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Kaplan, Gisela (2009). Animals and Music: Between cultural definitions and sensory evidence. Sign System Studies, 37 (3/4): 75-101.


Abstract and Figures

It was once thought that solely humans were capable of complex cognition but research has produced substantial evidence to the contrary. Art and music, however, are largely seen as unique to humans and the evidence seems to be overwhelming, or is it? Art indicates the creation of something novel, not naturally occurring in the environment. To prove its presence or absence in animals is difficult. Moreover, connections between music and language at a neuroscientific as well as a behavioural level are not fully explored to date. Even more problematic is the notion of an aesthetic sense. Music, so it is said, can be mimetic, whereas birdsong is not commonly thought of as being mimetic but as either imitation or mimicry and, in the latter case, as a 'mindless' act (parrots parroting). This paper will present a number of examples in which animals show signs of responsiveness to music and even engage in musical activity and this will be discussed from an ethological perspective. A growing body of research now reports that auditory memory and auditory mechanisms in animals are not as simplistic as once thought and evidence suggests, in some cases, the presence of musical abilities in animals.
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Sign Systems Studies 37(3/4), 2009
Animals and Music:
Between cultural definitions and
sensory evidence
Gisela Kaplan
Centre for Neuroscience and Animal Behaviour, S&T, Faculty of Arts and Science
University of New England, Armidale, NSW 2351.
Abstract. It was once thought that solely humans were capable of complex
cognition but research has produced substantial evidence to the contrary. Art and
music, however, are largely seen as unique to humans and the evidence seems to be
overwhelming, or is it? Art indicates the creation of something novel, not naturally
occurring in the environment. To prove its presence or absence in animals is difficult.
Moreover, connections between music and language at a neuroscientific as well as a
behavioural level are not fully explored to date. Even more problematic is the notion
of an aesthetic sense. Music, so it is said, can be mimetic, whereas birdsong is not
commonly thought of as being mimetic but as either imitation or mimicry and, in
the latter case, as a ‘mindless’ act (parrots parroting). This paper will present a
number of examples in which animals show signs of responsiveness to music and
even engage in musical activity and this will be discussed from an ethological
perspective. A growing body of research now reports that auditory memory and
auditory mechanisms in animals are not as simplistic as once thought and evidence
suggests, in some cases, the presence of musical abilities in animals.
1. Introduction
Music and other art forms are regularly regarded as a pinnacle of human
achievement and an enduring testament to human culture (Mithen 2006).
The very idea of considering a sense of music to be present in animals
(in the broadest sense of perception and even rudiments of its
Gisela Kaplan
production) seemed out of the question for a long time. The same was
said of complex cognition but it is now well recognized and documented
that animals, from different classes and orders, may possess complex
cognitive abilities including the ability to use tools, solve problems,
recall memories of past events and plan their future (summarized in
Rogers, Kaplan 2004). From an evolutionary point of view it is also
important to ask at least when aspects known to be part of human
musical abilities first appeared in hominids or non-human animals.
Studies of goldfish (Fay 1995) suggest that music discrimination may
have existed even in species of great evolutionary distance from humans.
Of course, discriminating sounds may be vital to survival for any
species in any acoustically rich environment and it seems a major leap
from this to the ability to learn and remember, let alone reproduce and
vocalize, sequences of sounds as well. Yet one notes that studies such as
those on carp (Chase 2001) and Java sparrows (Yamamoto, Watanabe
2008; Watanabe et al. 2005; Watanabe, Sato 1999) have shown that the
discrimination of music, even specific styles and rhythms, is sophisti-
cated rather than rudimentary. Being able to generalize between classical
music and modern western music and apply the principles to other
tunes seems a remarkably advanced ability in fish and songbirds. Like-
wise, the ability of rhesus monkeys to listen to melodies in transposed
octaves and still recognize a melody as the same even when transposed
by one or two octaves, but not by half octaves is comparable to the
musical ability of children to recognize a melody as a whole and not its
parts and to do so in different octaves (Wright, Rivera 2000).
Despite an increasing interest in the musical perception and abilities
of animals, various publications have hastened to add that animals
usually lack a key ingredient to appreciate, recognize, memorize, let
alone reproduce music. It is said to be uniquely human to combine in
music phonatory imitation with metric entrainment (Brown 2007) and
an exclusively human prerogative to have a “natural inclination to
engage with music”, a view attributed to one of Canada’s leading
neuropsychologists of music (see Gess 2007). There may also be an
implicit assumption that only humans practise music on their own
(Kenneally 2008), while others have emphasized the alleged human
uniqueness to perceive and synchronize rhythm because the latter
involves a tight integration and coordination between the auditory and
Animals and Music 77
motor system. Such statements seem highly premature given the
recency of research into this area and the very few species that have been
investigated in any depth.
