principles as music universals has increased
in recent years (e.g., see Arom, 2000).
While musicologists generally remain reluctant to envision biological
determinism in music and, consequently, do not engage in the active
search of relevant evidence in the various types of music around the
world, developmental psychologists have been more courageous. Infants
have been shown to possess precocious sensitivity to musical scales and
for temporal synchronicity. For example, six- to nine-month-old infants
process consonant intervals better than dissonant intervals (e.g., see
Schellenberg and Trehub, 1996) and exhibit learning preferences for mu-
sical scales (Trehub et al., 1999). In most musical cultures, musical scales
make use of unequal pitch steps. Infants already show a sensitivity bias
toward musical scales, since they have been shown to be better at de-
tecting a small pitch change in an unequal-step scale than in an equal-
step scale. On the time dimension, infants prefer music that is subject to
an isochronous temporal pulse. For instance, like adults, four-month-old
infants are biased toward perceiving regularity; they exhibit sensitivity to
slight disruptions of temporal isochrony (see Drake, 1998, for a review).
All of these aspects of auditory pattern processing suggest the presence
of innate learning preferences.
 The Biological Foundations of Music 441

Surprisingly, preference biases are rarely explored in adults. Adults are

studied as information-processing machines, not having emotional biases.
Part of this situation may be attributed to the widely held belief that
preferences, or emotional interpretations of music, are highly personal
and variable, hence preclude scientific examination. This belief is false.
A recent study of ours (Peretz, Gagnon, and Bouchard, 1998) showed
that emotional appreciation of music appears highly consistent across
individuals, to have immediacy, and to be available to the layperson with-
out conscious reflection and with little effort. Therefore, emotional ap-
preciation of music fits well both with the product of a specialized cortical
arrangement and the purpose of music. Emotional purposes are the first
reasons offered by people to explain why they listen to music (Panksepp,
1995; Sloboda, 1999).
It is probably the study of music as an emotional language that is the
most likely to tap the universal principles that are responsible for its
ubiquity. Until the 1960s it was believed that languages could vary arbi-
trarily and without limit. Today, there is a consensus among linguists that
there is a universal grammar underlying diversity. Similarly, it was once
thought that facial expressions of emotion could vary arbitrarily across
cultures, until Ekman (Ekman et al., 1987) showed that a wide variety
of emotions are expressed cross-culturally by the same facial movements.
Likewise, certain aspects of music are culture-specific, although the gen-
eral rules and processes subserving musical encoding of pitch and timing
can be universal. These universal principles may in turn be subserved by
neural networks that are shaped by natural selection.

What Is Music For?

If indeed music corresponds to a musical propensity that emerged early

in human evolution, that is universal and functional early in human devel-
opment, and that resides in a dedicated neural system, then the key ques-
tion becomes “why?” What adaptive function was served by music in
ancestral activities so as to provide its practitioners with a survival advan-
tage in the course of natural selection? There are two main explanations.2
The initial account was provided by Darwin himself (1871) who pro-
posed that music serves to attract sexual partners. This view has been
442 I. Peretz

recently revived by Miller (2000) who reminds us that musicmaking is

still a young male trait. However, the dominant view is that the adap-
tive value of music lies at the group level rather than at the individual
level, with music helping to promote group cohesion. Music is present at
all kinds of gatherings—dances, religious rituals, ceremonies—thereby
strengthening interpersonal bonds and identification with one’s group.
The initial step for this bonding effect of music could be the mother-
infant interactive pattern created through singing and motherese (or baby
talk, which refers to the musical way adults talk to infants), thereby fa-
voring emotional communion. These two different adaptive roles attrib-
uted to music do not need to be mutually exclusive. As pointed out by
Kogan (1994), individuals taking the lead in ceremonies by virtue of their
musical and dance prowess can achieve leadership status in the group, a
factor that contributes to reproductive success.
In support of the contention that music has adaptive value, particularly
for the group, is the fact that music possesses two design features which
reflect an intrinsic role in communion (as opposed to communication,
which is the key function of speech). Pitch intervals or frequency ratios
allow harmonious voice blending, and regularity favors motor synchron-
icity or grace. These two musical features are highly effective at promot-
ing simultaneous singing and dancing while admitting some degree of
autonomy between voices and bodies (Brown, 2000). This design appears
specific to music; it is certainly not shared with speech, which requires
individuality for its intelligibility. These special features fit with the im-
portant criterion, discussed by Buss and collaborators (1998), that for a
system to qualify as adaptive it must have a “special design” in order to
offer effective solutions to a problem. The bonding problem in the case
of music is to override selfish genes for the benefit of the group.

The Vogue Caveat

My major reservation toward the biological account of music is that it

subscribes to the biological vogue. Nowadays, all human phenomena,
from humor to rape, are explained in biological terms. Thus, the risk that
future research on music will lose track of the essential role of cultural
diversity, aesthetic freedom, and education is high. Fortunately, there is
 The Biological Foundations of Music 443

one notorious discrepant voice that reminds us to advance cautiously on

the biological path. This discrepant voice is that of Steven Pinker (1997)
who argues that music “is useless. . . . Music could vanish from our spe-
cies and the rest of our lifestyle would be virtually unchanged. Music
appears to be a pure pleasure technology. . . . All this suggests that music
is quite different from language and that it is a technology, not an adapta-
tion” (pp. 528–529). The merit in this position is that it provides us with
the necessary incentive to prove that music is more than just a game for
our mind or for our senses.3
. . . that music is a language which is understood by the immense majority of
mankind, although only a tiny minority of people are capable of expressing it,
and that music is the only language with the contradictory attributes of being
at once intelligible and untranslatable, make the musical creator comparable to
the gods, and music itself the supreme mystery of the science of man, a mystery
that all the various disciplines come up against and which holds the key to their
progress.
—Claude Lévi-Strauss (1969)

Acknowledgments

This chapter is based on research supported by grants from the Medical

Research Council of Canada and the Natural Science and Engineering
Research Council of Canada. I am grateful to Serge Larochelle, Emman-
uel Dupoux, and Carol Krumhansl for their valuable comments on a pre-
vious draft.

Notes

1. As attested, for example, by the recent meeting entitled “Biological Founda-

tions of Music” that was sponsored by the New York Academy of Sciences and
held in New York in May 2000.
2. One notable exception that is worth the detour is David Huron’s recent lec-
ture entitled “An Instinct for Music: Is Music an Evolutionary Adaptation?” The
lecture is available on the Web @ http:/ /dactyl.som.ohio-state.edu/music220/
Bloch.lectures/2.Origi ns.html
3. Certainly, the vacuity of music is helpless to explain how military music suc-
ceeds in leading sexually productive adults to death. The adaptive group-level
explanation does, albeit in its darkest effects.
444 I. Peretz

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This excerpt from

Language, Brain, and Cognitive Development.

Emmanuel Dupoux, editor.
© 2001 The MIT Press.

is provided in screen-viewable form for personal use only by members

of MIT CogNet.

Unauthorized use or dissemination of this information is expressly

forbidden.

If you have any questions about this material, please contact

cognetadmin@cognet.mit.edu.