99655Avanzini

INTERPLAY OF THE TWO CULTURES: NEUROAESTHETICS

Neuroscience and music

Giuliano Avanzini

Received: 3 November 2011 / Accepted: 27 April 2012 / Published online: 21 July 2012! Accademia Nazionale dei Lincei 2012

Abstract Music is part of all known human cultures,which suggests that musical competences are encoded in

the human brain. Attempts to define the cerebral organi-

sation of music perception and production date back to thebeginning of the modern era of the neurosciences, but the

subject has been adequately developed on only over the last

two decades. The combined approaches of neuropsycho-logical, neurophysiological and neuroimaging methods

have demonstrated that the musical information reaching

the primary auditory cortex undergoes a series of processesinvolving both hemispheres, but with predominance of the

right. Particularly interesting is the demonstration of

overlapping brain mechanisms that create and maintainlearned sound categories in both the linguistic and musical

domains i.e. Brocas area, whose functional properties

hierarchically organise perceptually discrete elements(words or musical tones) into structured sequences based

on syntactic principles. Only a few studies have addressed

the challenging task of investigating the neurobiologicalbasis of music by aesthetic categories. One possible

approach is to take advantage of neurophysiologicalmethods capable of detecting brain responses to novel

sound combinations as a signature of creativity. Most

sound data relate to the brain processing of music-relatedemotions. The results of studies carried out over the last

10 years demonstrate that intensely pleasurable responses

to music correlate with activity in the brain regionsinvolved in the emotional colour of biologically important

activities such as nutrition and reproduction. Whether such

structures are of general interest for the perception of theartistic quality of music is an exciting question that needs

to addressed by comparing the results obtained in other

domains of neuroaesthetics.

Keywords Music ! Neuromusic ! Neuroaesthetics ! Musicperception ! Music and languages ! Music and emotions

1 Introduction

It is probably impossible to find a single human beingbelonging to any culture who has no experience of music.

However, although everyone would say that he knows what

music is, any attempt to explain all of its relevant aspectswould be consistently inadequate. Even the popular dis-

tinction between noise and music is inappropriate, because

the noise produced by percussion instruments forms part ofthe acoustic material currently used in music. Conse-

quently, when searching for a definition of music, we areleft with the tautological statement that music is a set of

musically organised sounds. The definition of music by

aesthetic categories is even more elusive: what gives a setof musically organised sounds the quality of an artistic

product? Nobody could disagree with Edward Hanslicks

description of the concordance and opposition of sounds,their running away and catching each other, their growing

up and dying, this is what our spirit perceives as beautiful

(Hanslick 1854), but whether any given musical composi-tion is perceived as artistic by different listeners depends

on such a multifactorial process that the result is often

unpredictable.

This contribution is the written, peer-reviewed version of a paperpresented at the Golgi Symposium on Perspectives inNeuroaesthetics, held at the Accademia Nazionale dei Lincei in Romeon June 13, 2011.

G. Avanzini (&)Fondazione IRCCS Istituto Neurologico Carlo Besta,Via Celoria 11, 20133 Milan, Italye-mail: [email protected]

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DOI 10.1007/s12210-012-0180-6

These considerations suggest that any attempt to discuss

the role of the neurosciences in fostering our understandingof musical abilities in a neuro-aesthetic perspective is so

difficult that I am challenged to try.

2 Music and the brain

The relationship between music and brain functions or

dysfunctions has been a subject of interest since timeimmemorial. In ancient civilisations, chants and the sounds

of instruments were used to induce a positive effect on an

individuals cognitive and psychological functioning, andto cure melancholy and other psychopathological altera-

tions. However, it was only when the fine structure of the

brain and its correlations with sensory and motor functionscame to be known that it was possible to investigate the

cerebral organisation of musical competences.