More or less all these orthodoxies are beginning to be dented already,
however, by new studies that show some, often even compelling,
evidence to the contrary. “Snowball”, the sulphur-crested cockatoo
(Cacatua galerita eleanora) may have been an amusing sideshow on
YouTube but, when researchers investigated the rhythmic movements
that the bird performed to the music to see whether changes in beat, but
not in pitch, would result in the bird’s adaptation to the changed rhythm
(Patel et al. 2008), they found that was largely the case. This bird’s
performance may well meet the definitions of musical rhythmic
behaviour that Bispham (2006) described and analysed in humans. A
paper delivered at the 9th International Conference on Music Perception
and Cognition in Bologna in 2006 (Patel, Iversen 2006) presented
evidence that Asian elephants handling mallets on base drums
maintained a regular and stable drumming tempo over periods of half
an hour and over several days and it has been found that some African
apes use percussions in their natural environment (Fitch 2005). Male
palm cockatoos use a stick, and, together with some woeful screeching
(cockatoos are obviously not songbirds but “sing” all the same), they
drum a steady beat. With such acts Palm cockatoos are said to defend
their territory but also advertise themselves to attract a mate. Here is
thus an example of tool use and music making not in any shape or form
influenced by humans. Whales and dolphins may have complex song
sequences and some of these appear to be sung when an individual is
alone. Many songbirds sing by themselves and practise (and not just in
subsong) and quite a number of them also appear to appreciate species-
foreign sounds and even melodies well enough to integrate them into
their own song (Mathews, Schuyler 2004; Chisholm 1948) (more of this
Songs of animals of a number of orders are largely discoveries only
of the last five or so decades and so is the discovery that the brains of
songbirds possess an entire neuronal network, including a high vocal
centre, dedicated specifically to the task of learning songs, including the
abilities to memorize, produce and even improve and improvise on
songs. Many songbirds retain lifelong plasticity, that is, retain the
Gisela Kaplan
ability to learn new sounds throughout life, and many can mimic speech
of humans as well as sounds of other birds, mammal s and even
inanimate objects (Robinson, Curtis 1996; Kaplan 2000). Obviously,
reproduction of sounds, particularly of those that are not species-
specific, depends on the ability to form a memory of sound and have that
memory transfigured into production of sound.
Neuroscientists use the avian vocal system instead of nonhuman
primates as a model for human vocal learning (Zeigler, Marler 2004;
2007) because primates are not vocal specialists and learners. Indeed,
research on zebra finches (Arnold et al.1976; Konishi, Akutagawa 1985;
Margoliash, Fortune 1992; Vicario, Yohay 1993), canaries (Nottebohm
1977), sparrows (Konishi 1965) and a range of other songbirds (Notte-
bohm 1980) has assumed model status for the study of memory
formation and for the complex interaction between neural activity,
auditory feedback, plasticity, attrition and development of song. Hence,
the neocortex, once thought to be an indispensable precondition for
vocal learning, and as such a mammalian innovation de novo in evo-
lution, is being dramatically replaced by attention to the song control
system of birds as a way of understanding how vocal learning occurs.
2. Animal song: speech or music
As many of these hallmarks of human uniqueness begin to crumble
under the weight of emerging evidence of the abilities of animals, art,
and specifically music, is among the last vestiges of human uniqueness.
However, even in human cultures there appears to be no easy and totally
satisfying explanation for musicality or an aesthetic sense of music.
Many have called music the most unique of human behaviours and the
most intrinsic and defining feature of human culture (McDermott,
Hauser 2004), others have called music a mere useless adjunct, a ‘cheese
cake’ in culture (Pinker 1997) that has little to offer by way of
explanations of evolution and culture whereas language is invariably
considered an essential human trait.
There are writers who claim that human music has converged, quite
coincidentally, to share properties with birdsong and whale song (Mithen
2006), whereas the position of neuroscientists is that birdsong production
Animals and Music 79
helps us understand human language, not music. Debates on human
language origins, as is well known, have been acrimonious at times,
because of a failure to distinguish between language as a communication
system and the computations underlying the system (Hauser 1997). In so
far as human language is premised on acoustic memory and vocal
learning, there are at least a number of classes and orders of animals,
select though they may be, that share this trait with humans. These include
songbirds, cetaceans and bats (Pettigrew 1986). Complex vocal learning
has been shown also in parrots (Ball 1994; Dooling et al. 1995), and in
hummingbird species (Baptista, Schuchmann 1990), that is, in species that
are not closely related taxonomically (Sibley, Ahlquist 1990). This
suggests that the ability to learn vocalisations may have evolved
independently at least three times among birds alone (Gahr 2000).
Non-human primates do not feature greatly in this comparative
exercise of vocal learning among species, with some recent corrections,
and the emphasis on studies of songbird mechanisms has become the
dominant model. For anthropologists and evolutionary biologists,
however, the link of primates to humans remains of strong interest
(Owren, Rendall 2001) and the last few years have seen a strong interest
in auditory and vocal performance of primates (see below).
Music cannot be shown to be adaptive because any empirical
evidence is scant or lacking altogether. The latter is true enough because
music is ephemeral, fleeting, and early oral traditions of song and dance
have apparently not left as many clues as have paintings or architecture.
