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The Pleasure Principle: Part Two

The World of Animal Music
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Skull and hyoid bone.jpg
Skull and hyoid bone of a Venezuelan red howler (Alouatta seniculus), posterior view. Photo: Didier Descouens.
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The songs and sounds of animals surround us. In the wake of Darwin’s theories of evolution, the status of these sounds as music has increasingly come under fire. Skeptics see a fundamental difference between human and animal expression, relegating the latter to mere functional behavior. The closer that one listens, however, the more one detects how much creativity is involved in these performances — and how little we actually know or understand about our own human music.

Read The Pleasure Principle: Part 3 here. Or return to Part 1.


A Result of Evolution


Song is never just a cultural trait; it also relies directly on physical pre-conditions. And for humans, these pre-conditions took millions of years to develop. A string of spectacular fossil discoveries in recent decades has radically advanced our insight into our physical evolution.

Based on these discoveries, today it is contended that the Hominini vocal range and capacity continually expanded, through Homo ergaster, traced back to eastern and southern Africa between 1.4 and 1.9 million years ago, and through Homo erectus and heidelbergensis, all the way to Homo sapiens. 

That is, it now seems probable that changes in the human voice box allowed for the development of singing and more refined forms of vocalization. Crucial in enabling these capabilities is a tiny, horseshoe-shaped bone in our neck: the hyoid. Since the hyoid has a far more fragile structure than most of our bones, physical remains of it are hard to come by (although a few early fossils have been found). 

Combining these discoveries with contemporary knowledge of our anatomy, scientists can infer its development with some precision. Comparing different Homonin specimens, the hyoid appears to have gradually moved downward over more than a million years. By the time of Homo sapiens, the hyoid had concluded its long quest down the neck, and our voice box had essentially attained its current position and form. 


Illustration from Practical Human Anatomy: a Working-guide for Students of Medicine and a Ready-reference for Surgeons and Physicians. New York: William Wood & Co, 1886.


As soon as the anatomy was there, Iain Morley believes, Homo sapiens would have been equipped with the same vocal abilities that we have today.[6] And we put them to good use. Many believe (although the theory is contested) that it was through enhanced vocal capacity that Homo sapiens eventually displaced the Neanderthals and emerged as the dominant strain of Homonini. It would seem that music offered some kind of advantage as well. But what was it? Answers to this riddle can be grouped into two approaches: adaptationist (Darwinian) and non-adaptationist theories.

Adaptationists argue that human traits are the result of evolutionary processes that enhance a species’ ability to reproduce, whereas non-adaptationists explore the possibility that some traits could be byproducts of other evolutionary processes, genetic drift, or historical constraints, rather than being directly selected for their adaptive value. Although these approaches diverge, both are useful in understanding our relationship to animal sound and song. 

For adaptationists, the development of music is a direct result of evolution and can be an instrument for either of three things: sexual selection, parental care, or social cohesion. Referring to the first, Darwin wrote, “[M]usical tones and rhythm were used by the half-human progenitors of man, during the season of courtship, when animals of all kinds are excited by the strongest passions.”[7] 

It is hard to argue against there being at least some mating component in the development of music. Thanks to the cruel-but-groundbreaking research of Argentinian-born neuroscientist Fernando Nottebohm, we know that the areas of the brain responsible for bird song, which he calls “song nuclei”, will grow in size during mating season and return to their normal state after its conclusion.[8] 

But animals are known to sing not only in the courtship season but during other times of the year as well. And one would have to bend Darwin’s theory considerably for it to cover all of the different forms that music has taken. This is why, for some in the adaptationist camp, the concept of parental care has become a comparatively more attractive topic of study. 


Photo of Sergio Franchi performing on the Ed Sullivan Show.
Photo of Sergio Franchi performing "La La La (If I Had You)" on the Ed Sullivan Show, May 1970, copyright CBS Television.


As Homo sapiens started ruling the Earth, brain mass quickly increased in both adults and infants. As a result, a baby had to be born earlier in its development to still fit through the birth canal. These younger infants would grow to be more intelligent than their predecessors, but they also required more protection and attention in their early years. 

As their parents were unable to continuously hold them, they had to be sat down from time to time. To calm their babies, it is thought that parents would sing to them in an early form of song, today commonly referred to as “motherese”.[9] It is a theory that is plausible, especially as singing has remained, to the present day, an instinctive technique for soothing babies. 

