EMOTIONAL COMMUNICATION

series in affec tive science

Research on the role of emotions in acoustic communication and its evolution has often been neglected, despite its obvious role in our daily life. When we infect others with our laugh, soothe a crying baby with a lullaby, or get goose bumps listening to classical music, we are barely aware of the complex processes upon which this behavior is based. It is not facial expressions or body language that are affecting us, but sound. They are present in music and speech as “emotional prosody” and allow us to communicate not only verbally but also emotionally.

Evolution of

emotional communication From Sounds in Nonhuman Mammals to Speech and Music in Man

Altenmüller Schmidt Zimmermann

This groundbreaking book presents a thorough exploration into how acoustically conveyed emotions are generated and processed in both animals and man. It is the first volume to bridge the gap between research in the acoustic communication of emotions in humans with those in animals, using a comparative approach. With the communication of emotions being an important research topic for a range of scientific fields, this book is valuable for those in the fields of animal behaviour, anthropology, evolutionary biology, human psychology, linguistics, musicology, and neurology.

Evolution of  emotional communication

Why do we think that we can understand animal voices — such as the aggressive barking of a pet dog, and the longing meows of the family cat? Why do we think of deep voices as dominant and high voices as submissive. Are there universal principles governing our own communication system? Can we even see how close animals are related to us by constructing an evolutionary tree based on similarities and dissimilarities in acoustic signaling?

Cover images: chimpanzee © Peter-John Freeman/iStockphoto.com; bat © Valeriy Kirsanov/iStockphoto.com; concert pianist © alenavlad/iStockphoto.com

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e d i t e d by

Eckart Altenmüller Sabine Schmidt Elke Zimmermann

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In E. Altenmüller, S. Schmidt & E. Zimmermann (Eds.) (2013). Evolution of emotional communication. From sounds in nonhuman mammals to speech and music in man (pp. 313-335). Oxford: Oxford University Press.

Chapter 19

A contribution to the evolutionary basis of music: Lessons from the chill response Eckart Altenmüller, Reinhard Kopiez, and Oliver Grewe

Abstract in this article, we discuss the evoiutionary basis of music. We focus on the adaptationa l value of the ch ill response to music linked to strong emotions, feelings of pieasure, and nostalgia. in the first paragraphs, we briefly review the debate on whether music is an evolutionary adaptation or a more recent human invention without any adaptationa l value. A prominent protagonist of the former viewpoint was Charles Darwin, who proposed, in 1871, an analogy of human music to bird-song, linkin g it to courtship and ernerging language abi liti es. Later, the adaptational value of music promoting socia l coherence and we ll -being was emphas ized. in contrast, nonadaptationists argue that music is a more recent invention of humans, comparab le to the control of fire. However, according to this position, music re li es on resources wh ich are necessary for ianguage acquisition and which have developed previously in evolution . Subsequently, we argue that emotions induced by music may also refer to different evolutionary origins . Aesthetic emotions, not necessarily accompanied by an activation of the autonomic nervous system, may have developed relativeiy late in human evo lution, potentialiy in the context of the invention of the first musical instruments some 35,000 years ago. in contrast, strong emotionssuch as chi ll responses to music are linked to an activation of the sympathetic nervous system and the brain reward circuits . Chili responses occur in the presence of novel and unexpected musical events. Furthermore, they depend on individuallistening biographies and personaiity factors . Since chili responses support memory conso lidation, we speculate that they may have reinforced the development of human auditory perception and fine-tuned auditory pattern recog nition. We finally propose the hypothetical "mixed origins of music" theory (MOM theory) : Early roots of music may lie in an ancient affective signa ling system, common to many socia lly living mammals. Later, music was further developed; it induced aesthetic emotions and provided a safe piayground for auditory learning in general. Furthermore it promoted social cohesion and well -being.

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The difficult question about the origins of music There is general agreement that all human cultures possessed and still possess music. Here, we understand music as intentionally created, non-linguistic, acoustical events, structured in time and produced in social contexts (Altenmüller and Kopiez 2005). Amongst the oldest cultural artefacts, musical instruments such as bone and ivory flutes have been discovered in the Hohle Fels cave and the Geissenklösterle cave in the region of Swabia, South-West Germany (Conard and Malina 2008). These flutes, dating back to about 35,000 years, indicate a Paleolithical musical tradition at the time when modern humans colonized Europe. Intriguingly, they are tuned in line with a "modern" diatonic scale: The grip holes ofthe flute are arranged in such a way that an octave is divided into five whole steps and two half steps, with the half steps being separated by at least two whole steps. The tuning is so "modern" that the main theme ofJ.S. Bach's "The art of the Fugue" can be played on a reconstructed Geissenklösterle-flute (Münze! et al. 2002; see Fig. 19.1) . Nicholas Conard, the archaeologist who is incharge ofthe excavation in the Hohle Fels cave therefore speculates that there might have existed cultural traditions, which persisted from the Paleolithic ages until our times and preserved this diatonic scale locally in Central Europe (Conard et al. 2009). This is a strong claim, since performance parameters, i.e., the embouchure and the speed and width of the air-jet used to blow, may yield pitches, which vary more than a quarter tone (Liang 2002). Furthermore, such a presumed tradition is generally difficult to prove due to the Iack of continuity of records used in different sites at different times. It might belong to one of those romanticisms, frequently encountered when dealing with speculations about music and its evolutionary roots. We still do not understand the exact function of these flutes, since it is even unclear whether they were regarded as musical instruments and aesthetic objects or, for example, as signaling instruments, used by hunters or gatherers to indicate a temporary station or to require a specific action. Conard and Malina claim that the emotionallife of Paleolithic individuals was not different from ours (2008). They therefore suggest that these flutes were indeed used for playing expressive tunes and designed to influence early humans' well-being, emotions, group cohesion, and sense of beauty. In favor of this hypothesis is the fact that the manufacturing of these flutes was extremely time consuming and required fine manual skills and technical expertise (Münze! and Conard 2009). Earlier musical activities are likely to have existed although they arenot documented in artefacts or as cave art. Here, we consider instruments made from less durable materials, i.e., reed and wood, and furthermore of joint singing, hand clapping, or drumming as being connected to motor activities such as rhythmic movements and dancing. It is an open question though, to why these musical activities did emerge or persist, despite them being Iabor intensive and therefore costly in an environment of constant struggle for survival. From a scientific viewpoint the question of the origin of music is difficult, if not impossible, to answer. There is too little information available about the nature of musical activities in prehistoric times. Music does not fossilize and we rely on sparse documents, mainly artefacts such as the earlier-mentioned flutes. There are remarkably few cave paintings depicting musicians. Probably the earliest-though still debated-depiction of the use of a musical instrument in rituals is the "Schaman with the Mouth Bow" in the Cave "Le Trois Freres" dating back to about 14,000 years (Anati 1996) (see Fig. 19.2). Further indirect information concerning the origins of music can be obtained either from a comparative approach, for example, when analysing the acoustic communication of nonhuman mammals such as mice, bats, tree shrews, elephants, or primates (see this volume for chapters