Arguing that music may have been adaptive (that is, hominids sang
rather than spoke and the best (male) singers had a reproductive
advantage over less competent singers) has been based on no more than
conjecture, as also is the view that music never mattered in the evolution
of Homo sapiens. Neuroscientific studies have begun to research the
underlying mechanisms and neural coding of sounds found in the human
brain during the perception and production of music (for example,
Zatorre, Peretz 2003) but the evolutionary path and any precursors of
language and music (and their relationship to each other in terms of brain
function, see Koelsch 2005 or cultural themes, Merker 2005) in non-
human animals will require far more research, of course, before music can
be deemed a uniquely human trait.
Gisela Kaplan
For most modern humans, music is an essential art. Birdsong is
certainly music to us, but it has been a matter of debate whether the
songs that birds or other animals produce are music to the animals
themselves. There is little argument, however, that birdsong can be very
close to music and the evidence for such parallels is overwhelming
(Rothenberg 2005). The question is whether the birds so praised for
their music by human admirers (that is, by the many composers who
have actively incorporated birdsong into their own compositions)
actually share a sense of pleasure in their own song (Figure 1).
Figure. 1. Birdsong can function as advertising territory but it still sounds
3. Auditory perception
In the 1930s, hearing of birds was examined in the context of musical
sounds and musical ability. These studies, published in scholarly
musicology journals, tested whether birds could distinguish between
pure and noisy tones and whether their “musicality” allowed humans to
classify bird song in music annotation. For instance, are birds capable of
distinguishing intervals of a third, fourth and fifth and can they
Animals and Music 81
memorise a tune and transpose it to another key? This was tested in
budgerigars (Melopsittacus undulates; Knecht 1939), the small nomadic
parakeets of inland Australia, and now one of the most commonly
available pet birds worldwide, and these are not even songbirds. They
also used crossbills, that are songbirds, and it was found that these two
species, despite their differences in song production, were capable of
distinguishing between intervals that were considerably smaller than
full tone steps and they had no difficulty in transposing a song within
four octaves (Knecht 1939). The same was found to be true of pigeons,
Columba tartus (Wassiljew 1933). Memory of auditory cues was
ascertained for a difference as small as 1 to 2 Hz showing that this
ability is as well developed in some birds as it is in human hearing. In
another experiment budgerigars were conditioned to recognise one
specific call as a food call. On completion of this training the birds were
meant to be confused by being presented with sounds that embedded the
specific food call in a series of known and unknown sequences of sounds
and songs. The birds were able to identify the food call every time
despite the scramble (Knecht 1939). These findings suggest that
auditory communication in birds may well be extremely subtle and
complex and that the avian ear (not necessarily of all species) may well
be capable of very fine discriminations. Modern studies have confirmed
this in budgerigars (Dooling et al. 1995; Farabaugh et al. 1994, 1998)
and in many songbird species (Marler, Slabbekoorn 2004).
4. Sound distortion
Most research on avian auditory perception is of a relatively recent date.
One of the first tasks arising in this new subfield of neurobiology was to
map the avian brain and to understand its auditory feedback
mechanisms. The avian auditory pathways were mapped out in the
1960s, including the regions of the forebrain involved in processing
auditory inputs. It was found that, despite a lack of peripheral specia-
lisation in the avian ear, higher auditory centres process information
that is biologically relevant to each particular species (Konishi 1974). It
also needed to be explained how the avian hearing organ can deal with
identification of sound location and even with sound distortion
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(Taschenberger et al. 1995), how the sounds irrelevant to the messages
are filtered out and how the auditory pathways function in this process
of deciphering (Dooling 1982; Klinke et al. 1994).
In the 1990s it was discovered that budgerigars are capable of
distinguishing sounds stimulating one ear from sounds stimulating the
other ear (called large free-field binaural unmasking), an ability that had
been documented before only in animals with much larger heads (Dent
et al. 1997), and they show an unusually small signal-to-noise ratio
around 3 kHz (Farabaugh et al. 1998; Okanoya, Dooling 1987). They are
also able to classify a large number of types of contact call s and can
remember these for up to several months (Park, Dooling 1986). This
occurs even when the calls are degraded, as by filter ing or truncating
them (Park, Dooling 1986). The best signal-to-noise ratio attained by
the budgerigar auditory system is in a narrow spectral region of 2 to 4
kHz. The unusually sensitive hearing of budgerigars (called critical
ratio function), compared to that of other birds and mammals, is
characteristic of the species and not a result of domestication or
selective breeding (Farabaugh et al. 1998). Great tits (Parus major) also
show an unusual critical ratio function, which may have been an
adaptation to coping with broadband background noise of leaves and
branches generated by wind in the canopy (Langemann et al. 1998;
Boncoraglio, Saino 2007). Budgerigars do not naturally have many
trees in their arid Australian inland environment yet they have hearing
comparable to that of the great tits. The reason is probably that they also
have to hear against the constant sound of wind that generates
broadband background noise.