The first attempts to localise musical functions can befound in papers published in 1865 by Bouillaud and 1888

by Knoblauch, who coined the word amusia and ana-

lysed the disturbed musical functions of patients with braindiseases. However, although interest never ceased, there

were only relatively few further publications until 1977,

when the now classic book Music and the Brain, editedby Macdonald Critchley and R. A. Henson (Critchley and

Henson 1977), came out. This book substantially alerted

the scientific community to the neurological aspects ofmusic by providing a complete account of the literature on

the subject: no more than thirty papers, including that of

Alajouanine (1948) on Ravels degenerative encephalopa-thy and the paper by Luria and Tsvetkova (1965) on

Shebalins stroke.

Subsequently, various groups gradually emerged thatgave a new impetus to investigating the relationships

between the neurosciences and music. Although I risk

forgetting some important name, I would mention Robert JZatorre and Isabel Peretz in Montreal, Carol L Krumhansl

in New York, Diana Deutsch in San Diego (la Jolla),

Sandra Trehub in Toronto, Eckart Altenmueller inHannover, John Sloboda in Keele, Mireille Besson in

Marseille, Christo Pantev in Munster, Mari Tervaniemi

in Helsinki, Timothy D. Griffiths in Newcastle-upon-Tyne,Joseph P. Rauschecker in Washington, Gottifried Schlaug

in Boston. These and the many other groups that have

entered the field over the last 10 years were present at theconference The Biological Foundations of Music (New

York 2000) sponsored by the New York Academy of

Sciences, and at the series of Neurosciences and Musicconferences sponsored by the Mariani foundation of

Milano (Venice 2003, Leipzig 2005, Montreal 2008, and

Edinburgh 2010). These meetings originated a series ofvolumes published in the Annals of the New York

Academy of Sciences, which reflect the development of

Neuromusic in the last 10 years (Zatorre and Peretz 2001;Avanzini et al. 2003, 2005; Dalla Bella et al. 2009).

A Medline search shows that about 2,000 papers have

been published in qualified scientific journals since 1964,three-quarters of them in the last 12 years. This bears

witness of the growing interest in this field of the neuro-

sciences and, although this is not surprising (because thepresence of music in all human cultures indicates that

musical competences are encoded in our brain), what israther surprising is the fact that it developed so much later

than the interest in language, which has been one of the

main objects of brain research ever since Brocas pio-neering observations (Broca 1861).

The investigations have taken advantage of the wide

range of modern techniques and have led to impressiveadvances in our understanding of the musical brain. The

results were made possible by the development of methods

capable of analysing the specific musical quality of theacoustic stimuli used in the experiments.

3 Methods

3.1 Neuropsychology

The classical approach to the study of cognitive functions

is based on tests specially designed to assess the func-tion(s) of interest. To investigate musical abilities, Isabelle

Peretz has developed a battery of musical tests since 1987

(the Montreal Battery of Evaluation of Amusias, MBEA;Peretz et al. 2003) based on a modular model of the pro-

cessing components involved in music perception (Fig. 1).

She has subsequently adjusted and validated the MBEAusing different groups of individuals and neurological

patients, and has thus provided the scientific community

with a standardised instrument for assessing cognitivemusical functions (Peretz et al. 2003; Peretz and Coltheart

2003). The MBEA includes a series of tests designed to

assess the ability to recognise melodic organisation (con-tour, interval scale), temporal organisation (rhythm and

metre), and musical memory. The tests are based on a pool

of 30 musical phrases written with sufficient complexity toguarantee its processing as a meaningful structure rather

than a simple sequence of tones. The principle is to

introduce manipulations that alter the original sequences,and the task is to recognise that the manipulated stimulus is

different from the original. In the case of the melodic

organisation tests, the pitch is modified so that it is out ofscale (scale-violated melody), the contour is changed

without affecting the key (contour-violated melody), and

the interval is changed while preserving the original con-tour and scale (interval-violated melody). In the case of

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temporal organisation, the manipulation affects the dura-

tion of two adjacent tones, which leads to a change inrhythmic grouping while retaining the same metre (rhythm-

violated stimulus), or the metre itself (metre-violated

stimulus). The final test concerns the ability to recognise apreviously presented melody in a group of new melodies

(memory recognition).