Figure 19.1 Replicas of the Geissenklösterle and the Grubgraben flutes dating back to about 35,000

and 20,000 years respectively. The 22-cm \ong Geissenklösterle flute is made from the radius bone of a swan wing . The grip ho\es are arranged in a way that five notes of a perfectly tuned diatonic scale can be played. lt is unclear whether the horizontal carvings are ornaments or were used to determine the position of the grip holes. The 16-cm \ong Grubgraben flute is made from a reindeer tibia . lt is similarly tuned in a diatonic scale, however easier to play. The replicas are manufactured by Wulf Hein, pa\eotechnician. by Ehret (Chapter 4), Fitch and Zuberbühler (Chapter 2), Schmidt (Chapter 6), Snowdon and Teie (Chapter 9), Soltis (Chapter 7), Zimmermann et al. (Chapter 8)) or from cross-cultural studies, especially when comparing music production and appreciation in humans who have been isolated from Westernized cultures, such as the Mafa in the North of Cameroun (Fritz et al. 2009; see also Fritz and Koelsch, this volume, Chapter 18). Finally, conclusions can be drawn from considering ontogenesis, observing the individual developments of vocalizations and responses to music in infants (e.g., Mampe et al. 2009; Zentner and Eerola 2010). Undoubtedly, as many animal vocalizations do in conspecifics, music can evoke strong emotions and change the state of arousal when listened to attentively (Grewe et al. 2007a, 2007b; seealso Panksepp and Bernatzky 2002; for the role of attention, seeKämpfe et al. 2011). These strong emotions can even have effects on physiological functions, for example, on heart beat frequency (Nagelet al. 2008) and brain neurotransmitter production (e.g., Salimpoor et al. 2011) . According to the definition of emotion agreed in this book, such emotions are associated with basic evolutionary founded behaviors "originally evolved for governing fitness-relevant behavioral and physiological responses toward a particular stimulus or situation" (see Altenmüller et al. this volume, Chapter 20). Thus it is not far-fetched to speculate that our Iove for music may be based on evolutionary old mechanisms, linked to the very nature ofhumans. In the following, we will strive to find an answer to whether there is sufficientevidence supporting the claim that music is an evolutionary ingrained characteristic ofhumans. Furthermore we will discuss if making and listening to musicasameans to produce and experience strong emotions is a fitness-relevant behavior or not. Finally, we will propose a tentative model attributing the origins of music to a variety of either biologically-relevant sources or culturally "invented" activities, thus reconciling the opposing adaptationist-non-adaptationist Standpoints of"music

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Figure 19.2 The Schaman with the mouth bow from the cave "Le Trois Freres." Redrawing by E. Altenmül/er.

as part of the evolutionary founded endowment of humans" (e 0g 0 Brown et al 2000) . ". . versus ' 0 mus1c as an mventwn ofhumans, a transformative technology of the mindo" (Patel2010)

ls music an evolutionary adaptation? For theosake ofbrevity we will only summarize the discussion on a potentially adaptational value cultureso There are several recently published artides and books, reviewmg tim on~omg d1scusswn in more detail (eogo, Grewe et al. 2009a; Pate! 2010; Special Issue of M~tstcae Sctent:ae, 2009/2010)° Furthermore, we refer to the excellent dassie The Origins ofMusic edtted by Walhn, Merker, and Brown (2000) 0 Theoadaptationisot's viewpoint posits that our capacity to produce and appreciate music has an :vo~u~wnary adapttve,~al~e; it is the por~du~t of a natural selection process and contributes to the 0s~r vtval of the fit~esto It Imphes that 1t IS bwlogically powerful and based upon innate characterIstt~s of t~e or~amsm, for example, specialized brain networks refined by acculturation and edu~atiOno Histoncally, Charles Darwin, who proposed in his book The DescentofMan, and Selection 111 Re~att~n to Sex a~ an:,logy ~fhuman music to bird-song, has been the mostprominent exponent of t~Is VIewoHe wwte: MusiCaltonesand rhythm were used by half-human progenitors of man, dul!ng a season of courtsh1p, when animals of all kinds are excited by the strongest passions" (01871~2~06, ~ 0 1_209) 0 He further argued that the use of music might have been antecedent to our lmgmstJc abiiities, whLCh evolved from musico This thought has been recently elaborated in the 0_ mus!language model ofSteven Brown (Brown2000)o Indeed, the idea that music or at 1 t cal elements p d 0 eas mus1 0 ro ucmg strong emotwns could be precursors of our language capacity had been

~f mu:Jc 111 ~uma~1

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IS MUSIC AN EVOLUTIONARY ADAPTATION?

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alreadydeveloped in 1772 by Johann Gottfried Herder in his "Treatise on the Origin ofLanguage" (Abhandlung über den Ursprung der Sprache), which received the price of the Royal Academy of Berlino Here, Herder states that language may have evolved from a "natural" affective so und system, common to humans and animals, which aimed at the communication of emotions: "since our sounds of nature are destined to the expression of passions, so it is natural that they will become elements of all emotionso" According to Darwin and Herder, music is an acoustic communication system conveying information on emotions and inducing emotions, thus either (according to Darwin) promoting success in reproduction or (according to Herder) improving social cohesiono These two adaptationist arguments are still discussed: Geoffrey Miller (2000) has explored the sexual selection hypothesis, arguing that making music was a demonstration of hidden qualities in the struggle for mateso Playing a musical instrument means that resources for building such an instrument and investing time in practicing it are availableo Furthermore, the performance itself requires self-confidence, creativity, emotionality and, frequently, bodily features (such as skilled use of fingers) which can be conceived as the display of otherwise hidden qualitieso With respect to the social coherence hypothesis, recent research convincingly demonstrates that making music tagether promotes prosocial behavior in kindergarten children (Kirschner and Tomasello 2010)0 Furthermore, music seems to have the potential to initiate or reinforce the social bonding among individuals in a group by means of "emotional resonance" and shared emotional experienceso McNeill (1995, po viii) assumed that "moving our musdes rhythmically and giving voice consolidate group solidarity by altering human feelingso" In other words, keeping tagether in time creates social cohesiono This phenomenon, which the author called "muscular bonding" (po 2) is further explored by Fritz and Koelsch (this volume, Chapter 18)0 Ian Cross (2009) extended this theory to the effects of music on the human capacity for entrainmento According to Cross, listening to music and making music increases "the likelihood that participants will experience a sense of shared intentionalityo 0000Music allows participants to explore the prospective consequences of their actions and attitudes towards others within a temporal framework that promotes the al ignment of participant's sense of goal. As a generic human faculty music thus provides a medium that is adapted to situations of social uncertainty, a medium by means of which a capacity of flexible social interaction can be explored and reinforced" (Cross, 2009, po 179)0 In going further, he ascribes musicaroJe "as risk-free medium for the exercise and rehearsal of social interaction o" (Cross 2008) Besides sexual selection and group-cohesion, adaptationists frequently propose the roJe of musicaland music-like interactions during parental care as a third major group of evolutionary adaptive behaviorso Motherese, for example, is a specific form of vocal-gestural communication between adults (mostly mothers) and infantsoThis form of emotional communication involves melodic, rhythmic, and movement patterns as weil as communication of intention and meaning and, in this sense, may be considered tobe similar to musicoMotherese has two main functions: to strengthen bonding between motherand infant, and to supportlangnage acquisitiono Lullabies are universal musical activities designed to manipulate the infant's state of arousal, either by soothing overactive children or by arousing passive children (Shenfield et al. 2003) 0All of these functions enhance the infant's chances of survival and may therefore be subject to natural selectiono The importance of making music and listening to nmsic as a potentially adaptational feature of humanity is underlined by neurobiological findings linking our sense of music to hardwired neuronal networks and adaptations of neurotransmitterso Humans possess specialized brain regions for the perception of melodies and pitches oThis is impressively demonstrated by the selective loss of the sense of melody and pitch in congenital and acquired amusia, the former being a genetically transmitted deficit in fine-grained pitch perception, probably due to a dysfunctional

