The effect ofmusical experience on emotional self-reports

and psychophysiological responses to dissonance

DELPHINE DELLACHERIE,a,b MATHIEU ROY,c LAURENT HUGUEVILLE,d,e,f

ISABELLE PERETZ,c and SÉVERINE SAMSONa,baLaboratoire deNeurosciences Fonctionnelles et Pathologies, CNRS-UMR8160, University of Lille-Nord de France, Lille, FrancebEpilepsy Unit, La Salpêtrière Hospital, Paris, FrancecBRAMS Laboratory, University of Montréal, Montréal, Québec, CanadadUniversité Pierre et Marie Curie-Paris 6, Centre de Recherche de l’Institut du Cerveau et de la Moelle Epinière, UMR-S975, Paris, FranceeInserm, U975, Paris, FrancefCNRS, UMR 7225, Paris, France

Abstract

To study the influence of musical education on emotional reactions to dissonance, we examined self-reports and

physiological responses to dissonant and consonant musical excerpts in listeners with low (LE: n5 15) and high (HE:

n5 13) musical experience. The results show that dissonance induces more unpleasant feelings and stronger phys-

iological responses in HE than in LE participants, suggesting thatmusical education reinforces aversion to dissonance.

Skin conductance (SCR) and electromyographic (EMG) signals were analyzed according to a defense cascade model,

which takes into account two successive time windows corresponding to orienting and defense responses. These

analyses suggest that musical experience can influence the defense response to dissonance and demonstrate a powerful

role of musical experience not only in autonomic but also in expressive responses to music.

Descriptors: Emotion, Music, Dissonance, Musical experience, Psychophysiology, SCR, EMG, HR

Since Helmholtz (1877), musical dissonance has been considered

by psychologists and neuroscientists as a puzzling topic at the

interface between auditory perception and emotional processing.

The simplest definition of dissonance refers to an unpleasant

sensation induced by the simultaneous presentation of two

sounds. As proposed by Pythagoras two and half millennia ago,

this affective sensation produced by a pair of tones is related to

the ratio of the fundamental frequencies of the sounds that are

played together. When the ratio is simple, consonance is heard

(e.g., octave 2/1; perfect fifth 3/2) and when the ratio is more

complex, dissonance occurs (e.g., tritone chord, 45/32) (Plomp&

Levelt, 1965; Tramo, Cariani, Delgutte, & Braida, 2001). This

physical component of dissonance results in a perception of

beating that is perceived as unpleasant and is referred to as

‘‘sensory’’ dissonance, which appears when chords are played in

isolation (Tramo et al., 2001). This phenomenon should be dis-

tinguished from ‘‘musical’’ dissonance, which refers to disso-

nance manipulated by composers in a harmonic or melodic

context in order to induce tension and expectation effects

(Meyer, 1956). In the present study, we did not consider musi-

cal dissonance but focused rather on sensory dissonance.

According to neuropsychological, electrophysiological, and

brain imaging studies, it has been suggested that perceptual and

emotional processing of dissonance depends on distinct cerebral

structures. Whereas perception of dissonance requires the func-

tion of auditory cortical areas within the superior temporal gyrus

(Fishman, Volkov, Noh, Garell, Bakken, et al., 2001; Peretz,

Blood, Penhune, & Zatorre, 2001), its emotional processing in-

volves various brain structures including mesial temporal lobe

and precuneus regions (Blood, Zatorre, Bermudez, & Evans,

1999; Gosselin, Samson, Adolphs, Noulhiane, Roy, et al., 2006;

Koelsch, Fritz, Von Cramon, Muller, & Friederici, 2006).

At the behavioral level, it is well known that dissonant chords

induce more negative valence judgments than consonant chords

(Blood et al., 1999; Brattico, Pallesen, Varyagina, Bailey, Anour-

ova, et al., 2009; Gosselin et al., 2006; Khalfa, Roy, Rainville,

Dalla Bella, & Peretz, 2008; Koelsch et al., 2006; Koelsch, Remp-

pis, Sammler, Jentschke,Mietchen, et al., 2007; Pallesen, Brattico,

Bailey, Korvenoja, Koivisto, et al., 2005; Passynkova, Neubauer,

& Scheich, 2007; Peretz et al., 2001; Sammler, Grigutsch, Fritz, &

Koelsch, 2007; Schoen, Regnault, Ystad, & Besson, 2005). Even

infants prefer consonance to dissonance, suggesting a natural

The authors are grateful to Laurence Conty, Daniela Sammler, Séve-

rine Farley, and SeanHutchins for their helpful assistance and comments

on previous versions of the manuscript. This study was supported by a

PhD scholarship from the Regional Council of Nord-Pas de Calais to

Delphine Dellacherie and by a grant from ‘‘Agence Nationale pour la

Recherche’’ of the French Ministry of Research (project no. NT05-3-

45987) to Séverine Samson and from Eisai Inc. Isabelle Peretz is

supported by grants from the Canada Research Chair in Neurocognition

of Music and Mathieu Roy by a Canadian fellowship from NSERC.Address correspondence to: Séverine Samson, Department of Psy-

