Music evolution and neuroscience


There have been many attempts to discuss the evolutionary origins of music. We review theories of music origins and take the perspective that music is originally derived from emotional signals. We show that music has adaptive value through emotional contagion, social cohesion, and improved well-being. We trace the roots of music through the emotional signals of other species suggesting that the emotional aspects of music have a long evolutionary history. We show how music and speech are closely interlinked with the musical aspects of speech conveying emotional information. We describe acoustic structures that communicate emotion in music and present evidence that these emotional features are widespread among humans and also function to induce emotions in animals. Similar acoustic structures are present in the emotional signals of nonhuman animals. We conclude with a discussion of music designed specifically to induce emotional states in animals. © 2015 Elsevier B.V. All rights reserved.
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From Charles T. Snowdon, Elke Zimmermann and Eckart Altenmüller, Music evolution and neuroscience.
In: Eckart Altenmüller, Stanley Finger and François Boller, editors, Progress in Brain Research, Vol. 217,
Amsterdam: Elsevier, 2015, pp. 17-34.
ISBN: 978-0-444-63551-8
© Copyright 2015 Elsevier B.V.
Provided for non
commercial research and educational use only.
Not for reproduction, distribution or commercia
l use.
Music evolution and
Charles T. Snowdon*
, Elke Zimmermann
, Eckart Altenm
*Department of Psychology, University of Wisconsin, Madison, WI, USA
Institute of Zoology, Tiera
¨rztliche Hochschule Hannover, Hannover, Germany
Institute of Music Physiology and Musicians’ Medicine (IMMM), University of Music, Drama and
Media, Hanover, Lower Saxony, Germany
Corresponding author: Tel.: 1.608.262.3974; Fax: +1.608.262.4029,
e-mail address:
There have been many attempts to discuss the evolutionary origins of music. We review the-
ories of music origins and take the perspective that music is originally derived from emotional
signals. We show that music has adaptive value through emotional contagion, social cohesion,
and improved well-being. We trace the roots of music through the emotional signals of other
species suggesting that the emotional aspects of music have a long evolutionary history. We
show how music and speech are closely interlinked with the musical aspects of speech con-
veying emotional information. We describe acoustic structures that communicate emotion in
music and present evidence that these emotional features are widespread among humans and
also function to induce emotions in animals. Similar acoustic structures are present in the emo-
tional signals of nonhuman animals. We conclude with a discussion of music designed spe-
cifically to induce emotional states in animals.
adaptive value, cross-species parallels, emotional signals, emotions in music, evolution of mu-
sic, music and speech interactions
What are the origins of music? Is music unique to humans or does it have an evo-
lutionary history? Does music have an adaptive function and, if so, would this func-
tion have been of use to other species? What is the relationship between music and
This chapter is dedicated to the memory of Michael J. Owren (1955–2014) whose influential work on
emotional signals in human and nonhuman species has provided an empirical and theoretical basis for
our writing.
Progress in Brain Research, Volume 217, ISSN 0079-6123,
©2015 Elsevier B.V. All rights reserved.
Author's personal copy
language? Can music be related to emotional signaling in nonhuman animals? Are
there emotional universals in music and in animal signals? If music can induce emo-
tional states in listeners, can animal signals do the same? This chapter attempts to
provide some answers to these questions. We take the perspective that music was
derived from the emotional signals of other species and had as its initial primary
function to induce emotional states in listeners. We will briefly review various the-
ories of music origins and then provide data suggesting that music is adaptive in pro-
moting social cohesion and has beneficial physiological effects in humans and other
species. We then provide evidence that the emotional content of language is medi-
ated by music-like structures involved both in vowel harmonics and in prosody. Pros-
ody in human speech also influences the behavior of preverbal infants, as well as the
behavior of other species, suggesting an evolutionary continuum. Next, we will con-
sider the possibility of universals in the ways music induces emotions across cultures
and look for similar universals in animal emotional signals. We will provide evi-
dence on some experimental tests of playing music to animals and conclude with
some suggestions for future directions.
There is a variety of ideas about the evolution of music that focus on whether music is
adaptive or not, ranging from the “music as cheesecake” hypothesis of Pinker (1997)
that music is nice but has no adaptive function to the idea that music is sexually se-
lected and is important in mate choice (Charlton, 2014; Darwin, 1871; Kleinman, in
the first volume; Miller, 2000), to the Mixed Origins of Music hypothesis
uller et al., 2013) which maintains that the early roots of music may lie in
an ancient affective signaling system that is common to many socially living mam-
mals. However, later on music also induced aesthetic emotions and facilitated a safe
practice environment for auditory learning, promotion of social cohesion, and for
psychological and physiological well-being.
The origins of music have been hypothesized to be uniquely human following
after the evolution of language, since music requires many of the cognitive skills as-
sociated with language (Patel, 2008) or has evolved simultaneously with language
(the music language hypothesis; Brown, 2000). As an alternative to music being
unique to humans, Juslin and Va
¨ll (2008) and Levitin (2008) have proposed that
music has evolved from emotional communication and that the musical components
of speech provide honest communication about emotions. This is the view that we
will support in this chapter. We agree with Altenm
uller et al. (2013) that there is more
to music than simply affective or emotional communication, but from a phylogenetic
perspective we can focus only on observable behaviors.
In studying the evolution of a phenomenon, there are two separate questions that
need to be answered. The first question has to do with adaptation or function. Can we
discern obvious benefits to music that cannot be found with other types of auditory
inputs such as speech or other sounds? If there is no clear adaptive function that can
be detected then what we study might simply be an artifact of another evolved
18 CHAPTER 2 Music evolution and neuroscience
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function. Thus, music might simply have been an incidental component to the evo-
lution of a complex auditory system that is needed to process speech sounds. The
second question has to do with time course or phylogeny. A trait might be adaptive
solely for modern humans and could have evolved after branching off, or a trait may
have appeared even in nonhuman mammals and may thus be ancestral and shared by
other species as well.
There are two models of phylogeny-divergent and convergent evolution. Most peo-
ple are familiar with divergent evolution: that traits studied in one species might be
shared with a common ancestor. Thus, for humans, apes and monkeys are our closest
relatives and traits shared among several species suggest a common ancestor dating
back to when the lines diverged. Less well known is the concept of converging evo-
lution: that species with similar problems to solve may have developed similar adap-
tations regardless of phylogenetic closeness. Thus, many have argued that songbirds
are good models for human speech and music, since vocal signals appear to play a
much more important role for humans and songbirds than for our closest relatives.