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right frontotemporal neuronal network (Ayotte et al. 2002). Furthermore, humans have specific sensory motor networks to adapt to and entrain with rhythmic stimulation. These networks are an almost unique feature in vertebrates, with only few exceptions such as the dancing Cacadu Snowball (Pate! et al. 2009). Strangemotions whilst listening to music have been shown to affect various neurotransmitters, predominantly the serotonergic and dopaminergic systems. Serotonin is a neurotransmitter commonly associated with feelings of satisfaction from expected outcomes, whereas dopamine is associated with feelings of pleasure based on novelty or newness. In a study of neurochemical responses to pleasant and unpleasant music, serotonin Ievels were significantly higher when participants were exposed to music they found pleasing (Evers and Suhr 2000). In another study with participants exposed to pleasing music, functional and effective connectivity analyses showed that listening to music strongly modulated activity in a network of mesolimbic structures involved in reward processing including the dopaminergic nucleus accumbens and the ventral tegmental area, as weil as the hypothalamus and insula. This network is believed to be involved in regulating autonomic and physiological responses to rewarding and emotiona l stimuli (Menon and Levitin 2005). Blood and Zatorre (2001) determined changes in regional cerebral blood flow (rCBF) with positron emission tomography (PET) technology during intense emotional experiences involving chill responses accompanied by goose bumps or shivers down the spine whilst listening to music. Each participant listened to a piece of their own favorite music to which a chill experience was commonly associated to. Increasing chill intensity correlated with rCBF decrease in the amygdala as weil as in the anterior hippocampal formation . An increase in rCBF correlating with increasing chill intensitywas observed in the ventral striatum, the midbrain, the anterior insula, the anterior cingulate cortex, and the orbitafrontal cortex: again, these latter brain regions are related to reward and positive emotional valence. In a recently published study by the same group, the neurochemical specificity of [(11) C] raclopride PET scanning was used to assess dopamine release on the basis ofthe competition between endogenaus dopamine and [11C]raclopride for binding to dopamine D2 receptors (Salimpoor et al. 2011). They combined dopamine-release measurements with psychophysiological measures of autonomic nervaus system activity during listening to intensely pleasurable music and found endogenaus dopamine release in the striatum at peakemotional arousal during music listening. To examine the time course of dopamine release, the authors used functional magnetic resonance imaging with the same stimuli and listeners, and found a functional dissociation: the caudate was more involved during the anticipation and the nucleus accumbens was more involved during the experience of peak emotional responses to music. These results indicate that intense pleasure in response to music can Iead to dopamine release in the striatal system. Notably, the anticipation of an abstract reward can result in dopamine releaseinan anatomical pathway distinct from that associated with the peak pleasure itself. Such results may weil help to explain why music is of such high value across allhuman societies. Dopaminergic activation furthermore regulates and heightens arousal, motivation, and supports memory formation in the episodic and the procedural memory (Karabanov et al. 2010) and thereby will contribute to memorization of auditory stimuli producing such strong emotional responses.

ls music a human invention? The non-adaptationist theory postulates that music is a human invention and has no direct adaptive biological function. However, it can still be usefu l in terms of manipulating emotions,

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synchronizing group activities, supporting well-being and promoting health. An elegant analogy is the comparison of the ability to make and appreciate music to the ability ofhumans to control fire, which emerged probably some 150,000 years ago (Brown et al. 2009). Clearly, there is no "fire-making" gene and no neurological syndromesuch as a "apyretia"-the inability to make and control fire-but nobody would deny that making fire had an enormaus impactnot only on human well-being (heating, cooking, lighting) and nutrition (better digestion of protein-rich diets from animal meat), but also on physiological parameterssuch as the configuration of our gut and teeth . Why not considering music as such an ingenious invention in humans? Historically, this viewpoint dates back to Herbert Spencer and his essay "On the origin and function of music" (1857). Spencer argued that music developed from the rhythms and expressive prosody of passionate speech. The eminent psychologist William James followed this line of arguments when stating about music that it is a "mere incidental peculiarity of the nervaus system" (James 1890, Vol. 2, p. 419), which has "no zoological utility" (Vol. 2, p 627). Two decades later, German music psychologist C. Stumpf elaborated the non-adaptationist viewpoint. According to his theory, music is the result of correlative thinking, which allowed transgressing from sliding melodic contours to discrete pitches and intervals (1911). The mostprominent modern protagonist of a non-adaptationist position is Steven Pinker, who stated in his book, entitled How the mind works: "Music appears tobe a pure pleasure technology, a cocktail of recreational drugs that we ingest through the ear in order to stimulate a mass of pleasure circuits at once." (1997, p. 528) With respect to the biological significance Pinker comes to the same conclusion as James by stating: "As far as biological cause and effects are concerned, music is useless." (p. 534). An elegant way to conceptualize music as a human invention, while taking into account how human musicality can shape brain functions (Münte et al. 2002) and even influence our genetic information (be it by selection, or by epigenetic features) is the "transformative technology ofthe mind theory" (TTM theory) proposed recently by Aniruddh Pate! (2010). Basically, this theory has developed from a comparative approach, stating that there are aspects of music cognition rooted in other nonmusical brain functions, which are shared with other animals. The logic behind it is as follows: If music relies on other brain functions developed for other purposes, then it is not music which has shaped our genetic material by natural selection. As in fire making, which relies on skilled motor hand functions developed as a consequence of upright gait and adept use of tools, our ancestors invented music by transforming previously acquired abilities (e.g., refined pitch processing, ability to keepintime with an external beat; seealso Patel2008, p. 207f). This " invention of music," once established and tested for usefulness in several domains, does not preclude the later development of more specialized brain regio11s which may be of adaptational value-as is the length ofthe guts in fire-making humans. For example, the potential to memorize melodies and harmonies critically relies 011 superior right temporallobe functions, which are shaped by musical expertise (e.g., Hyde et al. 2009; P. Schneider et al. 2002). The same holds for the sensory-motor hand cortex, which adapts in function and structure to the breath-taking virtuosic skills of the hands in professional violinists and pianists (Bangert and Schlaug 2006; seealso Hyde et al. 2009 for the discussion of the farnaus "hen-egg problem"). Aniruddh Patel's argumentation in favor ofthe TTM theory is based on two generallines which we only briefly delineate here: Firstly, he focuses on tonality processing and on the differential use of scale pitches such that some are perceived as more stable or structurally significant than others. He argues that this "musical" feature leading to implicit formationoftanal hierarchies is not domain specific, but shared with cognitive processi11g of syntactic hierarchies in la11guage. Support for this theory is derived from the neuroscientific research 011 brain networks, which