Primates, by contrast, have generally not fared well in the new
scrutiny of sound production and reproduction, let alone in music as an
art form, because they cannot acquire new sounds and are said to lack the
capacity for vocal memory of anything novel (Zeigler, Marler 2004). To
an extent, these claims of suggested inabilities of primates are
exaggerated. Seyfarth et al. (1980) showed the learning of appropriate
vocal responses in vervet monkeys. Although the calls were not mo-
dified, as birds modify sounds, the context in which they were made was.
In other primates, it has been found that functionally referential signals
exist even in lower primates, such as tamarins, Saguinus fuscicollis and
Saguinus mystax (Kirchhof, Hammerschmidt 2006), and in alarm calls
Animals and Music 83
of sifaka, Propithecus verreauxi (Fichtel, van Schaik 2006). Recently it
was demonstrated that langur monkeys can remember which group
member had given alarm calls based on auditory cues alone (Wich, de
Vries 2006). These findings may have little to do with music but they
have to do with memory and auditory perception and both are vital
preconditions for musical ability and perception of music. Such recent
research shows that non-human primates may have finer auditory
discrimination and memory than they were hitherto credited to possess
(more below).
5. Memory of song
Song in avian species entails memory — a song that is sung in one
breeding season needs to be remembered in the following breeding
season — and poses complex questions related to how the song nuclei in
the brain manage to lose neurons in the non-breeding season and retain
full memory of the song next season (Nottebohm 1980; Konishi,
Akutagawa 1985), why some aspects of song are discarded (or “over-
learned”), why some song is spontaneous, and why other aspects of song
are imitated or improvised and also why and how some elements of
song are learned at all (Nottebohm et al. 1990; Nixdorf-Bergweiler
1995). Indeed, many factors may determine how long learning takes and
how strong a memory will be formed (Clayton, Soha 1999). Age is
important, and so are shaping events, such as approval or punishment.
Hence song practice in these species includes times of learning but not
reproducing all that has been learned in one season, as well as attrition
of elements and crystallization of song during the subsequent breeding
season (Marler, Peters 1982a; 1982b). The latter occurs either by selec-
tive memory or, as in cowbirds (Molothrus ater), as a result of shaping
in social contexts (Freeberg et al. 2002).
In some species, in which only the male sings, crystallised song may
not be entirely fixed because new syllables, phrases, indeed, a new
repertoire, may be produced in each successive season, as is the case in
lyrebirds (Robinson, Curtis 1996), nightingales (Luscini a mega-
rhynchos) and canaries (Serinus canaria), and these may have been
acquired via a process of improvi sation, rather than by rote learning
Gisela Kaplan
from a tutor, or they may be influenced by females, who may prefer
certain elements over others. For instance, female canaries respond to
higher trill rates in males with higher rates of solicitation displays and
thus they shape the song of adult males (Vallet, Kreutzer 1995).
Relatively little is known of song acquisition, song production or
memory formation and retention in passerines that are vocally
monomorphic. In particular, rather little attention has been paid to the
structure of the song control system in avian species in which both the
males and females sing the same amount of time and behavioural
dimorphism in song is minimal or absent (Kroodsma 1996). The
Australian magpie (Gymn orhina ti b i cen) belongs into this category.
Males and females both sing and there is no evidence to date to suggest
that song plays any role at all during the breeding season (Kaplan 2008).
The magpie is thus an interesting case and one in which musical
ability and “singing for joy” (an aesthetic sense of music?) may be tested.
And here it may be useful to resurrect an anecdote, written in 1903 by
Edgar R. Waite from the Australian Museum in Sydney (Waite 1903).
He supplied a small note for the journal Nature in which he reported his
musical experience with a magpie. He had acquired a nestling magpie
(Gymnorhina tibicen), Bird A, and taught it by playing a flute to sing the
following tune (Fig. 2):
Figure 2. The tune that the magpie learned from a flute play presented by E.
R. Waite (1903).
How the song was taught, how much time it took before the bird
acquired the tune and at what age it first gave a rendition of the tune is
not reported. At any rate, this might not have been the most interesting
aspect of the story. Many birds can mimic (Chisholm 1948) and such
mimicry may include the sounds of animals and inanimate objects, car
horns, telephone and other chimes and, presumably, this extends to
mimicry of any composed piece of music that is within range of its own
vocal abilities. The magpie is an excellent mimic, equally versatile in
producing mimicry as often heard in the male lyrebird’s vocal displays
Animals and Music 85
(Robinson, Curtis 1996; Kaplan 2000; 2003). However, when a second
magpie (Bird B) was added to the aviary, it learned the same tune from
the other magpie resulting in a duet in which both birds shared in a
portion of the tune, each time in the same manner. Bird A started the
tune and completed the first two bars. Bird B, according to the writer,
sang the last two bars. Moreover, once Bird A had commenced the tune,
Bird B adopted a vigilant posture and waited with half-open beak for its
turn and promptly took over to finish the song. Bird B later died and
thereafter Bird A resumed singing the entire song and did so in perfect
pitch, in F major (Waite 1903).