The MBEA principle of using acoustic material in whichmusically relevant parameters have been violated has been

widely adopted in investigations of the brain processing of

music, in which cerebral responses are revealed by meansof electrophysiological recordings or functional imaging

techniques.

3.2 Neurophysiology

While searching for perception-related electrical activitiesof the brain, Naatanen et al. (1978) discovered a negative-

going peak of event-related potentials (ERPs) that was

elicited by a deviance within a sequence of otherwiseregular stimuli (Fig. 2). Currently known as mismatch

negativity (MMN), this can occur in any sensory system

but has been most frequently studied in terms of auditionand vision. Auditory MMN is a fronto-central negative

potential with sources in the primary and non-primaryauditory cortex, and a typical latency of 150250 ms.

Further investigations by Mari Tervaniemi of the same

Helsinki group (Saarinen et al. 1992) demonstrated that

MMN provides a suitable index of the detection of deviant

sounds in a regular sequence (Tervaniemi 2001). MMN canalso be recorded by means of magneto-encephalography

(MEG), which has some advantages over electroencepha-

lography in terms of source analysis. The main MMNgenerator is located within and near the primary auditory

cortex, with a small contribution coming from the oper-

cular part of the inferior frontal gyrus.Another deviance-related component of ERPs is early

right anterior negativity (ERAN), which was first described

by Koelsch et al. (2000) and named by analogy with earlyleft anterior negativity (ELAN), an ERP effect that is

elicited by syntactic irregularities during language per-

ception (Friederici 2002). ERAN is usually maximalbetween 150 and 250 ms, has an anterior scalp distribution

and often right-hemispheric prevalence, and its main gen-

erators seem to be located in the inferior fronto-lateralcortex (Brodmanns area 44), with additional contributions

coming from the ventrolateral premotor cortex and the

anterior superior temporal gyrus (Fig. 3). Both ERAN andMMN are elicited by acoustic events that do not match a

preceding sequence, but they differ in terms of topography;

furthermore, the model of regularities relevant to MMN isextracted online from the acoustic environment, whereas

the structural model of ERAN processing is based onrepresentations of syntactic regularities that already exist in

a long-term memory format. In other words, the repre-

sentations of regularities building the structural model of

Acoustic input

Acoustic analysis

Pitch organization TemporalorganizationAcoustic-toPhonologicalconversion

Rhythmanalysis

Meteranalysis

Contouranalysis

Intervalanalysis

Tonalencoding

Emotionexpression

analysis

MusicalLexicon

PhonologicalLexicon

Vocal planformation

Associativememories

Singing Tapping Speaking

Acoustic input

Acoustic analysis

Pitch organization TemporalorganizationAcoustic-toPhonologicalconversion

Rhythmanalysis

Meteranalysis

Contouranalysis

Intervalanalysis

Tonalencoding

Emotionexpression

analysis

MusicalLexicon

PhonologicalLexicon

Vocal planformation

Associativememories

Singing Tapping Speaking

Fig. 1 Modular model of musicand language processingaccording to Peretz andColtheart (2003). The followingprocessing components: tonalencoding, interval analysis,contour analysis, musicallexicon, vocal plan formationand singing are specific tomusic. Component whosespecificity to music arecurrently unknown are: rhythmanalysis, meter analysis, andemotion expression analysis. Aneurological anomaly mayeither damage a processingcomponent or interfere with theflow of information betweentwo components (redrawn fromPeretz and Coltheart 2003)

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the acoustic environment are more sensory in the case of

MMN, and more cognitive in the case of ERAN (Koelsch

2009).We have used a different neurophysiological approach

in our laboratory to investigate brain responses to musical

material by analysing the ERPs elicited by individual notesbelonging to structured or unstructured sequences (Minati

et al. 2008, 2010). We consistently observed differences in

the latency, duration and topographical distribution of theN2 components elicited by structured or unstructured

stimuli (Fig. 4), but their significance in relation to the

musical quality of the stimuli is still under investigation.