chology, Université de Lille 3, BP 60 149, 59653 Villeneuve d’AscqCedex, France. E-mail: severine.samson@univ-lille3.fr

Psychophysiology, ]]] (2010), 1–13. Wiley Periodicals, Inc. Printed in the USA.Copyright r 2010 Society for Psychophysiological ResearchDOI: 10.1111/j.1469-8986.2010.01075.x

1

mailto:severine.samson@univ-lille3.fr

aversion to dissonance very early in life (Masataka, 2006; Trainor

& Heinmiller, 1998; Trainor, Tsang, & Cheung, 2002; Zentner &

Kagan, 1996). However, it has also been suggested that emotional

responses to dissonance can be influenced by musical experience.

As reported by Pallesen et al. (2005) and Schoen et al. (2005),

adults with musical experience considered dissonant chords as

more unpleasant than did musically untrained individuals. These

self-reported results suggest that aversion to dissonance could in

some instances be reinforced by formal musical training and pos-

sibly by learning mechanisms. Further lines of evidence suggest

that, at the neurophysiological level, musical experience could fa-

cilitate the processing of dissonance in musicians in comparison to

nonmusicians (Brattico et al., 2009; Minati, D’Incerti, Pietrocini,

Valentini, Scaioli, et al., 2009; Regnault, Bigand, & Besson, 2001;

Schoen et al., 2005). However, the effect of musical education on

emotional responses to dissonance is not a well-established phe-

nomenon (Brattico et al., 2009).

To test the effect of musical experience in emotional responses

to dissonance, we recorded psychophysiological correlates of

musical listening, which is an appropriate method to examine

objectivemeasures of emotional experiences (Bradley, Codispoti,

Cuthbert, & Lang, 2001; Bradley & Lang, 2000; Lang, Bradley,

& Cuthbert, 1998; Sanchez-Navarro, Martinez-Selva, & Ro-

man, 2006; Witvliet & Vrana, 1995). Several lines of evidence

show that physiological changes in skin conductance responses

(SCRs), heart rate (HR) and electromyographic (EMG) re-

sponses vary systematically with judgments of affective valence

or arousal to emotional stimuli, although evidence in the musical

domain is more mixed.

In general, SCR and HR responses that are indexes of au-

tonomic activity are shown to be higher for negative than for

positive emotion (Cacioppo, Bernston, Klein, & Poehlmann,

1997; Cacioppo, Bernston, Larsen, Poehlmann, & Ito, 2000;

Taylor, 1991). For example, by comparing the results obtained in

13 physiological studies, Cacioppo et al. (2000) found that heart

rate was significantly greater during negative than positive emo-

tions. The authors interpreted this effect as part of a ‘‘negativity

bias,’’ which was defined in behavioral studies as ‘‘the greater

impact of negative information on people’s evaluations than

comparably extreme positive information’’ (Peeters &Czapinski,

1990). Classical animal and behavioral studies (for review, see

Cacioppo et al., 2000; Taylor, 1991) as well as more recent phys-

iological studies (Ito, Larsen, Smith, & Cacioppo, 1998) inves-

tigated the concept of ‘‘negativity bias,’’ which can be considered

as a normalmanifestation of adaptive functioning. Cacioppo and

colleagues (Cacioppo & Bernston, 1994; Cacioppo, Gardner, &

Berntson, 1997) have then incorporated the negativity bias in a

more general model of evaluative space in which positive and

negative evaluative processes are assumed to involve separate

motivational substrates (Lang et al., 1998). In this context, neg-

ativity bias refers to a tendency for the negative motivational

system to respond more intensely than the positive motivational

system to comparable amounts of activation (Ito, Cacioppo, &

Lang, 1998; Ito et al., 1998). This is the definition that we

adopted in the present study, and we hypothesized that this neg-

ativity bias exists also with music.