It is puzzling, if one presumes the story to be true, that Bird B was
attending to the commencement of the song and did so in the specific
manner of vigilance. Further, it is noteworthy that bird A resumed
singing the entire song and in the correct sequence (and in pitch of F
major). There is no reason to presume that mimicry involves having to
copy sounds of this complexity in its entirety and in sequence. Since the
song had been divided into two by the birds themselves, the units had
been broken into two and could have been sung as part 2 first and then
part 1, interspersed with the magpie’s own vocalisations but, apparently,
that did not happen. Such anecdotes tend to raise more questions than
they answer. However, the experience described by Edgar Waite may
also suggest that the magpie had a musical sense (if not aesthetic
appreciation) of the tune having a certain internal integrity.
My own research on magpie (Fig. 3) mimicry (Kaplan 2005)
suggests that mimicry is not random. In a hand-raised magpie, over
months of testing and recording, the bird reproduced and practised
specific sounds that had to do with the sound scape of its own
envi ronment. Magpies are territorial and the sounds reproduced were
specific to those sounds that resided permanently within the bird’s own
“territory”. In its case, these consisted of human speech sounds and a
variety of vocalisations of other species (birds and dogs). Its mimicry
practice consisted to 73 per cent of practice of human speech, and then,
in various degrees, of other species that it had heard. Moreover, the focal
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magpie managed to improve its performance of mimicked sounds
substantially over time (Kaplan 20001).
Figure 3. An Australian magpie vocalising and presenting tuneful ‘warbles’,
often performed for hours. Note that a bird performs on its own, usually
away from the group and may sing all year round, male and female alike.
The species has specific territorial calls (carolling) and song plays no role
in the breeding season.
1 The results have also been presented in a paper titled Higher cognition and
communication in apes and birds, with special reference to the vocal repertoire of
Australian magpies (Gymnorhina tibicen) at Symposium, Development and
Evolution o f Higher Cognition in Animals at Australian Academy of Science in
Canberra, ACT, 4 May 2007.
Animals and Music 87
This behaviour perhaps undermines the conclusion that could have been
drawn from Edgar Waite’s description. If another magpie practises
speech with the same earnest application as his magpie practised the
tune, it is perhaps not so much a case of musical appreciation as the fact
that they need to learn and understand whatever is relevant and
important in their territory. The latter hints at a function for survival
and thus follows scientific principles that a behaviour that has been
preserved in a species ought to constitute an advantage for its survival.
By contrast, the conclusion that magpies may have musical appreciation
does not. Having said so, however, music has not been explicable
functionally in humans either. That is why Pinker (1997) could be so
provocative and call it a mere ‘cheese cake’ in human culture.
6. Effect of music
Birdsong in male seasonal singers has a clearly identified function. It
has evolved either to attract a mate on the basis of vocal performance or
to secure and maintain a territory (Catchpole, Slater 2008). The question
here is one beyond these functions, namely, whether music (composed
music) has any effect on animals. We do not know, of course, what
precisely animals perceive when we play music to them but we can
measure behavioural changes. The number of research papers reporting
effects of music (and specific kinds of music) on animals has steadily
increased and, perhaps not surprisingly, these have come from
researchers particularly interested in animal welfare.
In animal welfare it is now believed that music may have a role to
play not so much as a cognitive process (Dowling, Harwood 1986;
Krumhansl 1990) but as enrichment and as leading to beneficial physio-
logical responses. One such study was conducted at the Coulston
Foundation where singly housed chimpanzees were exposed to high beat
and low beat music. Results of the study showed an increase in activity
on presentation of high beat music (Harvey et al. 2000). Another study
of chimpanzees (Howell et al. 2002) in Arizona revealed a therapeutic
effect of music, showing a decrease in agitation and aggression and
promoting relaxation, confirming an effect that has repeatedly also been
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found in humans. As a result, the Arizona Primate Foundation
promoted a stereo music system as environmental enrichment for
captive chimpanzees (Howell et al 2003). Sutoo and Akiyama (2004)
found that music (specifically Mozart) decreased blood pressure in SHR
rats and a similar effect was found in Asian elephants (Wells, Irwin
Similar programs have been run for humans and the value and
efficacy of specific types of music on regulating heart beat in cardiac
patients and reduction of stress in Alzheimers patients have been tested
and examined, apparently with great success, according to an interview
with Arthur Harvey, music professor of the University of Hawai. He is
quoted as saying, on the basis of his own music intervention programs,
that “music can be a tremendous intervention. It can relieve pain and
stress, calm the heart rate and blood pressure, affect physical responses
for healing and growth, and stimulate creative thinking”2. The songs and
rhythms used correspond to near resting heart rate (62 beats a minute)
in the lower frequency. There are now also CDs of lullabies available for
pets, but to my knowledge, there are no scientific studies supporting or
rejecting the claimed soothing effects of specific heartbeat music for
dogs and cats.