Social cohesion is the most complex and arguably the most appealing adaptationist theory. The notion follows that music would have been used by ancestral cultures to create cohesion, or even adhesion, between members of a group, some of whom might otherwise have violently clashed or revolted. Music provided what might be described as a “master clock”, synchronizing breaths and motions, making physically demanding work more bearable, and allowing for conflicts to be resolved more peacefully.

Sound and song may also have been used to make individual members of a group realize that they were stronger as part of a collective. Björn Merker, from the Royal University College of Music in Stockholm, provides an analog from the world of animals, singling out chimpanzees, who use synchronized calls for their “carnival display”. 

If a chimpanzee discovers a valuable food resource, it will send out a call to other members of the group. As soon as a fellow chimpanzee arrives at the location, it will join in, thereby amplifying the signal and attracting even more chimps — including females from outside territories. 

Thus, Merker contends, synchronous vocal signaling serves three purposes: the sharing of resources among group members, which increases their chances of survival; the attraction of female chimpanzees to the territory, which increases their chances of progeny; as well as a sort of control on valuable calls and non-valuable ones (new chimpanzees arriving on the scene may retaliate in the event of a false alarm).[10] 



Just how closely social cohesion and musical expression are related in humans can be observed in individuals affected by Williams syndrome. Those displaying signs of the syndrome often exhibit significantly raised levels of oxytocin, a neuronal-signaling molecule and peptide hormone, making them far more sociable and providing them with a “cocktail party” personality. This increased sociability is closely linked to a significantly higher affinity for and interaction with music.[11] 

Within the adaptationist world, it is not hard to apply these criteria to both animals and humans. It is only when the focus shifts away from our shared roots and toward cultural and neurological factors that a divide starts to open up. 

Steven Pinker, the most prominent proponent of the non-adaptationist camp, disagrees with the notion that music constitutes an evolutionary mechanism. Music, Pinker believes, offers no Darwinian benefits to our survival — humans could have evolved without it. He describes music as an “auditory cheesecake”; it is pleasurable and fulfills human desire, but is biologically useless.

Pinker bases an important part of his argument on the idea that music is replete with harmonic sounds, sounds comprised of frequency components that are integer multiples of a fundamental frequency.[12] What he is essentially referring to is pitched sounds and how they combine into harmonies or dissonances to excite our brains. 

Within a natural environment, after all, a sound pitched at the level of human hearing is a rare phenomenon. So by singing a perfect B or C, or by creating melodies out of different tones, we are making a powerful statement — almost like drawing a perfect circle by hand. Initially, we may respond to these tones with fear or rejection, but ultimately our brain rewards this display of perfection with sensations of arousal and pleasure. This is the reason why we keep coming back to it.

Non-adaptationist theories also depend to a large degree on the assumption that music takes elements of language and sublimates them, leading to a place beyond words where the basic, visceral elements of speech — prosody, intonation, or even tiny details, such as the trembling of one’s voice — take over. Adherents of this perspective focus on the ways in which music activates different parts of the neurological pathways and creates experiences that can challenge and calm, excite, and pacify all at once. 

From what we know, imitation of non-linguistic sounds seems to have been a key element in the earliest forms of music. In some Mongolian tribes, for example, the imitation of wild animals is still a frequently performed ritual, and both the Tuvans and early Chinese said that it was animals who brought music to people.[13] 

Human music likely started out as a game of imitating nature and a celebration of life, as an affirmation of our ties with our surroundings. Composer Emily Doolittle sheds light on the complexity of these relationships, stating that the “connection between animals and music is not limited to one part of the world, one time, or one type of society… [T]he ways in which animal songs are used in music depends not on the region of the world… but rather on the type of relationship with music that they choose to cultivate.”[14] 

And yet, although we share many para-musical forms of behavior with other animals, there is decidedly a difference in intent. Many gelada monkeys (Theropithecus gelada), for example, are capable of using musical techniques, such as glissando and staccato rhythms, repetitions and variations with different accentuations, and creating “melodies that have evenly spaced musical intervals covering a range of two or three octaves”.[15]

Outwardly their sounds may be all but indistinguishable from the imitations of an early human singer. But the latter’s act of imitation elevates the song onto a different plane. If the original melody were a tune sung by a bird during mating season, creative transformation turns it into a song about a bird singing during mating season. The difference may seem subtle, but it goes a long way toward explaining why human music would add increasingly more layers of reflection and abstraction to the equation, heading in an entirely different direction.