In the musical domain, several studies have focused on the

effect of various musical variables on autonomic responses to

emotional stimuli (Baumgartner, Esslen, & Jancke, 2006;

Chapados & Levitin, 2008; Grewe, Nage, Kopiez, & Altenmull-

er, 2007; Khalfa, Peretz, Blondin, & Robert, 2002; Khalfa, Roy,

et al., 2008; Koelsch, Kilches, Steinbeis, & Schelinski, 2008;

Krumhansl, 1997; Steinbeis, Koelsch, & Sloboda, 2006; Witvliet

& Vrana, 2007), but only three studies specifically explored the

effect of musical valence on autonomic responses (Nater,

Abbruzzese, Krebs, & Ehlert, 2006; Roy, Mailhot, Gosselin,

Paquette, & Peretz, 2008; Sammler et al., 2007). A greater HR

deceleration in response to aversive (heavy metal) music was re-

ported by Nater et al. (2006). Sammler et al. (2007) also found a

significant decrease of HR induced by dissonant music in com-

parison to consonant music. This cardiac deceleration does not

fit with the classic cardiac defense response obtained with aver-

sive loud noises, which is mainly characterized by a late accel-

eration (Cook & Turpin, 1997). However, when unpleasant

pictures or sounds are presented, a marked cardiac deceleration

has been repeatedly reported (Bradley et al., 2001; Cook & Tur-

pin, 1997; Sanchez-Navarro, Martinez-Selva, & Roman, 2006).

Such a cardiac deceleration is characteristic of an orienting re-

sponse (Bradley et al., 2001; Cook & Turpin, 1997). Following

Cacioppo et al. (1997, 2000), this greater response to unpleasant

than to pleasant affective stimuli could be interpreted as resulting

from the negativity bias. According to these results, we could

predict a higher HR deceleration in response to dissonant stimuli

than to consonant stimuli.

Although SCR has been traditionally linked to variation of

arousal (Bradley et al., 2001; Bradley & Lang, 2000), some ev-

idence suggests that SCR duration and amplitude can also be

influenced by emotional valence (Cacioppo et al., 1997, 2000;

Norris, Larsen, & Cacioppo, 2007; Ohman & Wiens, 2003). In

the musical domain, Baumgartner et al. (2006) and Nater et al.

(2006) found that SCR was more elevated for unpleasant or

negatively valenced music than for pleasant or positively va-

lenced music. Therefore, SCR could be larger for dissonant than

for consonant music.

According to the defense cascade model (Bradley et al., 2001;

Lang, Bradley, & Cuthbert, 1997; Ohman &Wiens, 2003), emo-

tional autonomic responses to aversive stimuli display a two-step

dynamic characterized by an initial orienting response followed

by a subsequent defense response. This model describes an aver-

sive motivational circuit that triggers reactions ranging from

orienting to fight/flight. The orienting response elicits a large

SCR response as well as a HR deceleration in an early time

window and is part of an automatic lower level appraisal that

could be processed in a short period of time, without any ap-

parent implication of cortical association areas or of awareness

(Ohman & Soares, 1994; Ohman & Wiens, 2003). The defense

response defined as a controlled appraisal of the emotion is re-

flected by further increase of SCR and HR acceleration in a

subsequent time window. In addition to these two steps of re-

sponse, HR responses to emotional stimuli typically show a third

decelerative component that corresponds to the return to base-

line. Because this third component is not part of the affective

response per se, we focused here on the two first components of

the HR response. Here, we hypothesized that negative emotions

created by musical dissonance should elicit a similar pattern with

two steps of responses (an orienting response, followed by a

defense response).

EMG responses of facial expression provide another index of

somatic activity related to emotional valence (Cacioppo, Klein,

Bernston, & Hatfield, 1993). More specifically, corrugator ac-

tivity (used in frowning) is increased for unpleasant or negatively

valenced stimuli (pictures: Bradley et al., 2001; mental imagery:

Witvliet & Vrana, 1995; sounds: Bradley & Lang, 2000; films:

Ellis & Simons, 2005). Conversely, zygomatic activity increases

2 D. Dellacherie et al.

with pleasantness (Ellis & Simons, 2005; Lang et al., 1998;

Witvliet & Vrana, 1995). Although the corrugator activity in

response to emotional valence inmusic has been already reported

(Roy et al., 2008; Witvliet & Vrana, 2007), no study to our

knowledge has recorded corrugator and zygomatic EMGactivity

in response to dissonance. Based on previous results in non-

musical domains, we can hypothesize that corrugator muscles

known to be influenced by negative valence should be more ac-

tivated when listening to dissonance than to consonance. How-

ever, zygomatic muscles generally modulated by positive valence

should be more activated in response to consonance than to dis-

sonance. Since no study interpreted facial EMG responses in

relation to the defense cascade model, no specific prediction

about the dynamic changes of such responses was proposed.