However, the results of experiments testing whether chimpanzees
can appreciate music of one type over another have been mixed and often
contradictory. Partly, this might have been so because music is a
summary description for a vast variety of sounds and rhythms but this
was not always considered across experiments. Hence, more recent
studies began to look at types of sound and rhythm. Videan et al. (2007)
selected vocal versus instrumental music and, within these two
categories, classic versus ‘easy-listening’. Results showed that instru-
mental music increased affiliative behaviour in male and female
chimpanzees. Slow tempo ‘easy listening’ music decreased agonistic
behaviour in males more so than fast tempo classical music but had no
effect on females (Videan et al. 2007), suggesting that chimpanzees
respond differently to different types of music. In another study by
McDermont and Hauser (2007) their chimpanzees were found to prefer
2 Altonn, H. 2004. Music, especially by Bac h, helps reduce stress: heartbe at
music calms chimps.
Animals and Music 89
silence over any offering of music and similar differences were found in
studies of the music interests of gorillas (Wells et al. 2006). In one case,
at Melbourne Zoo, a goril la was accidentally found to be interested in
television and he shunned any music offerings, played as enrichment to
the apes in favour of a television program depicting parliamentary
debates in Canberra. He shunned auditory information for the rich
gestural and facial communication in Australian parliamentary politics
(personal communication 2009).
Choosing music on their own was one of the innovative ideas tested
at the Primate Foundation of Arizona. In the study already mentioned
above (Howell et al. 2003) chimpanzees were given their own juke box
with choices from Pavarotti singing to Indian flute playing. At the same
time, they were also given little plastic pianos with four keys and it was
found that they preferred to make their own music and totally ignored
the music they could call up (Fritz 2004). The fact that they never
seemed to tire of producing sounds themselves, in preference to
listening to music, even of their own choosing, may also mean that
chimpanzees like pushing buttons, just as children do or that a toy that
is partially interactive retains a certain fascination.
Hence the evidence about the musical interests and capabilities of
primates is by no means clear and, at times, contradictory. As far as the
experiments explored it, any demonstrated interest need have nothing to
do with aesthetic appreciation but with experience of sounds of the
uterus and with particular frequencies and rhythms for which the
auditory and perceptual apparatus of apes may be equipped, (that is,
these may belong to the psychoacoustical auditory perception of
animals and humans). Moreover, in attempts to assess the effect of
music on animals it would also seem important to revisit the methods
used. In quite a number of designs concerned with animal studies and
music, ‘music’ is defined into broad categories (such as ‘classical’, ‘easy
listening’, or ‘modern’) and one is therefore at a loss to assess to which
of the many aspects of any type of music an animal or a group of animals
may have responded. Finally, if an animal is to be tested for musical
“interest” it is paramount, of course, to establish first its own hearing
range and, more importantly, establish in which frequency range that
species’ own communication naturally occurs. For instance, testing
music perception and music choices in marmosets (a New World
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Monkey) and in rats would be constrained by the fact that their own
communication is largely in a range of frequencies well above those in
which music is composed for humans, even though some important
effects of classical music were recorded in rats (Lemmer 2008).
Marmosets often communicate in a frequency range of 15–24 kHz
(usually no longer audible to the human ear above 18kHZ) while most
human music is located in frequency bands well below that (1-8kHz). It
would also be important to consider structure and variability of their
own vocalisations so that the presentation of human-specific/composed
sounds fall at least within a range of sounds that are recognisable in
specific contexts to the animals being studied. I would not yet judge
primates as lacking an interest in music (McDermott, Hauser 2007)
without a great deal of further testing, simply because even a supposedly
soothing lullaby composed for humans, played, however, at a frequency
range in which, for instance, marmosets may only express fear (such as
low level “egg” calls; cf. Epple 1968) may well be very unattractive to
the primate listeners and thus tell us little about the primate’s ability to
appreciate music.
Neuroaudiological research on a variety of mammals, such as gerbils
(Schultze, Langner 1997), monkeys (Steinschneider et al. 1998; Fishman
et al. 2001) and cats (Schreiner, Urbas 1986; Wallace et al. 1991) has
long since established that the auditory cortex of mammals may contain
representations of amplitude modulation, be capable of periodicity
coding with different coding strategies for pitch and rhythm and, in
monkeys, have different mechanisms subserving pitch perception.
Howeve r, in birds, a vital component is added and that is a feedback
enabling auditory memory formation but also the capability of
reproducing sounds, even if these may not be part of its own species-
specific repertoire. It is the latter ability, shared with humans and some
cetaceans, which would appear to be a vital precondition for complex
music appreciation.
7. Music appreciation by animals?
Despite the substantial agreement by musicians that many songbirds
have a musically pleasing song, it is more difficult to turn the argument
Animals and Music 91
around and ask whether birds themselves have a musical sense or
whether the genetically fixed abilities serve specific functions and are
not at all appreciated as song and music by the birds themselves. The
memorising of song as mimicry, as described above for the magpie or
in the example described by Erwin Tretzel (1998), suggest that part of
some songbird’s endowment is a sense of music or it could not even be
reproduced. For instance, Tretzel described a case of a crested lark
(Galerida cristata) in Bavaria that had imitated the whistled commands
of a shepherd, which it then arranged in C major with “definite metric
construction that revealed a sense for musical form and proportion
(Tretzel 1998). Humans that lack musical ability are not able to re-
produce tunes accurately, let alone arrange them in a composition.