3.3 Neuroimaging

Because of their unique ability to describe neural activities

in real time at a scale of milliseconds, neurophysiological

techniques are first-line methods in neuroscience investi-gations, but they are much less satisfactory in terms of

spatial resolution, which is one of the main strengths of

imaging techniques. Combined neurophysiological andimaging investigations (such as can be obtained by making

neurophysiological recordings in the context magnetic

resonance imaging, MR) have been profitably used inneuromusic research as in many other fields of the neuro-

sciences. Moreover, functional magnetic resonance imag-

ing (fMRI) has effectively advanced our understanding ofthe musical brain whenever a high-time resolution is not

required. Figure 5 shows a good example of the power of

fMRI to investigate the cortical representation of the sound

Fig. 2 Mismatch negativity response to a 2.5 % pitch change.a Event-related potentials (ERP) to a frequently presented standardtone (thin line) and to an infrequently presented deviant tone (thickline). The x axis represents the time after the sound onset inmilliseconds and the y axis the strength of activity in microvolts.b The waveform is calculated by subtracting the ERP to standard tonefrom the ERP to deviant tone. On the schematic brain the recordingelectrodes (Fz, Rm) are indicated (from Tervaniemi 2001)

Fig. 3 Early right anterior negativity (ERAN) elicited by irregularchord sequences. a chords built on the tones of the C major scale.Within the tonal hierarchy the chord on the first scale tone (tonicchord) is the most stable chord, followed by the dominant and thesubdominant, whereas the submediant and the supertonic representless stable chords. The major chord on the second tone of a majorscale can be interpreted as the dominant to the dominant (squarebrackets). In majorminor tonal music, chord functions are arrangedwithin harmonic sequences according to certain regularities, forexample the dominant-tonic progression shown in b (left) is a typicalmarker for the end of a harmonic sequence. On the contrary theharmonic progression shown in b (right) ending on dominant todominant cord is highly irregular, c the final chord of the irregularsequence shown in b (right) elicits an ERAN better detectable in thedifference wave, which represents regular subtracted from irregularchords. d With MEG, the magnetic equivalent of the ERAN waslocalized in the inferior frontolateral cortex. The dipoles indicate thegrand-average of single-subject dipole solutions (modified fromKoelsch 2009)


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parameters relevant to music; the imaging subtraction

procedure is particularly useful when comparing differ-ences in the functional status of the brain with respect to a

specific system.

The measurement of regional cerebral blood flow bymeans of positron emission tomography (PET) is also

useful for mapping the cerebral areas involved in musicprocessing, not least because the images can be superim-

posed on MR images using fusion techniques. Figure 6

shows some examples of the results that can be obtained

using this method.

4 Anatomo-functional organisation of the musicalcompetences of the brain

Like any other type of acoustic stimulation, a musical

stimulus activates the peripheral and central systems that

Fig. 4 Event-related potential traces (solid line, average; dotted lines,average7SD) from the mesial frontal (Fz) and central (Cz) electrodes,for notes in melodies and unstructured note sequences. The two traces

clearly differ in the N2 component that is delayed and increased induration when evoked by structured stimulus. Left column musicians,Right column non-musicians (from Minati et al. 2008)

Fig. 5 fMRI activation elicited by different auditory stimuli. Theposition and orientation of the sections is depicted in the bottompanels. Note that unlike the standard use the left hemisphere is on theleft side and the right one to the right. Note also that the axial sectionis tilted relative to the horizontal plane to show the surface of theplanum temporale on which (structural MRI, bottom) the Heschlgyrus is represented as a white area. The differential activation,

obtained by contrasting noise to silence, fixed pitch to noise, tonicmelodies to fixed pitch and random melodies to fixed pitch aredepicted in different grey gradation (from darker to lighter). Theactivation area elicited by increasingly complex acoustic stimulimoves increasingly anteriorly to the primary auditory area andbecomes increasingly lateralised to the right hemisphere (fromPatterson et al. 2002)


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convey the information to the primary auditory cortex(Brodmanns area 41, the supratemporalis granulosa) inthe Heschl gyrus on the superior face of the temporal lobe