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s~rve both processin? of musical hierarchies and language syntax (for a review, see Koelsch and SJebel2005). Accordmgly, music tonality processing shares the same resources as syntactic Iangu~g~ processing; and both rely on much more basic cognitive operations, namely the general bu1ldmg of mental hierarchies or cognitive "reference points" (Krumhansl and Cuddy 2010) and the mechanisms of statisticallearning (McMullen and Saffran 2004). The second line of argumentation comes from Patel's work on entrainment to a musical beat. M~si~~l beat perception and synchronization is intrinsic to dance and to many other musical activJties, such as synchronizing work songs or choir singing. It does not appear tobe rooted in language, since at least prose language does not have temporally periodic beats and does not elicit period.ic rhythmic n:ovements from listeners (Pate! 2008, chapter 3). Furthermore, the ability to flexibly synchromze to changing tempi of musical beats seems to be unique to humans and ~o a few parrot species, who share with humans their excellent vocallearning ability. Here, it is ll~portant to note th.at adaptive rhythm and beat perception is essential for langnage acquisition. It IS already present 111 prenatal intrauterine auditory learning (Pate! et al. 2009). Summari zing, Pate! argues that synchronization to a musical beat relies on the brain systems designed for vocal learning, involving specialized auditory-motor networks not restricted to the cortex, but also to midbrain structures such as the periaqueductal gray and its homolog in parrots. Thus, the TTM ability to keep in time with an external beat is a by-product of vocallearning and its neuronal prerequisites. In summary, valid arguments are in favor of music as a human invention, based upon-or transformed fram-already pre-existent cognitive and motor capacities of our brain. However, the TTM theory neglects the strong impact of music on emotions and its possible origin and consequence with respect to its adaptational value. It is interesting to note that the emotional value of music has always been central to the adaptationists' viewpoint, beginning with Herder's and Darwin's ideas quoted earlier. In ~he following two paragrap hs we will demonstrate how music can elicit different types of emotiOns, namely aesthetic emotions and strong emotions. Aesthetic emotions are based on complex feelings with less salient physiological correlates, whereas strong emotions Iead to shivers down the spine and chills or thrills accompanied by physiologica l react ions of the autonomaus nervaus system. We will argue that the former constitute parts of a TTM, invented in Iater times, whereas the latter may point towards an evolutionary old acoustic communication system we share w1th many other nonhuman mammals.

Emotions induced by music Although most listeners agree that music can soundhappy or sad, there is Iess consensus about whether music truly evokes emotions. It is beyond the scope of th is article to review the issue in detail, or furthermo~·e whether and how music induces emotions. This disc ussion has recently been thoroughly rev1ewed by Hunterand Schellenberg (2010). Basically, two main theoretical Standpointsare held: the cognitivist and the emotivist position. In brief, cognitivists argue that happy- and sad-sounding music does not evoke true happiness and sadness in listeners. Rather, affective responses stem from the listener's evaluation of the music (Kivy 1990) . However, such an eva luation or "appraisal " of music can clearly induce emotions, and is in itself a constituent of emotions according to the "component theory of emotions" by Scherer (2004). For example, a boring and i.nac~urate rendering of a musicalmasterpiece might weil induce feelings of anger and ~rustrat10n 111 a music Iover based on his or her knowledge of other more adequate mterpretat10ns.

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In contrast, emotivists posit that music directly evokes and induces emotions. Several mechanisms accounting for such a role of music are discussed. Amongst them, cognitive appraisal is only but one of them. Juslin and Västfjäll (2008) have proposed six other mechanisms, namely (1) brainstem reflexes, (2) conditioning, (3) episodic memory, (4) contagion (5) visual imagery, and (6) expectancies that are fulfilled or denied. With respect to brainstem reflexes, Juslin and Västfjäll consider automatic reactions of individuals towards highly dissonant sounds as such an emotional effect of music, acting via a hardwired neuronal network of the brainstem. Although this phenomenon clearly exists the labeling is debatable, being in cantrast to "true" brainstem reflexes-for example, the constriction of the pupil following exposure to Iight-these reactions to music are interindividually variable, adapting to repetitions and strongly depending on learning. The emotional power of conditioning and episodic memory has been mas terly portrayed by Marcel Proust in the chapter "Swann in Iove," part of the novel In Sem·ch of Lost Time: The hero Swann falls in Iove with a Iady whilst a tune of Vinteuil, is played in the background. Subsequently, the piece becomes the "national anthem of their Iove," strongly linked to positive emotions of tenderness and longing. After breakup of the Iiaison, however, listening to the piece produces intense negative emotions in Swann, feelings of distress, melancholy, and hatred (Proust 1994). Here, associations of music with significant nonmusicallife events cause contrary emotions induced by the same piece of music. It should be mentioned, however, that the associative memories Iinked to music are less frequent than usually assumed : In a retrospective autobiographic study, Schulkind et al. (1999) could demonstrate that only 10% of the greatest hits of the last 60 years were linked to specific episodic memories. Emotional contagion is based on the idea of a sympathetic response to music invoking sad feelings in the presence of sad music (e.g., Levinson 1996) . Music-induced emotionsvia visual imagery can best be exemplified in opera and film-music, linking specific melodies or instruments to emotionally charged scenes or personalities. A suitable example is the mouth organ melody in the movie Once Upon a Time in the West directed by Sergio Leone, linking the chromatic tune to emotions of gloomy suspense and revenge personified by the actor Charles Bronson. Finally, with respect to the expectancies th at are fulfilled or denied, Leonard Meyer (1956) identifies the building up of tension and subsequent relaxation as a major component of emotional appreciation of music. Recently, David Huron has refined this idea in his book Sweet Anticipation (2006). Here, he develops the ITPRA (imagination-tension-prediction-responseappraisal) theory. He identifies five expectancy responses towards music. Two occur before the onset of the event and three afterwards. The first is the " imagination response," which consists of the prediction of what will happen and how will the listener feel when the musical event takes place. The second is the "tension response," which refers to the mentaland physiological preparation immediately before the onset of the event. After the event, the "prediction response" is based on the pleasure or displeasure depending on the degree of accuracy of the prediction. Furthermore, listeners evaluate the pleasa ntness or unpleasantness of the outcome in the "reaction response." Finally, in the "appraisal response," the conscious evaluation of the events occurs. According to Huron, the entire process can Iead to specific affective responses. When expectancies are met, music listeners get a certain degree of pleasure which is reinforced if the event and its evaluation are considered positive. If expectancies remain unfulfilled, this does not necessarily Iead to negative emotions; rather the result may be laughter, awe or chill responses: strong emotions that are frequently accompanied by physiological responses of the autonomaus nervaus system as will be specified later (for a recent update of this theory seealso Huron and Margulis 2010).