A boy playing and singing “Sipping Cider Through a Straw” at a pie supper in Muskogee County, Oklahoma, USA, 1940.
A boy playing and singing “Sipping Cider Through a Straw” at a pie supper in Muskogee County, Oklahoma, USA, 1940. Washington, DC: LOC, Farm Security Administration – Office of War Information Photograph Collection. Photo: Russell Lee.

The World Through Our Eyes


These arguments are convincing, but they are certainly not the entire story. No one will dispute the interrelatedness of different areas of the brain that deal with music and speech. But language most likely followed music and not the other way around.[16] What’s more, language and music are not as closely connected as some researchers make them out to be: It is often observed that even stroke patients with severely impaired speech abilities will still be able to sing demanding parts in a choir.[17] 

Although they are also occupied with sound, our processes of speech perception treat incoming data differently than our music-processing system. While speech perception involves the processing of articulatory movements (of a speech act), musical perception zooms in far more precisely on the melodic contours of a sound. 

Musical and linguistic perception are not so much extremes on a continuum but rather entirely different modes of observing similar phenomena. In fact, one could argue that the abstraction that language provides, its building of meaning from atomized units of sound and content, paved the way for a new understanding of music and for the development of themes that could be broken apart and reassembled, repeated and varied, and pitched from the deepest of bass registers to the highest of frequencies.

By abstracting music even further from its imitative and celebratory elements, it could now lead us away from the world and inwards to a sacred space. And within this greater picture, one could certainly argue that it is language which is the anomaly and, perhaps, our truly unique invention — not music.


American ethnomusicologist Frances Densmore.
American ethnomusicologist Frances Densmore sitting next to a horn made of elephant tusk and an early stringed instrument, 1924. Washington, DC: LOC, Prints and Photographs Division. Photo: Harris & Ewing.


Even if we do accept that certain genres of human music, such as classical composition or serialism, are far more complex and conceptually different than animal song, this does not mean that they are by default no longer functional. 

Over the course of centuries, driven by the constant search for new forms of expression and more powerful techniques to elicit even deeper emotions, artists have undeniably added several more layers of abstraction. However, the underlying motivation hardly seems to have changed: to move the listener and to stimulate deep-rooted feelings. It is a common occurrence for music lovers to say that they were moved to tears by Mozart, uplifted by Haydn, almost frightened by a Boulez piano sonata, or taken to a different place by Stockhausen.

Far less frequently will they muse about the move from the tonic to the dominant in bar 232, the exquisite stacking of tones in the powerful opening cluster chord, or the way every note in the piece was chosen according to an ingenious algorithm. Our techniques have become more elaborate, our palettes broader, and the range of possible sensations created by the music greater. Underneath all that, however, music is still as “functional” as it ever was.

Ultimately, of course, as humans, we see the world through our eyes, and we hear the world through our ears. And this naturally makes our music different from that of any other animal. There are plenty of pitched sounds around us — it’s just that animals don’t adapt their songs to our hearing range but to their own. This bias of our perception also means that we don’t know enough to make a qualified statement about most of the sounds and songs we hear in our environment. 

One great example is Jana Winderen’s album The Noisiest Guys on the Planet (Ash International, 2009). It captures the sounds of crustaceans, which combine into a living, planet-spanning installation of rhythmic crackles, snaps, pops, and stridulations. Some crustaceans produce sound by rubbing their pinnae against each other, and they hear by means of “tiny (microscopic) hairs all over their hard shells, and particularly on their antennae... mechanoreceptors [which] respond particularly to vibration or changes in water pressure”.[18]



As is typical for her approach, Winderen has arranged these sounds into a deep, disturbing, and dramatic piece, which takes its audience on an acoustic diving expedition. Anyone with an interest in sound will find plenty to admire here. But expecting us to ever be able to arrive at a meaningful understanding of crustacean creativity — and what their listening experience could be like when their entire apparatus for producing and receiving sound is so utterly different from ours — seems hopeful, to say the least. 

Even with far less exotic sounds, such as those produced by birds, we are only at the very beginning of deciphering meaning or intent. The discovery that birds have a different temporal resolution and are capable of singing and receiving information at a far higher speed than we do is still very recent. 