(also called the planum temporale). Area 41 is a granularcortex and, like all sensory cortices (also called konio-

cortices), is characterised by a predominance of non-

pyramidal neurons that distorts the eulaminated pattern ofnon-specialised isocortices (such as area 42 that surrounds

area 41). A topographically ordered frequency representa-

tion is clearly recognizable in the primary auditory cortexwith high frequencies being represented mediocaudally and

low frequencies rostrolaterally. Subjects with right lesionsinvolving the primary auditory cortex have deficits in pitch

perception. A detailed analysis of the topographical dis-

tribution of the responses to pitch and melody on the pla-num temporale has been carried out by Patterson et al.(2002). The experiments showed that the representation of

noise, notes with a fixed pitch, tonic melodies and randommelodies only partially overlap, because the topographical

representation of the activations elicited by increasingly

complex acoustic stimuli moves increasingly anteriorly tothe primary auditory area and becomes increasingly lat-

eralised to the right hemisphere (Fig. 5).

It has been shown that other cortical areas within oroutside the temporal lobe are activated when listening to

music or discriminating specific music components (e.g.

musical tempi and metre, Fig. 6). The order of activation(and therefore the significance of the activations) is

sometimes difficult to define because of the limited time

resolution of imaging techniques. The analysis of theauditory cortex organization supports the role of the

superior temporal gyrus for complex sound processing

(Rauschecker 1998). Comparative studies in non-human

primates and humans led Rauschecker and Scott (2009)

and DeWitt and Rauschecker (2012) to identify the supe-rior temporal gyrus and its projections to ventrolateral

prefrontal cortex as the auditory anteroventral stream for

language and music processing. Liegeois-Chauvel et al.(1998) studied the specific contribution of different tem-

poral areas in patients who had undergone temporal cor-

tectomies for the treatment of drug-refractory epilepsies,and confirmed the importance of the superior temporal

gyrus in melody processing. They also demonstrated adissociation between rhythm and metre, and the critical

involvement of the anterior part of the superior temporal

gyrus in metrical processing.Additional data on the cortical areas involved in music

perception will be provided below, in the sections dis-

cussing the relationships between music and language, andbetween music and emotions.

Further information on the organization of the musical

brain has been obtained from the analysis of brainmechanisms of musical performance. With this approach

Schlaugh (2001), Pantev et al. (2001) and Pasqual-Leone

(2001) provided the evidence of plastic adaptive changesin different brain areas of musicians playing musical

instruments. In particular Schlaugh (2001) and Schlaugh

et al. (2009) found that bimanual motor skill acquisitionthat is necessary to play a musical instrument is associated

to an increased size of the anterior section of corpus

callosum.

5 Music and language

Music and language are human activities that are common

to all human cultures and are both based on sounds that arehierarchically organised within a syntactic framework.

Their mutual relationship is demonstrated by a number of

facts. First of all, the ability of the brain to recognisetransposed pitch intervals, which makes it possible to

organise sounds on the basis of the same harmonic and

melodic patterns regardless of their height, is also a basicprerequisite for linguistic communication. Every vowel is

characterised by the first two formant frequencies deter-

mined by vocal tract resonance, which have a definiteinterval ratio between them and in relation to the funda-

mental frequency (Ross et al. 2007). The broad interindi-

vidual variability of the fundamental frequency (betweenmales and females) does not impair our ability to recognise

a vowel uttered by different speakers because of our ability

to recognise transposed intervals (Patel 2008).Furthermore, a number of studies have demonstrated

that a musical education facilitates the ability to reproduce

the non-familiar sounds of a foreign language and, moregenerally, the acquisition of a second language.