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Coming back to the question of the adaptational value of music induced emotions, it is reasonable to distinguish strong emotions, leading to the earlier-mentioned physiological responses fr~~ "~esthetic': emotions (Scherer 2005). Scherer groups emotions into two classes, namely utihtanan emotwns, such as, for example, the prototypical emotions anger, disgust, fear, happiness, sadness, surprise (Ekman and Davidson 1994) and aesthetic emotions. While the former ~an be objectively assessed by psychophysiological measures and have clear adaptational value 111 terms of fitne~s relevant behavior, the latter are characterized by a strong subjective feeling com~onent. Thelt' behavioral and physiological components remain frequently obscure and the emotwnal responses are highly individual. Zentner and colleagues (2008) have analyzed the vocabulary used in ~elf-reports of aesthetic emotions induced by music. They were able to group the common affectlve responses into one of nine categories: wonder, transcendence, tenderness, nostalgia, peacefulness, joyful activation, tension, and sadness. It is difficult to attribute an adaptational value to these highly elaborated feelings, although they clearly are beneficial for human well-being, adding meaning, consolation and security to our lives. Thus, aesthetic emotions are good candidates as a human invention forming parts of a TTM.

The c~ill response in music as an example of emotional peak expenence: Phenomenology and contributing factors "Chills," "thrills," or "shivers down the spine," terms used interchangeably, combined with

goose~bumps occ~r in manycontexts and are elicited in different sensory domains. Physiologically, the.chi~l response IS a consequence ofthe activation ofthe sympathetic nervous system. This acti-

vatwn mduce~ the hai~· to erect, event caused by a contraction of the minuscule arrectores pilorum musdes 111 the sk111. Furthermore, chills are frequently accompanied by other reactions of the sympathetic nervo~s system, for example, increases in heart rate, blood pressure, breathing rate,.and sweat productwn measured by the galvanic skin response. As already mentioned, chills are hnked to dopaminergic activation (Salimpoor et al. 2011), increase in arousal and motivation.thus supp?rting memory formation (for a review, see McGaugh 2006). In this way, events ~eadmg to a chiii response will be memorized more precisely and for a Ionger time. This fact is Importa~t as we will be later considering the adaptational value of chill responses in music. The chiii response seems tobe common in fUITed mammals and occurs as a response to cold or to anger and fear. In the former case, erect hairs trap air close to the warm body surface and create a layer of insulation (Campbell1996). In the latter, erect hairs make the animal appear !arger ~n order to fnghten enemies. This can be observed in the intimidation displays of chimpanzees, 111 .stressed mice. and rats, and in frightened cats, but also in the course of courtship of male ch~m~anzees (Nishida 1997; for a review see Kreibig 2010). A special case of acoustically-invoked chiils 111 mammals seems tobe in responsetomatemal separation calls in some monkey species. Panksepp (1995) argues that feelings of socialloss and social coldness in the offspring could thus be soothed by matemal vocalizations. In his opinion this could explain why, in humans, chills are frequently perceived in the presence of sad or bitter-sweet emotions (Benedek and Kaernbach ~Oll) ..Critically, it should be noted that no systematic study on the frequency, time course, and 111tensity of chill responses to separation calls in nonhuman primates exists . Therefore evidence for such a mechanisms remains, albeit frequently quoted, anecdotal and scientifically ill founded. In h~ma~s, chil~s can be induced through aural, visual, somatosensory, gustatory, and enterocep.tive stimu~atwn (Grewe et al. 2010). Although most research has focused on the previously mentwned music-evoked chills-which are in most instances linked to pleasurable and joyful,

THE CHILL RESPONSE IN MUSICASAN EXAMPLE OF EMOTIONAL PEAK EXPERIENCE 1 323

albeit sometimes nostalgic feelings (Grewe et al. 2007b; Guhn et al. 2007)-it should not be fm·gotten that aversive acoustic stimulation, such as the scraping ~ound of chalk on a blackboard or a dentist's drill, can induce such chill responses even more rehably (Grewe et al.2010). These aversive sounds are characterized by high intensity, high pitch, and frequently high degree of roughness in psychoacoustic terms. In the somatosensory domain chills are evoked by cold, as a thermoregulatory re~ex, and by tactile stimuli. The Iatter are frequently perceived as pleas~rable an.d are probably l111ked t_o grooming and sexual arousal, although research on this. topi.c IS lackmg. Gustatory chiils ar~ evoked by so ur and spicy food, and visual chills by aesthetic objects and feel~ngs of a"':e (Kone~m 2011), but also by viewing highly aversive pictures (Grewe et al. 2010). Fmall!, chiils are ~re­ quently elicited by mere mental self-stimulation, thoughts of pleasure a.nd.emotwn.al m~mone~, including musical ones. Allthese highly diverse chill responses hav.e srmiiar ?hyswlogical COIrelates, as assessed by measurements of skin conductance response, 111creases 111 heart r~tes, ~nd breathing rates (Grewe et al. 2010), and thus cannot be disti2gui.s~e~, by .psychophyswlogrc~l Iabaratory parameters. In the following, we will focus on the positrve chiii res.ponse to musrc Iinked to pleasurable feelings. We will briefly summarize our .find111g~ on musreal parameters and listeners' characteristics contributing to evoking these bodiiy reactwns. . Positive chill responses and emotional peak experiences in music ~r:e rar~ events. Accor.d mg to Goldstein (1980), about 70% of the general po~ulation are famii.Iar with the~e reactiOn S. Interestingly, there are differences between occupatwnal groups. Music students ai.e up to 90 0Vo more susceptible to chills as medical students (80%), and employees of an addictwn r~se~rch center (53%). In a preselected and susceptible group of avid music Iovers and amateur ch01r smgers, only 72% had a chill response when listening to emotionally arousing music for half an ho~r in a Iaboratm·y setting (Grewe et al. 2009a). 1t should be noted that these ~-esponses are fragi.le and not perfectly reproduced when playing the same musical passages on different days, even m individuals with high "chill susceptibility" (see Fig. 19.3) . . . Furthermore, they strongly depend on the context. For example, in an expenn~ent c?mpanng the effects of Iistening to favorite "chill-producing" music alone or in a group wrth ~nends, less chills occurred in the group condition, poilHing to another interesting facet of the chrll.response, at least in our Western culture: Chilisare frequently perceived as intimate and even hnked to a . . sense of shame (Egermann et al. 2011). In a series of studies we attempted to identify musical factors such as structural charactenstics, harmonic progressions, timbre of instruments/voices, and loudness .developments cont~· ibut­ ing to elicit a chill response. The results were quite disillusioning .. Fmt, ther~ was no simple stimulus-response relationship, i.e., even in music believed to be ~rghly emotwnally arousmg the chill responses remain rather casual and not simply repro~uCible . Secondly,. there was no combination of musical factors producing chills in a fairly rehable manner. Tim was already demonstrated by Guhn et al. (2007), who strived to maximize chill responses in listene~s by experimenter selected "chill music," unfamiliar to the subjects. Only 2~-35% of the subject~ perceived chills in the respective passages from works of Mozart, Chopm, and .Bruch. In. ~ur experiments, the only general factor identifiable as a necessary, how~ver not suf~Cient, conditr~n to induce a chill response was a change in musical structure, or, m the termmology of David Huron's ITPRA model, a nonfulfillment of expectancies (Grewe et al. 2007b). In a group of 38, quite heterogeneaus subjects (age range 11-72 years, 29 females.' five professianal musicians, 20 amateur musicians, and 13 non-musicians) we analyzed musical.parameters oftheir favorite music, producing a chill responsein the laboratory. In 29% ofthe pieces, the entry of a voice, irrespectively of whether it was human or of an instrument, could be identified