“The wren is a bird we describe as having a trill, and it can produce sixty-four separate notes in an eight-second burst of song,” field-recording maestro Chris Watson explains. “If you make a recording and slow it down, you can count the individual notes. You can count them, but you can’t understand them. In this blur of notes, there’s information about the bird’s sexual status, its location within the habitat, its breeding status. All of that is conveyed in a fraction of a second, in a speed that we can’t comprehend.” And this, he adds with an emphasis, “is a relatively simple creature”.



There are more examples demonstrating just how limited our perception of the world around us can be. In 1877, a British cotton baron named Joseph Sidebotham reported the first known instance of a singing mouse. Understandably, Sidebotham assumed he had stumbled upon an accident of nature. It would take another 120 years before Timothy Holy, at Harvard, using considerably more advanced equipment, arrived at the conclusion that, in fact, all mice sing. But since most of that occurs in ultrasonic frequencies, we just can’t hear them. 

Speaking to The Atlantic, Holy stated, “I noticed basically right away that these vocalizations were a lot more complicated than I had grown to expect based on the reading I had done of the literature. I remember joking with my postdoc mentor at the time that maybe these were like bird songs.”[19] 

So complicated are these songs — and so similar are the basics of mousean sound production to ours — that they are now being studied in depth to arrive at new insights into human speech disorders. In mouse songs, simple building blocks are continually recombined into complex sequences, all of which bear a different emotional meaning — just as in language and musical composition. 


Photo of a brown rat (Rattus norvegicus), a muroid common to every continent except Antarctica.


It seems almost irrational for non-adaptationists to classify these sounds as purely functional, considering their intricate arrangements and melodic contours, which make them appear very much like human music. On the other hand, the idea that ultimately any combination of sounds that has the outward appearance of music is music is equally problematic. 

In terms of its aesthetic appeal, bat songs, for example, sound nothing like human music. And yet, in many other respects, the way some bats learn their songs and use the sonic vocabulary at their disposal is surprisingly similar to our own.[20] Do we deny these songs the status of music just because they don’t conform to our aesthetic ideals?


Read The Pleasure Principle: Part 3 here. Or return to Part 1.



[6] Morley, Iain, The Prehistory of Music: Human Evolution, Archaeology, and the Origins of Musicality pp. 144–146. Oxford: Oxford University Press, 2013.

[7] Charles Darwin, The Descent of Man, and Selection in Relation to Sex, vol. 2, pp. 336–337. London: John Murray,1871.

[8] Loughnane, Sara, “Stem cell therapy – a bird-brain idea?”, Science Learning Hub – Pokapū Akoranga Pūtaiao, November 15, 2007.

[9] Falk, Dean,“Prelinguistic evolution in early hominins: whence motherese?” Behavioral and Brain Sciences, vol. 27, iss. 4, pp. 491-503, 2004.

[10] Merker, Björn. “Synchronous Chorusing and the Origins of Music”, Musicæ Scientiæ, vol. 3, suppl.1, pp.59–73, September 1, 1999.

[11] Schulkin, Jay & Greta B. Ranglin, “The evolution of music and human social capability”, Frontiers in Neuroscience, vol. 8, 2014.

[12] Patel, Aniruddh D, Emerging Disciplines, Houston: Rice University Press, pp. 91-144, June 15, 2010.

[13] Petrovic, Milena & Ljubinkovic, Nenad, “Imitation of animal sound patterns in Serbian folk music”, Journal of Interdisciplinary Music Studies, vol. 5, iss. 2, pp. 101-118, 2011.

[14] Doolittle, Emily, “Crickets in the Concert Hall: A History of Animals in Western Music,” TRANS – Transcultural Music Review, vol. 12, June 2008.

[15] Richman, Bruce, “Rhythm and melody in gelada vocal exchanges”, Primates, vol. 28, iss. 2, pp.199-223, April 1987.

[16] Montagu, Jeremy,“How Music and Instruments Began: A Brief Overview of the Origin and Entire Development of Music, from Its Earliest Stages”, Frontiers in Sociology, vol. 2, iss. 8, June 20, 2017.

[17] Carnegie, Joel,“These stroke victims can’t speak, but they’re still singing”, PRI’s The World, January 02, 2015.

[18] From the webpage of the Museum of Victoria, no longer available.

[19] Hare, Erin,“Can Singing Mice Reveal the Roots of Human Speech?”, The Atlantic, December 12, 2016.

[20] Shen, Helen H., “Singing in the brain”, PNAS, vol. 114, no. 36, pp. 9490–9493, September 5, 2017.