We need to evaluate both adaptation and phylogeny to understand the origins of music.
We first need to demonstrate how music can be adaptive. One of the best known pu-
tative adaptive advantages has been music as a sexually selected trait that allows
males to compete for females. This idea was initially suggested by Darwin (1871)
and subsequently advocated by Miller (2000).Haselton and Miller (2006) found in-
creased attractiveness of men expressing creative intelligence as short-term sexual
partners at the time of ovulation in women. Charlton (2014) has reported that peri-
ovulatory women show significant short-term mating preferences for men who are
attributed as composers of complex music. The “complex” music used by Charlton
(2014) is still relatively simple compared with most composed music, which may
make these short-term mate preferences even stronger with most music.
As articulated by Owren and Rendall (2001) for animal signals, emotional signals
can induce emotional states in others that can lead to social cohesion with shared
emotions and increased cooperation within a group. Mithen (2005) has suggested this
social cohesion function of music for our prehistoric ancestors. Emotional signals
can also influence cognition and have effects on the physiology and neuroendocrine
systems of listeners.
One contemporary study provides evidence for the social cohesion function of
music. Kirschner and Tomasello (2010) studied two groups of 4-year-old children.
In one condition, pairs of children marched around an artificial pond containing
toy frogs, while singing a song to musical accompaniment and picking up the frogs
in time to the song to wake them up. In the other group, pairs of children engaged in
the same actions but without singing. The children were then tested on a task that
involved cooperation with the other child and on a task where one child could choose
to help the other child. In the joint singing condition, children were significantly more
likely to cooperate with and to help one another than in the condition without music.
193 Music is adaptive
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Several cognitive and physiological effects of music have been demonstrated in
human and in nonhuman animals. When neuroanatomical terms are presented in the
form of a song, college students learned the terms more rapidly and retained more of
the terms when tested up to 10 days later (Panksepp and Bernatzky, 2002). Adding
speech to music (as in a song) may lead to greater memory. Weiss et al. (2012) mea-
sured recognition memory for old versus new melodies using piano, banjo, marimba,
and voice, with greater recognition occurring for sung melodies. Emotional mood
induction by music (happy or sad) can influence whether happy or sad memories
can be recalled (Parrott and Sabini, 1990).
Music has also been used in therapeutic situations with reports suggesting music
reduces anxiety and improves mood for medical and surgical patients (Kemper and
Danhauer, 2005), with specific effects on pain reduction and pain distress in the early
postoperative days in patients undergoing abdominal surgery (Vaajoki et al., 2011),
and with soothing music increasing oxytocin levels after open heart surgery (Nilsson,
2009). Music also reduces anxiety and depression, and blood volume pulse ampli-
tude in caregivers of cancer patients (Lai et al., 2011). Although music could not
have evolved initially to alleviate stress in patients, the more general conclusions
are that soothing music can influence physiological process that bring about en-
hanced physical and mental well-being, and these could have had important adaptive
Listening to music has been shown to modulate activity in a network of structures
associated with reward and pleasure in the brain. Using functional magnetic reso-
nance imaging (fMRI) and functional and effective connectivity analyses in human
participants, Menon and Levitin (2005) demonstrated activation of the nucleus
accumbens and ventral tegmental area with subsequent connections to the hypothal-
amus, insula, and orbitofrontal cortex. Salimpoor et al. (2013) used similar methods
with people listening to a piece of music for the first time and found that the aesthetic
rewards of music correlated with the interaction of the nucleus accumbens with the
auditory cortex, amygdala, and ventromedial prefrontal cortex. These results help
explain why listening to music is highly pleasurable.
Studies in nonhuman animals provide similar findings to those in humans, sug-
gesting some sort of continuity across species. Thus, music reduces the distress vo-
calizations produced by newborn chicks in isolation (Panksepp, 1998) similar to the
effects of injecting the social hormones, prolactin, or oxytocin into the brain
(Panksepp, 1996). Music also increases levels of dopamine and norepinephrine in
the brain, both of which are involved in processes of arousal and attention and lead
to rewarding effects (Panksepp and Bernatzky, 2002). Music has been shown to have
several other effects. For example, dogs in shelters were calmer after listening to
classical music and barked more after listening to heavy metal (Wells et al.,
2002). However, music by Mozart (Symphony #40) decreased heart rate in hyperten-
sive rats, whereas music by Ligetti (String Quartet #2) increased blood pressure in
hypertensive rats (Lemmer, 2008), suggesting that classical music should not be trea-
ted as a unitary genre. Playing of Mozart’s Adagio (from Divertimento #7, K. 205)
reduced blood pressure and stimulated dopamine synthesis in hypertensive rats
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(Akiyama and Sutoo, 2011), but only music in the range of rat vocalizations
(4–16 kHz) was effective, illustrating that the type of music played should be related
to the auditory system of the species being studied (see below). Prenatal exposure to
Mozart’s Piano Sonata (K. 443) led rats when adults to learn maze tasks more
quickly (Chikahisa et al., 2006). Ames and Arehart (1972) exposed lambs to music
of Montovani or to white noise and found decreased heart rate and decreased heart
rate variability in music-exposed lambs.
One would not normally expect fish to be responsive to music, but several studies
in fish have reported effects on growth rate and physiology. Gilthead seabream
showed increased growth rate and weight gain, but decreased dopamine levels when
exposed to Mozart’s Eine Kleine Nachtmusik (K. 525) (Papoutsoglou et al., 2008),
similar to results in common carp (Papoutsoglou et al., 2007).
In summary, taken together these human and animal studies suggest a role for
music in emotional induction and coordination of behavior, increased cognitive
skills, in beneficial physiological effects, and positive neurochemical changes. How-
ever, it is not clear what aspects of music have positive physiological and cognitive
effects in humans and animals. In many cases, the precise music being used is not
specified, and in other cases music by Mozart is used ostensibly to mimic the
now discredited “Mozart effect” on human cognition (Steele et al., 1999). It is likely
that different aspects of music—tempo, harmony versus dissonance, major versus
minor keys, note duration, and familiarity—may all have an influence on these pro-
cesses. Future work should examine with greater precision which aspects of music
have specific effects on both humans and animals. With nonhuman animals, re-
searchers should consider the range of auditory sensitivity in the tested species, as
well as typical tempos in animal vocalizations when testing with music, since the
literature also reports many studies where music has no effect on animal develop-
ment, physiology, or behavior.