Hence, expressed positively, the ability to do so should be regarded as
evidence of musicality. We would certainly not hesitate to call the
ability to produce a song in pitch musical ability if a human had done so.
Secondly, in some songbirds there arises the question why they
continue to sing once the function of song has been fulfilled (such as
attracting a mate). The point was recently made (Sound Archive Briti sh
Library) that black birds (Tu r dus m er ula) and willow warblers
(Phylloscopus trochilus) develop their song musically long after a mate
has been secured and it was therefore not easy to argue other than to say
that song was developed and sung for its own sake.
Moreover, there are songbirds that continue improvising and some
of them, among them outstanding singers like the Australian magpie,
continue to improve and find new ways of producing song they had
never sung before. Normally, we would call this creativity if it applied
to humans. The magpies I have recorded could sing over four octaves,
use crescendos and decrescendos, use the style of cadenza in accelerated
and retarded form (accelerando and ritardando) and have transitions of
phrases and resolution of sounds that seem to have a musical logic
which cannot be explained merely as accidents or faulty copying of a
species’ own template of species-specific calls. In other words, the
question is whether birds and other vertebrates might have evolved an
aesthetic appreciation of art or, specifically of music (Rogers, Kaplan
2006). There has been qualified support for this view from some
scientists of birdsong from as long ago as the 1950s. William H.
Gisela Kaplan
Thorpe, respected for his extensive work on birdsong, said in print
(Thorpe 1958):
The idea that bird song is often an expression of irrepressible joy can be
supported with some plausible arguments, and is certainly not without
some scientific justification. In so far as this may be true, the song of
birds can be regarded as a first step towards true artistic creation and
Such conclusions voiced by a scientist are warranted because a musical
sense has been relatively difficult to assess. However, there is now some
evidence from neuroscientific studies that singing may indeed be
pleasurable to the singer by increasing dopamine levels in the brain and
perhaps even inducing a state of euphoria (Sasaki et al. 2006; Feduccia,
Duvauchelle 2008).
To test whether animals have an aesthetic appreciation of music
might seem impossible but very basic tests have actually been
conducted on human infants (Zentner, Kagan 1996, 1998; Trainor,
Heinmiller 1998) and these designs can be translated into testing
animals because they involve simple choice tests. Playback experiments
can be instated and, in the case of music, alternative tunes be provided in
different, but freely accessible locations. This is precisely what
McDermott and Hauser (2004) did in a set of experiments with 6 adult
cotton-top tamarins (Saguinus Oedipus). The researchers allowed the
tamarins to choose between a number of sounds, paired for a)
familiar/unfamiliar sounds b) loud noise/soft noise and c) consonant
and dissonant sounds. The fist two sets of experiments (a-b) were
conducted simply to establish whether the tamarins responded to
auditory signals and whether their choices conformed to researcher
expectations. They did. The main set of experiments, however, tested
origins of musical preferences by choosing basic components such as
consonant and dissonant sounds. Consonant sounds are pairs of tones
(counting the fundamental frequencies) and these are related by simple
integer ratios (such as perfect 5th-c and g- interval ratio 3:2; or an octave
- c and c - interval ratio 2:1). Results showed that the tamarins had no
spontaneous preference for either pair of sounds, unlike their human
counterparts, and they concluded that this discriminatory ability may
well be unique to humans. The conclusion appears rather unjustified
Animals and Music 93
however, for reasons of differences in cultural appreciation of music
even within human cultures and sensory perception.
Although it will be impossible here to go into the complexities of
the theories of consonance and dissonance as developed by Pythagoras
and, in modern times, by Helmholtz (1863) it is important to indicate
that there are two ways of examining consonance/dissonance, one in a
musical sense and the other in a sensory, psychoacoustic sense. The
former is culturally determined and nurtured and thus depends on
learning and exposure. The latter is culturally invariant, concerns
isolated cords and is a sensory ability shared by humans and a wide
variety of animals (rats: Fannin, Braud 1971; starlings: Hulse et al. 1995;
Japanese macaques: Izumi 2000). In other words, as Fishman et al. (2001)
found, sensory consonance/dissonance is likely to be shaped by
relatively basic auditory processing mechanisms that are not music
specific. Hence, testing these in isolation may not have told us all that
much, certainly not about musical perception.
In summary, whatever the speculations might be, it seems relativel y
clear that auditory, psychoacoustic perceptual capabilities are more
sophisticated in non-human mammals than once thought and that the
musical abilities of songbirds and vocal learners, including parrots and
parakeets, have been underrated because human culture had predefined
such capabilities as uniquely human. This is circular thinking. A
growing number of studies has begun to show that auditory discri-
mination of animals can be very complex and, that in some species,
notably birds, discriminatory abilities of hearing are matched by output.