Finally, it has also been demonstrated that automatic phys-

iological responses to aversive stimuli can be modulated by our

own experience with the material. Some evidence in a non-mu-

sical domain suggests that the relevance of the task for a given

subject could modulate SCRs. For example, parachutists exhibit

larger skin conductance orienting response to parachutist-rele-

vant words and pictures than do inexperienced controls (Epstein

& Fenz, 1962; Fenz & Epstein, 1962). Since musical experience

acquired through lessons or practice can influence the emotional

response to dissonance, it seems relevant to explore the effect of

musical experience on emotional reaction to dissonance by com-

bining subjective rating judgments with objective autonomic

(SCR and HR) and somatomotor (EMG) measurements in re-

sponse to musical listening.

For this purpose, consonant musical excerpts and their dis-

sonant counterparts of the same musical pieces matched for

arousal were presented to participants with low and highmusical

experience that were carefully controlled in terms of age, sex,

education, and mood. Subjective ratings of emotional valence

(according to pleasantness) as well as autonomic (i.e., HR, phasic

SCR) and somatomotor (i.e., facial EMG) indexes related to

valence judgments were simultaneously recorded. We predicted

that dissonance will induce more negative valence judgments

than consonance, and this effect would be larger in high than in

low musically experienced participants. For all the participants,

we also predicted higher SCR and larger HR deceleration in

response to dissonance as compared to consonance, these effects

being modified by musical experience as well. Based on the de-

fense cascade model with two-step responses, two successive

patterns of autonomic responses were expected as a function of

the time window, early and late time windows corresponding to

orienting and defense responses, respectively. Therefore, we can

suppose that SCR and HR can be modified by musical experi-

ence differently in early and late time window. We also predicted

that facial EMG would be modulated by emotional valence: co-

rrugator muscles known to be influenced by negative valence

should be more activated when listening to dissonance than to

consonance, whereas zygomatic muscles generally modulated by

positive valence should be more activated in response to conso-

nance than to dissonance.

Methods

Participants

Twenty-seven participants (13 men and 14 women), aged be-

tween 19 and 31 years (mean age5 29.41. SD � 8.53) took part

in this study. Based on their responses on a musical experience

questionnaire, participants were sorted into Low Experience

(LE, n5 15) and High Experience (HE, n5 12) groups. The

musical experience questionnaire included items related to their

music listening habits (listening subscale) as well as their level of

musical education and actual practice (practice subscale), in ac-

cordance with a multi-dimensional definition of musicianship

(Ehrle, 1998). All HE participants, except one who was an au-

todidact, received training in classical music, and none of them

reported listening to music genres which generally have a lot of

dissonance, such as contemporary music or free jazz. As dis-

played in Table 1, the mean scores on both subscales were higher

for HE (mean listening score5 6, SD5 1.65 and mean practice

score5 6.92, SD5 2.57) than LE (mean listening score5 3.87,SD5 1.25; mean practice score5 3.87, SD5 1.04) participants(t(25)5 3.829, po.005 for listening and t(25)5 9.16, po.001 forpractice). Finally, the two groups did not differ on variables

unrelated to musical experience, such as age (mean age for

LE5 31.2, (SD5 9.08); for HE5 27.2 (SD5 2.18);

t(25)5 0.94, n.s.), years of education (mean years of educationfor LE5 14.33 (SD5 3.24); for HE5 15.83 (SD5 2.25);t(25)5 0.92, n.s.), and gender (for LE: 9M, 6F; for HE: 4M,

8F; w2(1)5 1.90, n.s). The results remain the same when onlyparticipants kept in the SCR and HR analyses were included in

the analyses.