The second evolutionary issue concerns whether music or music-like phenomena are
seen in other species. If we do see aspects of music in other species, then the origins
of music may predate our own species. There already have been several reviews on
this by Altenm
uller et al. (2013),Fitch (2006),Hauser and McDermott (2003), and
Patel (2008, 2010), each reaching different conclusions. Hauser and McDermott
(2003) assert that any features of music perception found in nonhuman animals must
be related to similar perceptual systems and not to music, since they assume that mu-
sic is not to be found in animals. Fitch (2006) is more open-minded and considers that
learned song in birds, whales, and other species might represent convergence to mu-
sic in humans whereas drumming by apes might represent a potential homology.
Patel (2010) argues in partial agreement with Hauser and McDermott that any as-
pects of music cognition that are based on brain functions were developed for other
purposes and cannot be part of the natural selective processes for music. However,
214 Music and phylogeny
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like Fitch he thinks that species with vocal learning might be able to display with
humans the ability to synchronize behavior to the changing of tempi in music,
and that this may represent a phylogenetic origin of the ability to keep time with
a beat. Indeed, Patel et al. (2009) have shown that a cockatoo is able to synchronize
to a changing beat. Here, we adapt the view of Altenm
uller et al. (2013) of two emo-
tional systems, with “strong” emotions having close parallels with emotional com-
munication in other species and “aesthetic” emotions being derived in humans.
We will detail support for this view in later sections but first address some other
data from animals. Patel and others have argued that vocal learning is a prerequisite
for beat synchronization and a study of rhesus macaques (which do not learn vocal-
izations) found that the macaques could detect rhythmic groupings but not the beat
(Honing et al., 2012). However, a recent study on one sea lion (Cook et al., 2013)
demonstrated the ability to entrain movement to rhythmic auditory stimuli. Thus, vo-
cal learning may not be a prerequisite for keeping the beat.
Many studies have demonstrated absolute pitch in nonhuman animals (Hulse and
Page, 1988), but only one study (in rhesus macaques) has demonstrated octave gen-
eralization, the ability to recognize melodies when transposed one octave higher or
lower (Wright et al., 2000). Interestingly, the macaques could generalize only when
melodies were taken from the diatonic scale; when they were tested with atonal mel-
odies, octave generalization disappeared. Most studies have used atonal melodies
and the success of generalization with the diatonic scale suggests that the diatonic
scale may have some fundamental perceptual features that can be found even in dis-
tantly related animals. It is interesting to note that research on bird songs (most com-
monly suggested as analogous to human music) has failed to find evidence of
harmonic intervals that match the chromatic, major diatonic, or major pentatonic
scales (Arala-Salas, 2012; Dobson and Lemon, 1977). Thus bird song is not really
musical. This suggests that, even if one used a diatonic scale, one would not find
octave generalizations or relative pitch in songbirds.
Several researchers have examined whether animals can discriminate between
different types of music and whether they show preferences. Porter and Neuringer
(1984) found that pigeons slowly learned to discriminate between music by Bach
and Stravinsky, but the pigeons showed rapid generalization to novel pieces by Bux-
tehude and Scarlatti with Bach, and to pieces by Carter and Piston with Stravinsky.
Human subjects showed similar generalization ability. Watanabe and Nemoto (1998)
found half of Java sparrows tested preferred Bach to Sch
onberg and subsequently
generalized to Vivaldi versus Carter. Watanabe and Sato (1999) reported that five
of seven Java sparrows discriminated between Bach and Sch
onberg and generalized
to novel examples from the same composers, as well as to music by Vivaldi (for
Bach) and Carter (for Sch
onberg). Otsuko et al. (2009) trained rats to discriminate
between music by Bach and by Stravinsky and found that rats could generalize to
novel examples, but they also found that, although rats could discriminate between
composers, they did not exhibit any preferences. Since the auditory range of rat vo-
calizations (unlike pigeons and sparrows) is much higher than that of human music,
and because rats have subsequently been found to react only to the high-frequency
22 CHAPTER 2 Music evolution and neuroscience
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components of music (Akiyama and Sutoo, 2011), it is difficult to interpret the lack
of preference for human music by rats. We address this general issue in greater detail
later in this chapter.
There have been contradictory findings with respect to whether animals have a
preference for consonant over dissonant music. Sugimoto et al. (2010) found that one
infant chimpanzee showed a preference for consonant over dissonant music, whereas
Koda et al. (2013) reported that Campbell’s monkeys did not show any preference for
consonance. However, Chiandetto and Vallortigara (2011) found that chickens did
prefer consonant music. It is hard to make sense of this pattern of results in terms of
In summary, there is considerable controversy about the degree to which music-
like phenomena are found in nonhuman animals, and results from different species
do not suggest consistent phylogenetic homologies or a consistent pattern of conver-
gent analogies. We think that the data are much clearer with respect to emotional
signals. In the next section, we shall consider musico-emotional effects in human
language and emotional communication and then shall seek parallels in animal
Human vowel sounds are based on the chromatic scale. In a series of studies, Purves
and collaborators have shown that the statistical structure of human speech shows a
probability distribution with peaks at frequency ratios that match the chromatic scale.
This appears to be a direct result of the resonances of the human vocal tract, and sug-
gests that music and speech are closely linked. Peaks in the distribution were espe-
cially prominent at the octave, the fifth, the fourth, the major third, and the major
sixth forming the intervals of the pentatonic scale and most of the intervals on a dia-
tonic scale (Schwartz et al., 2003). The authors sampled not only English speakers
but speakers of Tamil, Farsi, and Mandarin and found similar relationships within
each language. Han et al. (2011) examined music and speech from three tonal lan-
guages and three nontonal languages and found that changes in pitch direction oc-
curred more frequently and had larger changes in pitch direction in tonal
languages, and that the music typical of the cultures with tonal language also showed
similar frequent and large changes in pitch direction, suggesting a coevolution of mu-
sic and language. Gill and Purves (2009) showed that the most widely used scales
across time and across cultures are those that are similar to harmonic series. The au-
thors suggest that humans prefer tone combinations that reflect the spectral relation-
ships of human vocalizations. Bowling et al. (2010) sampled speech spectra from
excited versus subdued speech and found that the spectral distribution of excited
speech showed similarities to the distribution of major intervals, whereas the spectral
distribution of subdued speech matched the spectral pattern of minor intervals. This
was particularly noteworthy with respect to major and minor thirds. Thus, the
235 Music and emotion in human speech and parallels in other species
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harmonic structure of speech closely parallels that of music across cultures, and af-
fective changes in emotion are evident in different harmonic structures of speech just
as they are in music.