Fig. 6 Activations in left medial frontal cortex (Broadman area 9) fornon-musicians (a) and musicians (b) during the discrimination ofmusical tempi. The regional blood flow, as an index of neural activitywas measured by PET and shown as group mean PET images for eachtask (contrasted with rest) overlaid on MRIs (from Parsons 2001)


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One decisive demonstration of the overlapping brain

mechanisms that create and maintain learned sound cate-gories in both the linguistic and musical domains came

from electrophysiological studies demonstrating that the

ERPs evoked by linguistic and musical incongruities wererecorded from the same cortical region: i.e. Brocas area

(Fig. 3) (Maess et al. 2001; Koelsch et al. 2002; Koelsch

2006; Patel 2003). It is interesting to note that the relevantstimulus in both cases was a syntactic musical or linguistic

irregularity, thus leading to the conclusion that Brocasarea is endowed with functional properties related to

organising perceptually discrete elements (words or musi-

cal tones) into hierarchically structured sequences on thebasis of syntactic principles. These findings that support the

common nature of the mechanisms by which the brain

processes language and music need to be evaluated withreference to the existing differences between the musical

and linguistic domains, which formed the basis of

Aniruddh Patels book Music, Language and the Brain(2008). He exhaustively reviewed evidence taken from the

cognitive and neurosciences and discussed it with reference

to the shared syntactic integration resource hypothesis,which postulates that the two domains have distinct and

domain-specific syntactic representations, but share neural

resources for activating and integrating them during syn-tactic processing (Patel 2003) and can account for obser-

vations of a double dissociation between aphasia and

amusia (Luria and Tsvetkova 1965; Peretz et al. 1994]. Theimpressive bulk of the information supports the authors

statement that music and language are complex constel-

lations of subprocesses, some of which are shared andothers not..Exploring this network of relationscanimprove our understanding of how the mind assembles

complete communicative abilities from elementary cogni-tive processes.

6 Neuroscience and music as an aesthetic experience

At the beginning of this chapter, I quoted EdwardHanslicks statement concerning musical beauty, which

indicates the essence of the aesthetic musical experience as

being captured by the dynamics of the sounds that relatetheir tempo to our own experience of time. As a lover of

music, I completely agree with Hanslicks view but, as a

neurophysiologist, I find it desperately difficult to sub-stantiate this feeling using neurophysiological tools.

Nevertheless, I will try to discuss some data that may be

relevant to our perception of music from an aesthetic per-spective, although I am aware that they will be only indi-

rectly related to the core of the music. In doing this, the

main risk is to give Western tonal music (the subject ofmost studies of the aesthetic aspects of music) a universal

value. This is obviously not true because the tonal system

is by no means the universal paradigm of music even inWestern cultures, and has attained its dominant position

only over the last three centuries or so. What we can learn

from these studies may therefore not apply to other musicalsystems, including those used in Western music before the

introduction of the so-called baroque style or in the

twentieth century.One aspect that qualifies a human activity as artistic is

creativity. The difference between a common figurativeworker and an artist was happily expressed by Pablo

Picasso, who once said that there are painters who trans-

form the sun to a yellow spot, but there are others who,with the help of their art and their intelligence, transform a

yellow spot into the sun. Artistic creativity is what sets the

boundary between an ordinary and an artistic musicalproduct, and can be defined as an ability to establish novel

melodic, harmonic, rhythmic and timbre relationships

between the notes.There is no doubt that humans can perceive the signa-

ture of a creative act in a musical composition but the

question is: can such a perception be objectively describedusing a neurobiological approach? Most experiments

involving the brain and music have been carried out using

rather trivial musical material that is devoid of any creativevalue, and therefore does not seem to be adequate to pro-

vide an answer to the question. However, the responses to

melodic and harmonic irregularities (MMN and ERAN)may provide an interesting means of investigation. In the

literature, the alterations that evoke these responses are

negatively seen as violations, but Stefan Koelsch (seeKoelsch 2009) has shown that ERAN can be elicited by

Neapolitan chords even in a regular dominant to tonic

harmonic progression. The Neapolitan chord characterisingthe Neapolitan school has been widely used by non-

Neapolitan composers, and is not perceived as a viola-

tion of musical rules, although it may be unexpected to alistener of a dominant to tonic progression.

It can therefore be realistically hypothesised that the

perception of a musical passage that sounds unexpected tothe listener because of its originality could be revealed by

recording brain responses, which would provide research-

ers with an objective means of studying the effect ofmusical creativity.