Mood Questionnaires

Twomood questionnaires were administered: the State and Trait

Anxiety Inventory (STAI; Spielberger, 1983) and the Profile of

Mood Scales (POMS;McNair et al., 1992). The STAI comprises

two subscales, one assessing the individual general level of anx-

iety (i.e., trait anxiety), which is presumed to be stable over time,

and the other one assessing the present level of anxiety (i.e., state

Psychophysiology of musical emotion 3

Table 1. Description of the Musical Experience of the Two Groups

of Participants with Low (LE) and High Experience (HE)

Lowexperience

(LE)

Highexperience

(HE)

(A) Scores on the musical experience questionnaireGlobal score 4.13 (1.25)n 12.91 (3.21)n

Listening subscale score 3.87 (1.24)n 6 (1.65)n

Practice subscale score 3.87 (1.03)n 6.92 (2.57)n

(B) % of the participants reportingListening HabitsListen music every day 80 91.60Listen music with attention 20 66.60Listen classical music 33 58.30Go to concert 46.60 75

PracticePractice actually 0 33Have received an institutional training 6.60 75Have received lessons during at least 3years

6.60 92

Self-educated 0 8.30(C) Details on practice for HE

Age at start of study – 8.54 (3.56)Duration of the practice – 8.41 (4.30)

Notes: (A) Mean scores on the musical experience questionnaire (stan-dard deviation in parentheses). (B) Percentage of participants reportingdifferent listening habits and musical practice details. (C) For HE par-ticipants, mean age at start of study andmean number of years ofmusicalpractice (standard deviation in parentheses).npo.05.

anxiety). The POMS is a questionnaire comprising 30 items, each

consisting of an emotional adjective for which subjects have to

rate to what extent it describes their current mood (0F‘not atall,’ 4F‘extremely’). Six emotional sub-scales can be derivedfrom the 30 items: Tension, Depression, Anger, Vigor, Fatigue,

and Confusion.

Stimuli

Tenmusical excerpts of classical music were taken from a set used

in previous studies (Gosselin et al., 2006; Khalfa, Guye, Peretz,

Chapon, Girard, et al., 2008; Peretz et al., 2001; Peretz, Gagnon,

& Bouchard, 1998). They were instrumental in that they were not

originally sung with lyrics. The excerpts were selected to evoke

happiness and were played in major mode (e.g., ‘‘Brindisi’’ from

Verdi’s ‘‘Traviata’’), with a fast tempo (the quarter note value

varies from 80 to 255, conventionally written by M.M. for Met-

ronome Marking). All stimuli were transcribed for piano, com-

puter-generated, and delivered with a piano timbre. Each excerpt

had duration of 7 s. The dissonant versions of each original

consonant excerpt were created by shifting the pitch of the tones

of the leading voice by one semitone either upward or downward,

leading to 20 dissonant versions of the 10 original consonant

excerpts. During the experiment, each consonant version was

presented twice to match the number of dissonant excerpts.

Experimental Procedure

The general time course of the experiment is depicted in Figure 1.

All participants started the experiment by filling out the musical

experience and the mood questionnaires. Then, the physiological

sensors were attached while the participants sat comfortably in a

quiet room. Before the experiment started, participants per-

formed a handgrip squeeze task in which they were asked 4 times

to squeeze repeatedly on a handgrip for 7 s. This control taskwas

performed in order to obtain an index of individual electrodermal

reactivity in each participant before the beginning of the exper-

iment. The experimental task comprised four testing blocks.

Each block consisted of 10 excerpts, including 5 consonant

and 5 dissonant excerpts presented in an alternating order. The

orders of presentation of the consonant and dissonant trials

within each block were counterbalanced within subjects follow-

ing an ABBA design, as well as between subjects with half of the

participants starting with a consonant excerpt and the other half

with a dissonant one. All blocks started with a loud and sudden

burst of white noise (50 ms, 85–90 db) intended to elicit a startle

response. These startle responses served as another index of in-

dividual electrodermal reactivity and of habituation effect on the

signal throughout the experiment.

Each trial started with a warning signal asking the subject not

to move in order to avoid contamination of physiological re-

cordings by movement artifacts. This signal was followed by a

rest period lasting between 7.1 s and 7.7 s, in which subjects were

asked to do nothing. The length of the rest period varied ran-

domly in order to reduce subject’s expectancies and response

habituation. Themusical excerpt was then played for 7 s andwas

followed by another rest period of 1 s, in which subjects were

again asked not to move. This second rest period allowed the

recording of music-related physiological activity overflows and

also prevented the activity recorded at the end of the musical

excerpt from being contaminated by response preparation arti-

facts. The end of this 1-s rest period was indicated by another

signal allowing participants to move. Then, participants were

asked to judge the excerpt they just heard using a rating scale for

emotional valence (from � 55 ‘unpleasant’ to 55 ‘pleasant’)and arousal (from � 55 ‘relaxing’ to 55 ‘stimulating’). No timeconstraints were given for the ratings. All participants signed

informed consent to participate in the study.