A second source of music in language is prosody—the intonation contours of
speech. It seems quite likely that we detect emotional signals more clearly through
pitch and intonation contours than we do through actual words. A clear test of this is
in studies of communication between human parents and preverbal infants, where
specific prosodic (musical) features have been identified that can influence the be-
havioral state of the infant (Fernald, 1992). Several short, upwardly rising staccato
calls lead to increased arousal. Long descending intonation contours have a calming
effect, and behavior can be stopped with a single short plosive note. These patterns
were observed across speakers of several different languages. Interestingly, similar
features appear in the calls and whistles used by humans to control the behavior of
working animals (dogs and horses) (McConnell, 1990, 1991). The convergence of
signal structure that humans use to communicate with both preverbal infants and
nonhuman animals suggests that these signals are effective across species. The com-
munication of affect through voice is not unique to humans, and the acoustic struc-
tures involved must have similar effects on the nervous system of both human infant
and animal recipients.
Prosody can be used to induce behavioral changes in others. In the case of humans
who are attempting to manage the behavior of infants and animals, the speakers need
not be directly experiencing the emotion they are trying to induce. Rather, they are
using specific signal types to induce a form of emotional contagion in their listeners.
We know very little about the effects of natural animal signals on inducing emotions
in other animals, a point we will try to address below.
Juslin and Laukka (2003) examined a large number of studies that evaluated how
emotions were conveyed in spoken language and in music performance, and they
found notable similarities between the two modes in the accuracy with which lis-
teners could identify discrete emotions and the specific types of acoustical cues used
to convey discrete emotions in both music and speech. There is indeed a very close
relationship between emotional communication in speech and language.
Given that nonhuman animals respond behaviorally to the same affective sig-
nals that human infants do, we must next ask whether humans have the ability
to distinguish affective states in the calls of other species. Belin et al. (2008) pre-
sented humans with positive and negative affective vocalizations from humans,
cats, and monkeys, and they found that humans were adept at discriminating human
affective calls, but were at chance level with the cat and monkey vocalizations.
However, when the same participants were presented with the same stimuli while
undergoing fMRI of their brains, they found that the animal vocalizations activated
the same areas that human vocalizations with similar valence activated. Specifically
bilateral regions of the auditory cortex were activated more by negative vocaliza-
tions from all three species; bilateral regions of the lateral inferior prefrontal cortex
were activated more by positive vocalizations of all three species; and the right
orbital frontal cortex responded more to negative vocalizations of all species.
24 CHAPTER 2 Music evolution and neuroscience
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Thus, although the human participants did not display conscious recognition of
different animal calls of different emotional valences, their nervous systems distin-
guished between these calls.
In another study of cat calls, Nicastro and Owren (2003) found a modest ability of
humans to discriminate between positive and negative calls, and also found that
participants who owned cats or interacted frequently with cats were more adept
at discrimination. Similarly, Scheumann et al. (2014) tested human ability to dis-
criminate between agonistic and affiliative calls of humans, dogs, chimpanzees,
and tree shrews and found that whereas discrimination of human calls was easy
for everyone, there was a clear effect of familiarity with a species and with the
contexts of agonism or affiliation, leading to more accurate discrimination. Thus,
although human brains appear responsive to affiliation and agonistic calls of other
species, conscious discrimination of these calls appears to require significant famil-
iarity with the species.
In summary, music and speech appear to be closely linked, and the linkage is
clearest at the level of emotional expression. Both the prosody of speech and the
spectral distribution of speech sounds can convey emotional meaning. These same
characteristics are effective in altering the behavior of nonverbal human infants and
of working animals (e.g., horses, herding dogs), suggesting that these emotional sig-
nals are effective across species. The brain areas in humans involved in distinguish-
ing between positive and negative emotions in human and animal calls appear to be
the same and can be activated even if the human is unable to make a conscious dis-
crimination between the affective calls of another species. However, with experi-
ence, humans can make accurate discriminations. Let us now turn to the question
of whether there exist emotional universals in human music, and then consider
whether similar universals are present in animal calls.
Emotions can be expressed in music and there have been several attempts to describe
the structures that convey emotions. Scherer (1995) suggested that sadness is con-
veyed by slow tempos, a narrow frequency range, decreases in pitch, and a slow rate
of articulation. (This is similar to the intonation contours that lead to calming in pre-
verbal infants and nonhuman animals.) Joy is conveyed by fast tempos, increasing
pitches that are highly variable, and by increased rates of articulation. (This is similar
to the intonation contours that lead to increased activity and arousal in preverbal in-
fants and nonhuman animals.) Anger is conveyed by an increase in fundamental fre-
quency and by higher intensity (amplitude), and fear is shown with an increase in
fundamental frequency, many high-frequency components, and a faster rate of
Snowdon and Teie (2013) hypothesized that harmonic structures and pure tones
would be associated with positive states, whereas dissonant (or noisy) structures
would be associated with aggression, fear, and defense. Staccato calls would be
256 Are there emotional universals in human music?
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arousing, whereas legato notes would be calming. Regular rhythms should be asso-
ciated with positive states or events, whereas irregular rhythms would be associated
with negative states or events.
In a review of several studies on how emotions were expressed in both speech
andinmusic,Juslin and Laukka (2003) reported that the structural patterns
matched very closely the predictions made by Scherer (1995).Bresin and
Friberg (2011) experimentally tested the validity of Scherer’s classification by
having 20 trained musical performers manipulate seven different variables (i.e.,
tempo, sound level, articulation, phrasing, register, timbre, and attack speed) to
communicate five emotions (i.e., neutral, happy, sad, fear, and calm). Happiness
was communicated by a fast tempo, staccato articulation, high register, high inten-
sity, and fast attack. Fear was communicated by a fast tempo, staccato articulation,
moderate intensity, low register, and slow attack rate. Sadness was communicated
by a slow tempo, very low intensity, legato articulation, mid-range register, and
slow attack speeds. Calmness was communicated by a slow tempo, low intensity,
legato articulation, high register, and slow attack rate. Thus, when musicians were
asked to express different emotions in the same piece of music, they explicitly used
the same acoustic variables that Scherer hypothesized to be involved in emotional
Gomez and Danuser (2007) studied the relationship between the emotional as-
pects of music and psychophysiological response to music. Participants evaluated
the degree of pleasantness and arousal of different types of music, while simulta-
neous measurements were made of skin conductance, heart rate, and respiration.