324 / A CONTRIBUTION TO THE EVOLUTIONARY BASIS OF MUSIC THE CH ILL RESPONSE IN MUSICASAN EX AMP LE OF EMOTIONAL PEAK EXPERIENCE Chili reactions to "Breit über mein Haupt"

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Chiliresponse data (top panel) for one participant across 7 days in response to Strauss' re1 u er me1n Haupt" with · . ch·/1 . . accompanylng psychoacoustlc analysis presented in the bottom pane/ an~ f~~~~ontse co~slstency 15 evident at t = 1 min . Psychoacoustic parameters loudness roughness . ua IOn s ow peaks at this point in time. However, the other chi/1 responses va~y consider-' ab/y over the 7 days Wlth a general tendency to habituation. Modified from Grewe et al. 2007a with ~~::~~~ Reproduced from Grewe, 0, Kopiez, R., and Altenmüller, E. (2009) The chi/1 para~eter: 61-7 c ps and shlvers as promlslng measures in emotion research . Music Perception, 27 (1), . 4 . © 20f09 by the Regents of the Un1vers1ty of California. Reprinted with permission from the Un1vers1ty o Cal1forn1a Press.

(Grewe et al. 2007b). Furthermore, in 19% a peak in loudness andin 14% a peak in sharpness was found. When looking closer to the latter parameters, the increase in loudness was prominent in the high register (between 920-4400 H z), thus contributing directly to the parameter "sharpness." Less salient was the increase in roughness. In 12% of the chill responses an increase in roughness linked to a reduced signal-to-noise ratiowas observed. The latter reflects an increase in acoustic "density" (Grewe et al. 2007b; Nagelet al. 2008). Transferred to music, this occurs, for example, when more instruments are playing or more voices are singing with high er loudness and tempo. Behaviorally, all these acoustic changes are accompanied by an increase in arousal, which was confirmed in real-time self-reports using the device "EMuJoy" that allowed monitaring the self-declaration of feit valence a nd arousal on a two-dimensional coordinate system (Nagelet al. 2007). A typical example including all the mentioned criteria is the "Barrabas-Call" in St Matthew's passion ofJohann Sebastian Bach. Interestingly, this example was the most frequently quoted in John Sloboda's first pilot study on strong emotions when listening to music (Sloboda 1991) . However, the chill responsein the "Ba rrabas-Call " is not reflex-like, since it varies depending on many factors, such as the listening situation of the individual, overall well-being, attentional factors, and day-form (Grewe et al. 2009b). With respect to the listeners' factors in the earlier mentioned heterogeneaus group, strong chill responders differed from those not perceiving chill responses in several respects: they were more familiar with classical music, rated music as more important for their Jives, identified more with the music they preferred, and listened more readily to music in everyday life (Grewe et al. 2007b). Of course, it could be discussed further whether these features are a consequence of the participant's proneness to pleasurable experiences in music or whether they contribute a priori to a higher susceptibility for chill responses in music. Concerning psychological tra its, chill responders showed a general tendency for less intensive stimuli, as operationalized by Zuckerm an's sensation seeking questionnaire (Litle and Zuckermann 1986) and were more reward dependent, i.e., they especially liked approval and positive emotional input from their environment. Since familiarity with musical genre and personal emotional memories seemed tobe important factors in the production of chill responses, we addressed the roJe ofthe individual musical biography in a furth er experiment (Grewe et al. 2009) . The goal of this study was to induce chill responses more reliably and to ga in further insights into the factors influencing it. We recruited 54 subjects from three different amateur choirs who had performed Mozart's requiem, furth er referred to as "Mozart group," and 41 participants from gospel and pop choirs, further referred to as "control gro up," who were unfa miliar with theMazart requiem and with classical music in general. We exposed these subjects to emotionally moving excerpts from Mozart's requiem ("Lacrimosa," "Confutatis," "Rex tremendae," "Tuba mirum ," "Dies irae"), which were either recordings of themselves or of professionals . Furthermore, excerpts from the Requiem ofPuccini and from the Bach motet "Our life is a shadow," which had been sung in each case by only one of the three choirs of theMazart group, were played. As measurements, subjective real-time rating of the intensity of the feelings, and perceived chill responses were recorded using the softwa re EM uJoy (Nagelet al. 2007). Additionally psychophysical measures such as sk in conduction response (SCR), sk in conduction Ievel (SCL), heart rate (HR), and breathing rate (BR) were assessed. Fig. 19.4 shows the time course of psychophysiological data 10 s prior, during and 10 s after the chill responsein a grand average. The two most sa lient features of the physiological responses are (1) the increase of SCR abou t 2 s before the chill reaction and (2) the response after the chill of about 4-6 s which has recently been ca lled th e "afterglow effect" (Schubert 2012) .

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Overall, comparable to previous results of Goldstein and Guhn, only about two-thirds of the participants reported a minimum of one chill during the experiment. There was high variability in chill responses, ranging from a maximum of 88 chill responses in one subject to no chills at all in others. On average, each participant experienced nine chills during the experiment. Interestingly, chill responses showed no relation to age, gender, or knowing and liking of classical music. However, familiarity with the stimuli influenced the frequency of chill responses. Chilis occurred more frequently in the Mozart group than in the control group (72% vs. 56% of the participants), and the overall number of chills was much higher in the former than in the latter (679 vs. 173 chill responses). Furthermore, whilst listening to the Bach motet and the Puccini excerpts, chill responses were significantly morefrequent in the choir members familiar with the respective pieces. However, it seems not to be very important to Iisten to one's own interpretation, since only the Confutatis interpretation of one choir produced slightly more chill responses in the choir members as compared to the professional version (in average 0.95 chills vs. 0.11 chills). Thus, obviously, familiarity with the stimulus is an important factor in eliciting chill responses. Musical biography and individual associations, for example, the remembrances of a successful performance in an awe inspiring gothic cathedral, may weil promote emotional susceptibility. Summarizing this paragraph with respect to the overall topic of this review, namely the evolutionary adaptive value of music, the ch ill response is biologically grounded in an ancient reflexlike response of the sympathetic nervous system related to thermoregulation and intimidation displays. It is biologically linked to arousal and facilitates memory formation. In humans, the chill response occurs in the aud itorydomain in the context of negative arousal and alarm, mainly linked to aversive loud and high-frequency noise, andin the context ofhighly pleasurable events leading to activation of the dopaminergic reward system in the brain. Factars facilitating these positive chill responses include: structural changes, beginning of something new, increase in loudness in the high register, and personal emotional memories linked to positive associations and liking of the music. Chiliresponsesare morefrequent in moresensitive and social personalities. In the next section we will demonstrate how the chill response may be linked to an adaptive value of music in human evolution. Lastly, we will develop our model of "mixed origins of music" in human evolution.