Data Acquisition and Transformation

SCR, HR, and EMGs were monitored continuously using an

MP150 Biopac system (Biopac Systems, Inc., Goleta, CA) at a

sampling rate of 2000 Hz and processed using AcqKnowledge

4 D. Dellacherie et al.

A musicalmusicalexpertence

+ handgripexpert

+ handgrip+mood

handgripsqueeze Experimental Experimental Experimental Experimental

+mood

handgripsqueeze Experimental Experimental Experimental Experimental

questionnaires task block 1 block 2 block 3 block 4 questionnaires task block 1 block 2 block 3 block 4

B

llstartleprobe DCDDCDCCDCstartleprobe DCDCDCprobe

+DCDDCDCCDC

Musicalt i l

orprobe

+DCDCDC

Musicalt i l

orrecoveryperiod CDDCDCCDCD

trialsrecoveryperiod CDCDCD

trialsperiodperiod

10 sec10 sec ≈ 350 sec

C

“ l “valence

and“ l “valence

and“ l “valence

and“pleasedon’t rest

“youcan

andarousal

“pleasedon’t rest

“youcan

andarousal

“pleasedon’t rest

“youcan

andarousaldon t

move” rest period musical excerptrest

periodcan

move”arousalratings

don tmove” rest period musical excerpt

restperiod

canmove”

arousalratings

don tmove” rest period musical excerpt

canmove”

arousalratings

2 sec 7 1 to 7 7 sec 7 sec 1 sec 1 5 sec7 1 to 7 7 sec 7 sec 1 sec 1 5 sec7 1 to 7 7 sec 7 sec 1 sec 1 5 sec ≈ 10 sec2 sec 7.1 to 7.7 sec 7 sec 1 sec 1.5 sec7.1 to 7.7 sec 7 sec 1 sec 1.5 sec7.1 to 7.7 sec 7 sec 1 sec 1.5 sec 10 sec

Figure 1. Experimental procedure: (A) Participants first filled out the musical experience andmood questionnaires and performed the handgrip squeeze

task. The musical excerpts were then presented in four experimental blocks. (B) Each block started with a startle probe. Five consonant and 5 dissonant

excerpts were presented in an alternate order. The blocks either started with a consonant ‘‘C’’ or dissonant ‘‘D’’ excerpt. (C) Each musical trial started

with a rest period in which participants were asked not to move. At the end of each musical excerpt, participants were asked to remain still for 1 s before

they had to rate the valence and arousal of the excerpt.

software. Skin conductance was recorded on the index and mid-

dle finger of the non-dominant hand using the EDA finger

transducer BSL – SS3LA filled with an isotonic conducting gel.

The signal was filtered between 0.05 and 10 Hz. Then, the SCRs

to each musical trial were visually inspected and checked for

failures of the measuring device as well as movement-related ar-

tifacts. Trials in which there was electrodermal activity prior to

the onset of the musical excerpts were excluded. Responses were

selected if they occurred in a 1–4-s latency window following

stimulus onset. Three participants from the low expertise group

that showed no or very feeble responses to grasping and startle

trials were considered as non-responders and excluded from the

analysis.

Facial EMGs were recorded over the left corrugator and

zygomatic sites as recommended by Fridlund and Cacioppo

(1986), using 8 mm Ag/AgCl shielded electrodes, and were fil-

tered between 100 and 500 Hz. After the recording, EMG was

rectified using the root mean square function of the software and

then smoothed by a factor of 200 samples. HR was recorded

using a bipolar montage with an electrode on the right carotid

artery and another below the left ribs. The signal was filtered (0.5

to 35 Hz). Instantaneous inter-beat (RR) intervals (in ms), cor-

responding to the inverse of HR, were calculated from the elec-

trocardiogram using a peak detection algorithm to detect

successive R-waves and obtain a continuous RR tachogram.

Careful examination of the electrocardiogram and the tacho-

gram ensured that the automatic R-wave detection procedure

had been performed correctly. Careful examination of each par-

ticipant’s electrocardiogram signal led to the exclusion of four

participants (3 LE and 1 HE) due to technical failures. Electro-

cardiograms of the remaining participants were then cleaned by

removing the trials in which the presence of artifacts prevented

the exact extraction of RR Intervals (RRI). The participants who

were excluded from the SCR analyses were not the same ones

who were excluded from the HR analyses.