There was a close connection between self-reported emotional evaluation and the
physiological responses with mode, harmonic complexity and rhythmic articulation
differentiating between negative and positive valences and tempo, acceleration and
rhythmic articulation discriminating between high and low arousal. Thus, partici-
pants not only evaluated the music appropriately, but the music actually induced
emotional responses.
However, all of these studies have been done using Western listeners and musi-
cians as well as with Western music. Does emotional communication generalize to
music of different cultures, and are listeners who are unfamiliar with music from an-
other culture still able to distinguish emotions? Balkwell and Thompson (1999) pre-
sented Western listeners with no prior experience with Indian ragas with excerpts
from ragas recorded in the field in northern India. Each excerpt was intended to con-
vey one of four emotions (i.e., joy, sadness, anger, and peace), and Western partic-
ipants were able to identify the ragas associated with joy, sadness, and anger,
although peace was confused with sadness. Among the key features for discrimina-
tion were rising notes and a fast tempo for joy, and falling notes and a slow tempo for
sadness—again reflecting the prosodic features used by humans with preverbal chil-
dren and with animals to arouse or calm them, respectively. Despite the great differ-
ences between Indian and Western music, the same structural features appear to
encode the strong emotions of joy, fear, and anger.
In summary, one can find acoustic structures in music that reliably communicate
different emotions. Experienced musicians can manipulate these structures when
26 CHAPTER 2 Music evolution and neuroscience
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asked to communicate a specific emotion, and naive listeners can identify emotions
even within musical genres that are unfamiliar to them. There appear to be some
emotional universals in music.
Based on the results on how music communicates and induces emotions in humans,
we can now ask if similar structures are found in the calls of other species. If we can
find similar acoustic variables influencing emotional calls in nonhuman species, then
it seems likely that the “strong” emotions (see Altenm
uller et al., 2013) could have
served as precursors for human music. The best known model of affective signals in
animals is the motivational-structural model of Morton (1977). Morton evaluated
call structures in fear and aggressive contexts in a variety of bird and mammal spe-
cies and suggested that high-pitched, narrow-band, legato calls were used in fear con-
texts and that low-pitched, broad band (or noisy) calls signaled aggression.
Snowdon and Teie (2013) applied their framework of emotional structures in mu-
sic to the calls of cotton-top tamarins. Recordings of spontaneous calls were pre-
sented to musicians, who evaluated the timbre, tempo, rate of articulation, and
pitch of calls without knowing the context in which the calls were given. Five dif-
ferent clusters of calls were found and subsequently associated with the actual con-
texts in which they were given: calls used for affiliation had harmonic structure,
legato articulation with ascending pitch, and narrow bandwidth; calls used for high
arousal and threat were characterized by broadband staccato calls with clear har-
monic intervals; calls used in fear contexts were characterized by noisy, dissonant,
staccato sounds; and calls signifying confident threats were characterized by legato,
harmonic sounds with rising pitches; the approach context was characterized by calls
in triple meter with moderately long notes displaying harmonic structure but with
both rising and falling intonations. The acoustic properties hypothesized for human
emotional expression and music also appear to have parallels in the vocal repertoire
of tamarins.
Research on several other species provides supporting evidence for some of these
acoustic structures being involved in emotional communication. Yang et al. (2013)
removed estrus females from male mice and found an increase in ascending compo-
nents of ultrasonic vocalizations (indicating arousal) and a return to flat frequency
calls (indicating calm) when reunited with females. In contrast, Brudzynski
(2013) found alarm and threat calls (initiated by release of acetylcholine) in both rats
and cats were characterized by low frequency, constant pitch, and long notes,
whereas positive appetitive vocalizations (initiated by release of dopamine) were
higher in pitch with frequency modulation and short notes (equivalent to the prosodic
features that lead to arousal in preverbal infants and working animals). Soltis (2013)
reported that dominance interactions in African elephants were associated with in-
creased amplitude and duration of calls, whereas social agitation was associated with
increased and more variable fundamental frequency and shorter duration notes.
Aggression and mating were also characterized by high-frequency vocalizations.
277 Are there emotional universals in animal calls?
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In nonhuman primates, Zimmermann (2009) showed that gray mouse lemur calls
increased in pitch and in duration during conditions of high arousal. A startled lemur
produces loud, noisy, and plosive grunts, and females rejecting male mating ap-
proaches also produce short, frequency-modulated calls. Males courting females,
however, use long, frequency-modulated or broadband calls. Infant mouse lemurs
gave short frequency-modulated calls in threats, longer and less modulated calls
when isolated, and low-pitched purrs while being groomed. Lemasson et al.
(2012) studied three species of arboreal Old World primates under two conditions
of affect intensity. As seen with mouse lemurs, each of these species produced higher
pitched calls of longer duration when in the high-intensity condition. Thus, it appears
that increased arousal in these species is communicated with longer calls, rather than
the short, frequency-modulated prosodic variables that humans use to induce arousal.
However, in cats Scheumann et al. (2012) reported intensity was coded by longer
duration calls with shorter intercall intervals but decreased fundamental frequency.
In a comprehensive review of 39 studies across the mammalian order,
Zimmermann et al. (2013) found increases in call rate were associated with
alarm/disturbance and agonistic contexts and to some extent with affiliation as well.
Call duration was longer with both affiliative and agonistic contexts, and increased
fundamental frequency was seen with both alarm and agonistic contexts. However,
there was also much variation between species, with call rate showing the most con-
sistent correlation with arousal.
All socially living animals have to discriminate between individuals as well as
context, and several studies have looked at whether different acoustic parameters
are used for individual recognition or contextual information. In general, the results
suggest that source and filter-related variables (e.g., fundament frequency, peak fre-
quency, bandwidth) code for individual recognition whereas temporal (e.g., call dura-
tion, intercall interval), source-related, and tonal (e.g., voicing, harmonicto noise ratio)
parameters code arousal (for cats, see Scheumann et al., 2012; for baboons, see
Rendall, 2003). In addition, different types of affective calls differ in the likelihood
of coding individual features. Thus, mother baboons could easily discriminate the
contact calls of their own infants, but not their distress screams (Rendall et al.,
2009). There may be adaptive value in structuring a distress scream for an immediate
response without taking additional time and resources to encode individuality.