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The evolutionary adaptive value of the chill response is at hand when considering the ea rlier-mentioned biological concomitants. Negative chill responses may have been direct, aro using reactions towards the piercing sounds of a hunting predator or by the shrieking calls of conspecifics attacked by an enemy or predator (Owren and Rendall 2001) . They may be part of an evolutionary ancient inter- and intraspecific affective signaling system of alarm calls and pain shrieks, observable today in many socially living mammals, for example, in tree-shrews and vervet monkeys (see alsoZimmermannet al. this volume, Chapter 8). Thesesounds furthermore support avoidance behavior in order to increase the distance to the sound source. In this way, close contact to a potentially dangeraus predator is prevented, but also the delicate sensory organ of the cochlea with its hair cells, susceptible to high so und pressure Ievels damage, is protected (Subramaniam et al. 1995). Finally, in agonistic contexts, an intimidation display is activated to frighten the predator. In human evolution, the roots of such a behavior may weil dateback to some 3.2 million years when we, or better our about 1-m tall evolutionary ancestors

328

THE MIXED ORIG IN S OF MUSIC THEORY

I A CONTRIBUTION TO THE EVOLUTIONAR Y BASI S OF MUSIC

Australopithecus afarensis, prowled through the high grass of the Centrat African dry steppes, haunted by the piercing sounds of some !arge African eagles, hunting for prey. Explaining the evolutionary adaptive value of the positive chill response is more difficult. There are two proven and one hypothetica l features, which can be related to an adaptive value of music. First, it is the fact that "surprise," for example, the earlier-mentioned "not fulfillment of expectancies," contributes regularly to the chill response. Since accompanying arousal and activation of the reward system improves memory formation, this kind of acoustic Stimulation may weil have enlarged the repertoire of auditory patterns remernbered by our ancestors and furthermore fuelled curiosity to detect novel auditory stimuli. This, in turn, was of evolutionary relevance, since fast and precise classification of acoustic stimuliwas a prerequisite for optimal adap tations of behavior (for example, avoidance of a sneaking predator at night and perception of subtle nuances of intraspecific affect voca lizations). We therefore specu late that one of the driving forces of the development of our superior auditory memorywas the reward gained by identification of novel acoustic patterns. We wi ll even go further claiming that the first song-like voca li zations, the first artificial sounds produced by primitive instruments (for example, wooden drumsticks hit on hollow stumps) may have constituted a safe playground to train auditory discrimination. Furthermore, vocal production abilities improved and reinforced curiosity to detect novel sounds long before language emerged, thus establishing prerequisites to develop the latter. The second feature of the positive chi ll response implicating an evolutionary adaptive value is pleasure induction by music. Rooted in the activation of the sympathetic nervaus system and of centrat nervaus reward circuits, music as a TTM could add moments ofhappiness and comfort to the hard lives of early modern humans living in hostile environments. The Hohle Fels and Geissenklösterle caves, for example, were located in alpine tundra at the time the flutes were constructed, 35,000 years ago. Average temperatures were comparable to present day Green land . Albeit food was readily available, due totherich wi ldlife, musculoskeletal diseases, gastrointestinal infections, parasites, toothache, and the omnipresent cold rendered life cumbersome. Music could provide moments of well-being, of forgetting the daily hardship not only by producing aesthetic emotions but also by giving rise to occas ional emotional peak experiences, which then reawakened Iove oflife. Finally, the third potential feature with evolutionary adaptive value is the frequently quoted "separation ca ll " theory by Jaak Panksepp (1995) . It proposes that the evolutionary origin of music-induced positive chill responses is a soothing and "warm ing" function of matemal monkey vocalizations on the offspring. Unfortunately, this theory has not yet been verified by empirical research. An argument speaking against such a mechanism is the lacking evidence of aco ustically evoked chill responses in infants and toddlers, for example, when listening to soothing lullabies . Possibly, such a phenomenon may have been overlooked up to now. However, in informal interviews with children and adolescents, it seems that the ea rliest descriptions of positive chill responses emerge just before reaching puberty. Admittedly, systematic empi rica l research on this interesting topic is missing. In short, when hypothetically summarizing the long history of chill responses, we argue that in the beginning it was predominantly related to a reflex-like mechanism, involved in thermoregulation. This was based on neuronal networks of the autonomaus nervaus system, involving thermoceptive afferents from the skin and efferent activation of the sympathetic nuclei. These reflexes were additiona lly activated by exposure to aversive stimuli, such as shrieking sounds, sour food, or enteroceptive pain, producing a threatening display and enlarging the appearance

I 329

. . . nditioned reflexes, the trigger of such a threatening display could be by hatr-rars111g. D.ue to co . . the chill res onse activated memory formation. Finally, after modified by learn111g and vrce ver·stal . pth contextofthermoregulation and threatening ·fi 1 ffur ehr responses 111 e , . 1 d ld be used for other purposes. The acoustically the human-spect c oss o . b b" 1 · ally meamng ess an cou drsp1ays ecame 10 ogrc d conditioned cou1d be powerfully . . fl 1.k .. h "ll ses prev10us 1y re ex- 1 e an ' mediated posrtrve c r . respon : . d r risin acoust ic stimuli with chill responses used for auditory learmn~. Reward111g ~ew ~n1 su p d su~J·ective feelings ofwell-being cou1d be . . . db d ·phm and dopam111e re ease an accompame Yen 01 . . constituting a prerequisite of drfferentrd" the most important driving force of au rtory 1.e~t n111g,

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ated communication be part of our human condiWhat are then the ongms ~f musrc :~~~r MOM theory. This will be achieved by integrating tion? In the follow111~, we wrll ~xpo~ ada tive or later acquired, or recently refined properties p , h h . . evolutr"on and anthropolsevera1 aspects of anoent evolut10nary h" h . many ot er t eones 111 of music. We are aware that t rs .t eor~, ~s . . ords of musical activities until the first ogy, cannot b~ directly proven, Sll~e~o::v::.e:eos:r~i~e to strengthen our arguments by drawhuman-made mstr~ments appeare . "b1, d by referring to physiological and neuronal . h" t . ing on a comparatrve approach when possr e an

adaptations which m~st proba~l: dateback ~~~g in ~u:~~h~:=~;~:~~h:: ;r:~ic may have several As we have exemphfied earher wrt.h thbe clt respmr"l1r·o' ns ofyears and some acquired in later . h 1 t " some datmg ac < many roots 111 um an evo u 10n, . . f fi (B. t 1 2009). Allthese posTTM com arable to the mventron o re rown e a . . . trmes possrbly as a . , p 11 lusive· rather they demonstrate its richness and mulsib1e origins of musrc arenot mutua y e~c , 11 ,1 r·n the many effects music can have in Th . ts of musrc may we exp a tifaceted nature. e many roo f h t t. development of music out of an ancient humans. In Fig. 19.5 we provrde a scheme o t e pu a rve affective signa ling system, e1aborating our MOM theory.