Results

Before analyzing the data, we checked for group differences be-

tween LE and HE participants on mood questionnaires. Then,

we tested the effects of dissonance and musical experience on

valence and arousal ratings, as well as on the physiological re-

sponses to the musical excerpts. Finally, we used regression an-

alyses to explore the relationship between valence ratings and

personal (mood and musical experience) and physiological

(SCR, facial EMGs, HR) variables. For all analyses, the Fmaxstatistic was used to test if the homogeneity of variance assump-

tion was met. Following the guidelines of Tabachnick and Fidell

(2001) for similar samples sizes (i.e., within a ratio of 4 to 1), the

Fmax was in an acceptable range (i.e., below 10) for all analyses,

indicating that the variances between samples were sufficiently

homogeneous to proceed with the analyses. Partial eta-squared

(Z2) were used as the effect sizes for the analyses of variance(ANOVAs). According to Cohen’s (1988) guidelines, Z2 5 .01corresponds to a small effect, Z2 5 .09 to a medium effect, andZ2 5 .25 to a large effect.

Mood Questionnaires

The mean ratings of the mood questionnaires STAI and POMS

for the LE andHEgroups are displayed in Table 2, alongwith the

results of the t-tests to verify whether baseline mood levels dif-

fered between the LE andHEgroups. As can be seen in this table,

there were no differences on any of the mood parameters that

were assessed, confirming that both groups did not differ in this

regard.

Valence and Arousal Ratings

Separate ANOVAs with one repeated measure (consonance vs.

dissonance) were carried out on the valence and the arousal rat-

ing scores. Figure 2 shows the mean ratings of valence and

arousal for consonant and dissonant excerpts as a function of

musical experience. As expected, dissonant excerpts were rated as

more unpleasant than consonant ones (F(1,25)5 73.19, po.05,Z2 5 0.75). This effect was modulated by the degree of musicalexperience of the listener, as revealed by the significant Musical

experience by Dissonance interaction (F(1,25)5 8.32, po.05,Z2 5 0.25). HE participants rated dissonant excerpts as moreunpleasant than LE participants did (F(1,25)5 8.81, po.05,Z2 5 0.26), whereas their ratings of consonant excerpts did notdiffer across groups (F(1,25)5 0.02, p5 n.s., Z2 5 0.001). Theanalysis of arousal ratings showed no effect of Dissonance

(F(1,25)5 0.28, p5 n.s., Z2 5 0.015) or of Musical experience(F(1,25)5 0.97, p5 n.s., Z2 5 0.51) nor any interaction(F(1,25)5 0.08, p5 n.s., Z2 5 0.003).

Psychophysiology of musical emotion 5

Table 2. Inter-Group Comparisons of the Mean (� SD) Ratings on the Mood Questionnaires and Electrodermal Reactivity Tests

Musical experience

Dependent variable Low experience High experience Result of t-test

STAI subscalesTrait anxiety 37.33 (� 8.86) 34.83 (� 7.63) t(25)5 1.51, p5n.s.State anxiety 34.40 (� 7.15) 30.58 (� 5.65) t(25)5 0.77, p5n.s.

POMS subscalesAnger 1.40 (� 1.45) 0.73 (� 1.10) t(25)5 1.28, p5n.s.Anxiety 1.73 (� 1.62) 1.55 (� 2.25) t(25)5 0.25, p5n.s.Depression 1.33 (� 1.80) 0.91 (� 1.64) t(25)5 0.62, p5n.s.Confusion 4.27 (� 1.87) 4.90 (� 0.83) t(25)5 1.06, p5n.s.Vigor 9.80 (� 2.68) 11.55 (� 3.59) t(25)5 0.42, p5n.s.Fatigue 3.13 (� 2.92) 4.18 (� 2.27) t(25)5 0.99, p5n.s.

Electrodermal reactivityHandgrip squeeze 0.45 (� 0.46) 0.48 (� 0.23) t(22)5 0.28, p5n.s.Startle 0.38 (� 0.34) 0.58 (� 0.40) t(22)5 0.86, p5n.s.

Note: n.s.5not significant.