Relating this to timing and melody is yet another daunting task awaiting
researchers (Janata, Grafton 2003). Such musical production and
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Животные и музыка:
культурные определения и сенсорные доказательства
Ранее считалось, что только у людей имеется способность к комплекс-
ной перцепции, но к настоящему времени научные исследования
представили достаточно доказательств об обратном. Так, например,
искусство и музыка считаются чем-то исконно человеческим, и
научные доказательства также это подтверждают. Но так ли это?
Искусство означает создание чего-то нового, и его наличие или
отсутствие у животных очень трудно доказать. Более того, взаимо-
отношения между музыкой и языком до сих пор недостаточно
исследованы на неврологическом и этологическом уровнях. Понятие
«эстетической перцепции» еще более проблематично. Утверждается,
что музыка может быть и миметической, но, тем не менее, птичье
пение считается или чистым подражанием или мимикрией, т.е. в
данном случае мы якобы имеем дело с так наз. «неосмысленным»
действием (напр. попугаи, которые автоматически повторяют
Animals and Music 101
сказанное). В данной статье я привожу несколько примеров, где
животные выказывают определенную восприимчивость к музыке и
даже совершают музыкальные действия, и анализирую их поведение
с этологической точки зрения. Все большее число научных
исследовани й свидетельствует, что слуховая память и слуховые
механизмы у животных не столь примитивны, как считалось ранее, и
есть свидетельства, что у некоторых животных даже могут быть
музыкальные способности.
Loomad ja muusika:
kultuurilised määratlused ja sensoorne tõendusmaterjal
Varem arvati, et vaid inimestel on kompleksse taju võime, kuid teaduslikud
uurimused on praeguseks esitanud küllaldaselt tõendeid vastupidisest.
Siiski peetakse kunsti ja muusikat enamasti millekski vaid inimestele
eriomaseks ning tundub, et ka teaduslikud tõendid selle kohta on veenvad.
Kuid on nad seda? Kunst tähendab millegi uudse loomist, mida keskkonnas
niisama ei sünniks, ja selle puudumist või olemasolu loomadel on väga
raske tõestada. Veelgi enam, muusika ja keele soesed ei ole tänaseni ei
neuroteaduslikul ega käitumuslikul tasemel veel piisavalt läbi uuritud.
“Esteetilise tunnetuse” mõiste on aga veel problemaatilisem. Väidetavalt
võib muusika mimeetiline olla, kuid linnulaulu kohta seda tavaliselt ei
arvata ning seda peetakse kas pelgaks jäljendamiseks või mimikriks,
misjuhul on tegemist „mõttevaba aktiga“ (nt automaatselt öeldut järele
korrutavad papagoid). Käesolevas artiklis esitan mitu näidet, kus loomad
näitavad üles teatud vastuvõtlikkust muusika suhtes ning sooritavad isegi
muusikalisi tegevusi, ning analüüsin neid näiteid etoloogilisest
vaatevinklist. Järjest suurem hulk teaduslikke uurimusi tõendab, et
loomade kuulmismälu ja kuulmismehhanismid ei ole nii lihtsakoelised, kui
varem arvati, ning tõendusmaterjal osutab mõnel juhul isegi musikaalsete
võimete olemasolule mõnedel loomadel.
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Full-text available
The detection of signals in noise is important for understanding both the mechanisms of hearing and how the auditory system functions under more natural conditions. In humans, the auditory system gains some improvement if the signal and noise are separated in space (binaural masking release). Birds with small heads are at a disadvantage in separating noise and signal sources relative to large mammals, because interaural time differences are much smaller. Two binaural phenomena in budgerigars related to the detection of tones in noise were examined. Budgerigars show 8 dB of free-field binaural masking release when signal and noise are presented to their right side and correlated noise is presented to their left side. Budgerigars also show a spatial masking release of 9 dB when a signal and noise are separated in azimuth by 90°. These results are similar to those found in humans and other mammals with much larger heads.
The voices of birds have always been a source of fascination. Nature's Music brings together some of the world's experts on birdsong, to review the advances that have taken place in our understanding of how and why birds sing, what their songs and calls mean, and how they have evolved. All contributors have strived to speak, not only to fellow experts, but also to the general reader. The result is a book of readable science, richly illustrated with recordings and pictures of the sounds of birds. Bird song is much more than just one behaviour of a single, particular group of organisms. It is a model for the study of a wide variety of animal behaviour systems, ecological, evolutionary and neurobiological. Bird song sits at the intersection of breeding, social and cognitive behaviour and ecology. As such interest in this book will extend far beyond the purely ornithological - to behavioural ecologists psychologists and neurobiologists of all kinds.
Growing evidence indicates that syntax and semantics are the basic aspects of music. After the onset of a chord, initial music-syntactic processing can be observed at about 150-400 ms and processing of musical semantics at about 300-500 ms. Processing of musical syntax activates inferior frontolateral cortex, ventrolateral premotor cortex and presumably the anterior part of the superior temporal gyrus. These brain structures have been implicated in sequencing of complex auditory information, identification of structural relationships, and serial prediction. Processing of musical semantics appears to activate posterior temporal regions. The processes and brain structures involved in the perception of syntax and semantics in music have considerable overlap with those involved in language perception, underlining intimate links between music and language in the human brain.