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THE MIXED ORIGINS OF MUSIC THEORY 330

I A CONTRIBUTION TO THE EVOLUTIONARY BASIS OF MUSIC

In the very beginning, intraspecific and interspecific affective communication amongst prehumans included shrieking calls of threatened conspecifics and of alarm calls of threatening predators, producing a heightened arousal which may have been accompanied by aversive chill reactions as it can be witnessed today in many socially living mammals. An important step in evolution must have been the generalization of these chill reactions to affiliative sounds and vocalizations with positive emotional valence. These may have been related to parent-offspring communication, although experimental evidence for the separation-call theory is stilllacking. However, irrespectively of the emotional valence induced, the chill reaction fastered auditory memory formation by activating the brain's reward systems when new acoustic patterns were perceived. It is unclear whether vocalizations, probably some hundred thousands yea rs ago, were related to music in the narrow sense of the word or whether they were part of an ancient affective signaling system comparable to Steven Mithens "hmmmmm" proto language (2005, p. 172), which means that music was part of a communication system which was holistic, multi-modal, musical, and mimetic. However, the brain's reward system activation and improved memory consolidation linked to detection of violations of auditory expectancies may have triggered the superior auditory discrimination faculty of early humans, which in turn constituted the prerequisite of language acquisition and production. Here, we argue that chill responses to novel melodic contours, timing subtleties, timbre variations, and structural breaks Iead to superior classification abilities and in turn to a !arge repertoire of language-like vocalizations, apt to replace, for exa mple, maimal gesturing as a means to organize distributed Iabor in complex social groups (e.g., Corballis 1992). In parallel with this very fundamental aspect ofhuman auditory learning, music also contains aspects of a TTM, which may have developed much later than language. Besides the previously mentioned positive chill response as a source of pleasure when exposed to refined tunes and playful manipulation ofincreasingly complex melodies, harmonies and rhyt hms, other facets of music are good candidates for a TTM: As Pate! (2010) exemplified, group cohesion, and coop erative behavior are supported by joint clapping and dancing, relying on the human ability to synchroni ze in time with an external beat. According to Pate!, this capacity is related to vocal learning, and may thus be an epiphenomenon of our language abilities. However, as demonstrated ea rlier, our language capacity in turn may be grounded in our auditor y classification abilities facilitated by the chill responses. A more convincing musical TTM aspect is the processing of musical hierarchies in tonality. These cognitive processes rely partly on the same neuronal resources as syntactic language processing. Brain activation studies showed overlapping neuronal networks when violating either harmonic musical rules or linguistic syntactic rules (for a review, see Koelsch and Siebel2005). According to Pate!, both tonal music and la nguage rely on the same basic cognitive operations, namely the general building of mental hierarchies. Thus, our ability to identify and create tonal hierarchies in music and syntax in language emerges from a common evolutionary old capacity, which is on one side transformed into linguistic syntax and on the other into musical tonal hierarchies. Here, a word of caution is necessary since many forms of music such as MinimalMusic, Rap, or m any types of Ethnic music in Africa, do not contain tonal hierarchies (for a review, see Stevensand Byron 2009). On the other hand, hierarchies and rules, whether tonal or temporal, are almost universally found in both music and language and thus may indeed share a common "rule detector" mechanism in the brain, which is ancient and evolutionary adapted (Brown and Jordania 2011). Many other effects of music may additionally be considered as constituents of an evolutionary late acquired TTM. To name but a few, the roJe of music in improving health status and

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. . d t" ts with basal ganglia disorders may serve as examples rehabilitation II1 stroke patients an_ p~ Jen! 2010· Thaut et al. 2001). Similarly, improving mem.. .. t 1 2008· S Schneidereta · , l"d (e.g., Särk_amo_e a . . , . . Vink et al. 2003) and supporting memory conso I aory functiOns 111 Alzheimer patient~ (e.g., 1 h . d " "d ls (Wallace et al. 1994) may be regarded tion of complex linguistic contents 111 hea t y 111 IVI ua .. "d fc ct of such a musical TTM. . . t as a positive s1 e e 1e . f . Cl.ent affective commumcatiOn sys em, th t n the basis o a very an . . . In summary, we argue a o . ·ng r·efinement of auditory discnm111a. . d d and Jed to an 111creas1 auditory learmng was rewa~ ~ h" h laid the ground to acquisition of langnage . . . . h d tirning T IS may ave d" tion abiht1es 111 pltc an · . t"t t d a sace playground for new au 1tory f . h "ch 111 turn cons Ill e 1' and also to our Iove o music, w I . 1 functi·ons increasing our chances of . · dapted for many soc1a , . expenences. Later, musi_c _was a d b ddin pleasure and aesthetic emotions to our survival by better orgamz111g of groups an y a g hard Jives . . t . h k of brevity many aspects of our MOM theory This review is not exhaustive and OI t e sabe d " d cri"tically For instance, we did not . d ffi . tJ and may e !SCUSSe . . could not be considere su cien y . . d"t" that strongly supports the exlstf gemtal amusia, a con 1 1on . . comment on the phenomenon o con I ·k d signed for refined pitch discnm111a. ld . l"zed neurona networ s, e . ence of evolutiOnary o , speCia 1 .h t t th positive chill response we admlt that . A 1 2002) W1t respec o e tion (for a review, see yotte et a . . : I . d. .d I linked to personal memories, and even in present times this phenomenon lS high y ~~/VI ~:~thermore it is usually elicited by highly unknown to about 30% ofthe Western popu a .IOn. h or a ~eatles song. We do not know . . h as a Bruckner symp ony complex acoustlc pattel ns, suc . . th H hle Fels and Geissenklösterle, but .· d by our ancestms 111 e o how flute tun es were expetlence . f low exposure to music, and artificially pro. . . bl to assume that at times o . . we belteve lt IS reasona e h . t could have a strong emotiOnal Impact. . 1 ·mple monop omc une duced sound 111 genera, even a SI .. h"ll ·esponse isamusicaluniversal or whether · · h th r the pOSitiVe C I 1 Another open questiOn IS w e e_ . . h .t h domain restricted to a limited number of it is predominantly linked to vanatiOns II1 t. e PI lcl c nd this would weaken our argument of ·· h ·u were not umversa Y1ou , music cultures. I f positive c I s . Th" b . ngs us to another perspective, namely that . · ( al reactiOn. IS n . · an evolutiOnary anCient emo Ion d to li.sten to music seated 111 a chatr nce of our mo ern way b the chill response may e a conseque_ h h h mparable to a "sublimation" of our "b"l" bodlly to t e r yt ms, co ' without the possl I lty to move . h th impact ofbodily movements on the naturalurge to move to music. Systematic researc on e n experience contains many facets and may positive chill response is still_lacking. 1 .· h h uma To end with, music as an Immense y nc have many effects: Orpheus with bis Jutem ade trees, And the mountain tops that fre eze, Bow them selves, when he did sing: To his music plants and f\owers Ever spr ung; as sun and showers There had made a Iasting spring. Every thing that heard him play, Even the billows of the sea, Hung their head s, and then lay by. In sweet music is such art, Killing care and grief ofheart Fall asleep, or he ar ing, die. (W illiam Shakespeare, He11ry VIII, 3.1.4-lS .)

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Acknowledgments This work was supported by the DFG (AI 269-6). Furthermore we would like to thank the many friends and colleagues who gave valuable input to this paper in many discussions on the origins of music. Here, we wou ld like to thank especially Dr Andre Lee, Dr Thomas Fritz, Floris van Vugt, Professor Dr Elke Zimmermann, Dr Sabine Schmidt and the members ofthe IMMM. We furthermore would like to thank Marta Beauchamp for carefullanguage editing.

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