Emotional contagion is frequently seen when animals vocalize. Singing birds, du-
etting titi monkeys and gibbons, pant-hooting chimpanzees, howling wolves, and
many other species show emotional contagion. When one animal or pair begins to
call, others of the same species join in, until many members of one group or pair
are calling to members of other groups and pairs. The contagious calling serves to
reinforce social relationships within a pair or group, and serves to keep others away
from the pair or group, just as music was hypothesized to promote social cohesion in
our human ancestors.
In summary, there are many parallels between the structures of signals used to
communicate specific emotional states across animal species, just as there are among
humans. This is especially clear with respect to arousal and less clear with respect to
28 CHAPTER 2 Music evolution and neuroscience
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states such as fear, aggression, or affiliation. Emotional contagion is common in
many animal species and serves to promote social cohesion among group members
and to keep others away.
Although there are some similarities in emotional signaling, there are also species
differences that must be considered. This becomes most obvious in the case of play-
ing human music to nonhuman animals. Akiyama and Sutoo (2011), in an effort to
see if playing Mozart would have any effect on blood pressure in hypertensive rats,
found that, if they filtered the music, the components above 4 kHz were as effective
as playing the unfiltered music. Given that rats use frequencies into the human ul-
trasound range for communication and are sensitive to a higher frequency range than
humans, this makes sense. However, many other studies have failed to consider the
ecological relevance of human music to other species.
McDermott and Hauser (2007) tested common marmosets and cotton-top tama-
rins for preferences for Mozart versus heavy metal and found a preference for
Mozart. But when they tested Mozart against silence, they found the monkeys
preferred silence, and they concluded that monkeys are indifferent to music. How-
ever, the monkeys they tested have small bodies and communicate in a frequency
range three octaves above human speech and at a tempo at least twice as fast. It seems
premature to conclude that monkeys are indifferent to music.
Playbacks of animal sounds are often used as the gold standard for evaluating the
functional significance of animal signals, but Owren and Rendall (1997) have pro-
posed an affect-conditioning model of primate affective signals. If affective re-
sponses are conditioned to calls, then it becomes difficult to find naive subjects to
evaluate emotional responses. One solution to this problem is to create species-
relevant music in the frequency range and with the tempos appropriate to the species
being tested, and then to build features into the compositions that are hypothesized to
be of affective significance. Using this strategy, Snowdon and Teie (2010) presented
cotton-top tamarins with music composed in their frequency range and tempos, and
compared their responses with music composed for humans having similar features.
Tamarins responded to “tamarin music” with arousing features with increased activ-
ity and increased signs of anxiety, and sought increased social interactions with
group mates. In contrast, they responded to “tamarin music” with calming features
by reducing activity, increasing foraging, and decreasing social contact. Thus, dif-
ferent emotional states could be induced in monkeys with appropriate species-
specific music. However, music composed to induce similar affective responses
in humans had no effect on the tamarins (similar to results of McDermott and
Hauser, 2007).
Recently, Snowdon et al. (in review) have used music designed to be relevant to
cats (higher pitched than human music with tempos similar to purring or sucking) and
298 How do animals respond to species-relevant music?
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found that cats preferred this music to calming music composed for humans. Further-
more, cat music led to a significant increase in calm behavior in the cats. The use of
species-relevant music may have many practical effects on behavior of animals in
laboratories, zoos, and shelters, but to date most facilities use human music—
generally the genres preferred by the caretakers—and the results evaluating the ef-
fects of music have been inconsistent.
In summary, there is much contradictory literature about the effects of human-
based music on nonhuman species with some authors claiming other species have
no appreciation for music. However, there has been little effort to consider the effects
of specific types of music and even less effort to make music ecologically relevant to
other species. When music is composed that takes into account the ecological differ-
ences between humans and another species, music has been shown to be effective in
inducing emotional responses.
In this chapter, we have argued that music has adaptive functions for humans includ-
ing increasing cooperation and helping, and modulating physiological responses. It
may also have value in mate selection, but in our view, this would be a more recently
evolved effect. Musical structures are found in the distribution of harmonics in
speech and in the prosodic features of speech that communicate emotions. Prosodic
features are also used by humans to manipulate the activities and emotional states of
preverbal infants and working animals. Although humans find it difficult to con-
sciously identify the emotional valence in the calls of other species without direct
exposure and experience with those species, there is some evidence of unconscious
discrimination of affective state in animal calls using brain imaging.
Emotions in music can be differentiated by both musicians and nonmusicians,
and Western listeners unfamiliar with Hindu ragas can nonetheless discriminate
the emotional intent of the composers of the ragas. Many of the acoustic features seen
in how emotions are presented in music are also seen in similar emotional signals in
many mammalian species ranging from rodents to primates. This consistency in the
acoustic structures underlying different affective states supports our notion that mu-
sic has emerged in humans based on strong emotional signals and consequently has
early phylogenetic origins.
However, there is still critical research to be done. Are the differences in affective
signals in some species real or due to different paradigms and different definitions of
behavioral contexts? Can researchers manipulate the affective states of animals
through music that is species relevant? Music is frequently used as psychological
enrichment in shelters, laboratories, and zoos, but rarely does the selection of music
relate to the specific goals of enrichment (does one want more active or calmer an-
imals?), nor are species-relevant aspects considered. More work needs to be done on
the intriguing possibility that human brains might be capable of analyzing animal
sounds at a subconscious level. Finally, the music we enjoy listening to is not just
30 CHAPTER 2 Music evolution and neuroscience
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about strong emotions. Our species has developed a complexly structured corpus of
music that affects us emotionally but also affects us aesthetically (Altenm
uller et al.,
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... Music is a near-universal expression in both individuals (Blacking, 1973;Koelsch, 2012;Tomlinson, 2013;Trehub, 2001) and societies, throughout history and across cultures (Brown & Jordania, 2013;Cross, 2003;Snowdon et al., 2015;Titon & Slobin, 1996). Evolutionary theories concerning the utility of group singing, as the first and most fundamental form of joint music-making (Bannan, 2012;Fitch, 2006;Mithen, 2009), argue that it creates a shared, positive emotional state, facilitates group cohesion, and increases empathic responses (Cross, 2001(Cross, , 2003(Cross, , 2007(Cross, , 2008Greenberg et al., 2015;Harvey, 2018;Loersch & Arbuckle, 2013;Perlovsky, 2010Perlovsky, , 2011Savage et al., 2020;Schulkin & Raglan, 2014;Snowdon et al., 2015). ...