6.1 Music and emotions

The idea that music expresses emotions is a commonplace

that is often assumed to explain the essence of the art ofmusic. However, it is important to say that intensity of

emotional feeling is not always a measure of the artistic

quality of the music that elicits it. Leaving aside theemotional responses aroused by a piece of music simply


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because of its associations with a certain memory (as is

typical in the case of tunes related to a privileged love

experience), there are other instances in which music-associated emotion is not related to the musics artistic

quality, such as the sound of an organ in a church, military

marches, anthems, etc., to which the emotional response oflisteners may be very intense even if the artistic quality of

the music is desperately poor. I will consider here only the

emotions that seem to be primarily related to the intrinsicfeatures of music.

Rigorous studies of the effects of musical tunes chosen

to induce pleasant or unpleasant subjective responses havebeen carried out by Blood et al. (1999), Koelsch (2005),

Koelsch et al. (2006) using PET, fMRI and EEG record-

ings). Pleasant emotions were induced by joyful andmainly consonant musical tunes, whereas musical pieces

with various degrees of dissonance were used to induce

unpleasant emotions. The main changes induced byunpleasant music were observed within an extensive neu-

ronal network of limbic and paralimbic structures consist-ing of the amygdala, hippocampus, parahippocampal

gyrus, and temporal poles. During the presentation of

pleasant music, activations were observed in the ventral

striatum and the anterior superior insula. Additional acti-vations in response to consonant (in comparison with dis-

sonant) music were observed in the larynx representation

within the Rolandic operculum (Broadmanns area 43) andthe inferior frontal gyrus (Broadmanns areas 44i, 45/46).

Using a slightly different approach, Blood and Zatorre

(2001) tested the effects of musical stimuli capable ofeliciting intensely pleasurable responses in a group of

subjects who described them as shivers-down-the-spine or

chills.Such chills were found to be highly reproducible in the

same individual by a specific type of music, thus providing

the investigators with the possibility of studying emotion-related changes in regional cerebral blood flow by means of

a PET scan and MRI. The results demonstrated the acti-

vation of a circuit that included the ventral striatum, thedorsomedial midbrain, the amygdala and hippocampus that

correlated with the intensity of the chills (Fig. 7). Thesestructures form part of a reward/motivation circuit that is

Fig. 7 Brain activations associated with music-related chills. Regres-sion analyses were used to correlate regional blood flow (measured byPET) with ratings of chills intensity (010). Neuroanatomical regionswith positive (left column) and negative (right column) correlations,are shown as t-statistic images superimposed on correspondingaverage MRI scans (x, y coordinates refer to location in stereotaxicspace. a (sagittal section, x = 4 mm) shows positive rCBF correla-tions in left dorsomedial midbrain (Mb), right thalamus (Th), AC,

SMA, and bilateral cerebellum (Cb). b (coronal section, y = 13 mm)shows left ventral striatum (VStr) and bilateral insula (In; also AC).c (coronal section, y = 32 mm) shows right orbitofrontal cortex (Of).d (sagittal section, x = 4 mm) shows negative rCBF correlations inVMPF and visual cortex (VC). e (sagittal section, x = 21 mm) showsright amygdala (Am). f (sagittal section, x = -19 mm) shows lefthippocampus/amygdala (H/Am) (from Blood and Zatorre 2001)


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known to be activated during pleasurable emotions, such as

those related to nutrition, sexual activities or drugs ofabuse.

The idea that music-related pleasurable emotion is asso-

ciated with the activation of the same structures as thoseinvolved in the emotional colour of such biologically

important activities as nutrition and reproduction is certainly

interesting. These findings may open up the way towards afurther understanding of the brain processing associated with

the arousal of aesthetic emotion that invades us when lis-tening to or playing a piece of music we love.

Acknowledgments The generous support of the Mariani Founda-tion to the organization of the series of meetings The Neurosciencesand Music is gratefully acknowledged.

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Neuroscience and musicAbstractIntroductionMusic and the brainMethodsNeuropsychologyNeurophysiologyNeuroimaging

Anatomo-functional organisation of the musical competences of the brainMusic and languageNeuroscience and music as an aesthetic experienceMusic and emotions

AcknowledgmentsReferences

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