... Music is a near-universal expression in both individuals (Blacking, 1973;Koelsch, 2012;Tomlinson, 2013;Trehub, 2001) and societies, throughout history and across cultures (Brown & Jordania, 2013;Cross, 2003;Snowdon et al., 2015;Titon & Slobin, 1996). Evolutionary theories concerning the utility of group singing, as the first and most fundamental form of joint music-making (Bannan, 2012;Fitch, 2006;Mithen, 2009), argue that it creates a shared, positive emotional state, facilitates group cohesion, and increases empathic responses (Cross, 2001(Cross, , 2003(Cross, , 2007(Cross, , 2008Greenberg et al., 2015;Harvey, 2018;Loersch & Arbuckle, 2013;Perlovsky, 2010Perlovsky, , 2011Savage et al., 2020;Schulkin & Raglan, 2014;Snowdon et al., 2015). Empathy is linked to the development of pro-social behaviours Eisenberg & Miller, 1987;Telle & Pfister, 2015), which in turn reinforce group bonds and cohesion. ...
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Human musicality is a mystery. Theorists have proposed that it was evolutionarily adaptive through its ability to create a shared and positive emotional state, increase a sense of social cohesion, and encourage pro-social behaviours. This research found that group singing provides immediate socio-emotional wellbeing benefits but longer-term benefits are confined to emotional domains. These effects were not unique to group singing, but were similar across comparison groups. Wellbeing was facilitated by both group characteristics (music, movement, socialising) and individual mindset towards participation (motivation, flow), with greater benefits for exercise groups. Implications for social prescribing and similar interventions are discussed.
... At least two other perspectives, however, do. The first is the idea that music evolved from vocalizations that were used to communicate emotions (Snowden et al., 2015), perhaps as part of a pre-linguistic precursor of both music and language (Darwin, 1871;Masataka, 2009). Just as names presumably evolved from vocative addresses to individuals into nouns that could refer to the individuals in their absence, so sounds that communicated a current emotional state (fear, distress, love) came to express (or symbolize, as Langer, 1957, would argue) these affects themselves. ...
... 220). Sloboda and Juslin (2001) described such musical qualities as iconic sources of emotion, and Snowden et al. (2015) showed parallels across species: "the acoustic structures involved must have similar effects on the nervous system of both human infant and animal recipients" (p. 24). ...
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Some accounts of the evolution of music suggest that it emerged from emotionally expressive vocalizations and serves as a necessary counterweight to the cognitive elaboration of language. Thus, emotional expression appears to be intrinsic to the creation and perception of music, and music ought to serve as a model for affect itself. Because music exists as patterns of changes in sound over time, affect should also be seen in patterns of changing feelings. Psychologists have given relatively little attention to these patterns. Results from statistical approaches to the analysis of affect dynamics have so far been modest. Two of the most significant treatments of temporal patterns in affect—sentics and vitality affects have remained outside mainstream emotion research. Analysis of musical structure suggests three phenomena relevant to the temporal form of emotion: affect contours, volitional affects, and affect transitions. I discuss some implications for research on affect and for exploring the evolutionary origins of music and emotions.
... While some see music as nothing other than an evolutionary 'cheesecake' -pleasant but non-essential (Pinker, 1997) -there is growing evidence showing that music may have played a vital role in shaping essential evolutionary functions. The underlying neurological mechanism of music has been linked with essential adaptive functions such as emotional communication, social bonding, mate selection and language (Snowdon, Zimmermann & Altenmüller, 2015). It has also been hypothesized that while the early roots of music may have been an affective signalling system common to many living mammals, it became more specific for humans, inducing aesthetic emotions and facilitating auditory learning, promotion of social cohesion, and psychological and physiological well-being (Altenmüller, Kopiez & Grewe, 2013). ...
Neuroplasticity has been increasingly discussed in phylo-ontogenetic terms the last few years, with a rising number of studies and scientific publications demonstrating its importance in the whole life span learning, development, and well-being domains. This chapter, focusing specifically on the neuroplastic changes happening in the infant brain when provoked from music, attempts to discuss the basic features and principals permeating this connection, bringing to the fore their combined value in terms of enriched development and extended social inclusion. The chapter content offers a steppingstone to both academics and practitioners alike, upon which they can update, ‘rephrase', and specialize their knowledge in the particular interdisciplinary topic, while further reflecting towards the more sensitive and special in education and development practice contexts.
... Listening to music can have positive effects on mood and quality of sleep [5], decrease symptoms of depression [6] and decrease fear [7]. Research has shown that listening to music increases activity in the 'rewarding system' in the brain [8]. ...
Conference Paper
How can smart technology support people with mild-moderate dementia to benefit from the positive effects of listening to music in daily life? The quality of life of people with dementia decreases rapidly when they experience difficulties in using everyday products and lose initiative. With a focus on the interaction with music, we study how smart technology can enable human-product-interaction while adapting to loss of initiative. As a result, knowledge on interaction design will be developed to help designers create better products for people with dementia.
... Consonant patterns lead to calmness and relaxation feelings, whereas dissonance induces anger or fear feelings 22 . It has been hypothesized that the understanding of those patterns is also present in non-human animals 30,31 . Although research in this field in non-human animals is still scarce, a predilection for consonant music has been suggested in some primates and chickens 32,33 . ...
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There is a lack of clarity on whether pigs can emotionally respond to musical stimulation and whether that response is related to music structure. Qualitative Behavioral Assessment (QBA) was used to evaluate effects of 16 distinct musical pieces (in terms of harmonic structure) on emotional responses in nursery pigs (n = 30) during four periods: “habituation”, “treatments”, “breaks” and “final”. Data were evaluated using Principal component analysis (PCA). Two principal components (PC) were considered in the analysis: PC1, characterized as a positive emotions index, included the emotional responses content, playful, sociable, and happy, whereas PC2, characterized as a negative emotions index, included fearful, inquisitive, and uneasy with positive loadings, and relaxed and calm with negative loadings. Musical stimulation (treatment) increased (P < 0.01) both emotional indices, compared to other periods and this response was influenced by harmonic characteristics of the music. We concluded that pigs have a wide variety of emotional responses, with different affective states related to the music structure used, providing evidence of its potential use as environmental enrichment for this species.
... This study shows that, much like humans, animals can psychologically interpret musical pieces by showing interest in biologically and socially important features [65]. It is believed that music has evolved from acoustic structures used by various species for emotional communication [66]. ...
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The welfare of animals, especially those kept in intensive production systems, is a priority for modern agriculture. This stems from the desire to keep animals healthy, to obtain a good-quality final product, and to meet the demands of today’s consumers, who have been increasingly persuaded to buy organic products. As a result, new sound-based methods have been pursued to reduce external stress in livestock. Music therapy has been known for thousands of years, and sounds were believed to improve both body and spirit. Today, they are mostly used to distract patients from their pain, as well as to treat depression and cardiovascular disorders. However, recent studies have suggested that appropriately selected music can confer some health benefits, e.g., by increasing the level and activity of natural killer cells. For use in livestock, the choice of genre, the loudness of the music and the tempo are all important factors. Some music tracks promote relaxation (thus improving yields), while others have the opposite effect. However, there is no doubt that enriching the animals’ environment with music improves their welfare and may also convince consumers to buy products from intensively farmed animals. The present paper explores the effects of music on livestock (cattle, poultry and pigs) on the basis of the available literature.
... The therapeutic use of music in its different forms is the subject of several studies in clinical and non-clinical fields [1][2][3][4][5][6][7][8][9]. However, the therapeutic applications of music require further scientific investigation aimed at validating the results and at standardizing the stimuli. ...
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Music influences many physiological parameters, including some cardiovascular (CV) control indices. The complexity and heterogeneity of musical stimuli, the integrated response within the brain and the limited availability of quantitative methods for non-invasive assessment of the autonomic function are the main reasons for the scarcity of studies about the impact of music on CV control. This study aims to investigate the effects of listening to algorithmic music on the CV regulation of healthy subjects by means of the spectral analysis of heart period, approximated as the time distance between two consecutive R-wave peaks (RR), and systolic arterial pressure (SAP) variability. We studied 10 healthy volunteers (age 39 ± 6 years, 5 females) both while supine (REST) and during passive orthostatism (TILT). Activating and relaxing algorithmic music tracks were used to produce possible contrasting effects. At baseline, the group featured normal indices of CV sympathovagal modulation both at REST and during TILT. Compared to baseline, at REST, listening to both musical stimuli did not affect time and frequency domain markers of both SAP and RR, except for a significant increase in mean RR. A physiological TILT response was maintained while listening to both musical tracks in terms of time and frequency domain markers, compared to baseline, an increase in mean RR was again observed. In healthy subjects featuring a normal CV neural profile at baseline, algorithmic music reduced the heart rate, a potentially favorable effect. The innovative music approach of this study encourages further research, as in the presence of several diseases, such as ischemic heart disease, hypertension, and heart failure, a standardized musical stimulation could play a therapeutic role.
... Consonant patterns lead to calmness and relaxation feelings, whereas dissonance induces anger or fear feelings 22 . It has been hypothesized that the understanding of those patterns is also present in non-human animals 30,31 . Although research in this field in non-human animals is still scarce, a predilection for consonant music has been suggested in some primates and chickens 32,33 . ...
Full-text available
There is a lack of clarity on whether pigs can emotionally respond to musical stimulation and whether that response is related to music types. Qualitative Behavioral Assessment (QBA) was used to evaluate effects of 16 distinct musical pieces (in terms of harmony and vertical density) on emotional responses in nursery pigs (n=30) during four periods: “habituation,” “treatments,” “breaks,” and “final.” Data were evaluated using Principal component analysis (PCA). Two principal components (PC) were considered in the analysis: PC1, characterized as a positive emotions index, included the terms content, playful, sociable, and happy, whereas PC2, characterized as a negative emotions index, included fearful, inquisitive, and uneasy with positive loadings, and relaxed and calm with negative loadings. Musical stimulation (treatment) increased (P < 0.01) both emotional indices, compared to other periods and this response was influenced by the compositional characteristics of the music. We concluded that pigs have a wide variety of emotional responses, with different affective states related to the type of music used, providing evidence of its potential use as environmental enrichment for the species.
... Many of these same structures are seen in the emotional features used in human music: short quick notes are arousing, long harmonic notes are calming, dissonance induces feelings of anger or fear and harmonic patterns lead to feelings of calm and relaxation. These emotional components to music have been hypothesized by musicologists and biologists [7,8] to be present in nonhuman species and serve as the original functional origins of music. Neuropsychological studies of brain activity [9] suggest that these emotional structures of music have different and specific effects on the different brain areas associated with processing different emotions. ...
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Playing music or natural sounds to animals in human care is thought to have beneficial effects. An analysis of published papers on the use of human-based music with animals demonstrates a variety of different results even within the same species. These mixed results suggest the value of tailoring music to the sensory systems of the species involved and in selecting musical structures that are likely to produce the desired effects. I provide a conceptual framework based on the combined knowledge of the natural communication system of a species coupled with musical structures known to differentially influence emotional states, e.g., calming an agitated animal versus stimulating a lethargic animal. This new concept of animal-based music, which is based on understanding animal communication, will lead to more consistent and specific effects of music. Knowledge and appropriate use of animal-based music are important in future research and applications if we are to improve the well-being of animals that are dependent upon human care for their survival.
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Long before the internet provided us with a networked digital system, music exchanges had created a global networked analog system, built of recordings, radio broadcasts, and live performance. The features that allowed some audio formations to go viral, while others failed, fall at the intersection of three domains: access, culture, and cognition. We know how the explosive growth of the hip hop recording industry addressed the access problem, and how hip hop lyrics addressed cultural needs. But why does hip hop make your ass shake? This essay proposes that hip hop artists were creating an innovation in brain-to-brain connectivity. That is to say, there are deep parts of the limbic system that had not previously been connected to linguistic centers in the combination of neural and social pathways that hip hop facilitated. This research is not an argument for using computational neuroscience to analyze hip hop. Rather, it is asking what hip hop artists accomplished as the street version of computational neuroscientists; and, how they strategically deployed Black music traditions to rewire the world’s global rhythmic nervous system for new cognitive, cultural, and political alignments and sensibilities.
In the first comprehensive study of the relationship between music and language from the standpoint of cognitive neuroscience, the author challenges the widespread belief that music and language are processed independently. Since Plato's time, the relationship between music and language has attracted interest and debate from a wide range of thinkers. Recently, scientific research on this topic has been growing rapidly, as scholars from diverse disciplines including linguistics, cognitive science, music cognition, and neuroscience are drawn to the music-language interface as one way to explore the extent to which different mental abilities are processed by separate brain mechanisms. Accordingly, the relevant data and theories have been spread across a range of disciplines. This book provides the first synthesis, arguing that music and language share deep and critical connections, and that comparative research provides a powerful way to study the cognitive and neural mechanisms underlying these uniquely human abilities.