Physiological Recordings

Electrodermal Responses

After exclusion of three non-responder participants, there

were 12 LE and 12 HE left in the SCR analyses. To make sure

that the LE and HE groups exhibited similar levels of electrode-

rmal reactivity, the mean SCRs for the grasping and startle trials

were compared by means of t-tests (Table 2). The results showed

that there were no differences in baseline electrodermal reactivity

between the two groups (Handgrip: t(22)5 0.28, p5 n.s.; Star-tle: t(22)5 0.86, p5 n.s.). The SCRs to the consonant and dis-sonantmusical excerpts were then analyzed for the two groups of

participants. First, the mean SCR values were extracted by slices

of 500 ms and averaged by musical condition and experimental

group in order to provide average response curves for the dis-

sonant and consonant excerpts and LE and HE participants (see

Figure 3A). Visual inspection of the resulting graph led to the

identification of two response periods according to the guidelines

proposed by Dawson (2007). The first response ranged from 1.5

to 6 s and corresponded to the orientation response triggered by

the onset of the excerpt. The second response ranged from 6 to 8

seconds and appeared to reflect later responses to the musical

excerpts. The amplitude of each SCR was then computed by

extracting the maximum value of the response in each time win-

dow and subtracting it from the baseline level. For the first time

window, the baseline period was the mean skin conductance level

of the 1-second baseline preceding the onset of the excerpt. For

the second time window, the mean skin conductance level be-

tween the 5th and 6th second served as the baseline. The resulting

values were transformed in log (SCR11) and averaged for each

participant.

An ANOVA with Dissonance (consonance vs. dissonance)

and Time window (early or later) as within-subjects factors were

carried out on the SCR values for the two groups of participants

(LE vs. HE). The averaged peaks for the LE and HE groups are

displayed in Table 3. SCRs were higher when participants lis-

tened to dissonant than to consonant excerpts (F(1,22)5 9.85,po.05, Z2 5 0.31), and this effect was influenced by musical ex-perience, as revealed by the significant interaction

(F(1,22)5 4.32, po.05, Z2 5 0.16) which demonstrates that thedifference in SCRs between consonance and dissonance was

higher for HE (F(1,11)5 7.89, po.05, Z2 5 0.42) than LE(F(1,11)5 2.05, p5 n.s., Z2 5 0.16) group.

The effect of Dissonance alsomarginally interactedwith Time

window (F(1,22)5 4.04, p5 .057, Z2 5 0.001), the effect of Dis-sonance being more pronounced in the early (F(1,22)5 9.01,

po.05, Z2 5 0.29) than in the late time window (F(1,22)5 1.63,p5 n.s., Z2 5 0.066). Finally, the interaction between Disso-nance, Time window, andMusical experience was not significant

6 D. Dellacherie et al.

Figure 2. Mean valence and arousal ratings of the dissonant and consonant musical excerpts for the low and high experience groups.

Table 3.MeanValues (� SD) of the Physological Recordings for Consonant andDissonant Excerpts byTimeWindow andLevel ofMusicalExperience

First window Second window

Consonant Dissonant Consonant Dissonant

SCR amplitude Low experience 0.08 (� 0.08) 0.10 (� 0.07) 0.01 (� 0.03) 0.01 (� 0.03)Log (maximum 11), mS High experience 0.12 (� 0.08) 0.20 (� 0.17) 0.04 (� 0.05) 0.07 (� 0.03)Corrugator EMG Low experience 0.03 (� 0.28) 0.06 (� 0.24) 0.03 (� 1.19) 0.94 (� 1.53)Area under the curve, mVnsec High experience 0.36 (� 0.90) 0.43 (� 0.65) 0.10 (� 01.17) 1.92 (� 4.53)Zygomatic EMG Low experience � 0.12 (� 0.22) � 0.03 (� 0.56) � 0.05 (� 1.43) 0.10 (� 2.51)Area under the curve, mVnsec High experience � 0.26 (� 0.42) � 0.11 (� 0.26) 0.26 (� 2.44) 3.03 (� 6.70)RR intervalIn the first window : Maximum acceleration, msec Low experience 33.73 (� 13.78) 29.50 (� 11.02) � 36.82 (� 28.55) � 35.92 (� 26.96)In the second window : Minimum deceleration, msec High experience 24.41 (� 12.64) 29.27 (� 11.44) � 32.06 (� 20.57) � 32.38 (� 14.66)

Note: All displayed values are differences from baseline.

(F(1,22)5 0.17, p5 n.s., Z2 5 0.075). For exploratory reasons,we tested if the interaction between Dissonance and Musical ex-

perience was different in the two time windows by performing

ANOVAs in the first and second time window with Dissonance

and Musical experience as factors. The results showed that the

effect of Dissonance did not interact with Musical experience in

the early time window (F(1,22)5 1.78, p5 n.s., Z2 5 0.075), butit did interact withMusical experience in the second time window

(F(1,22)5 4.11, po.05, Z2 5 0.16), indicating that SCRs werehigher for dissonant than for consonant excerpts in the late time

window. This effect holds for HE (t(11)5 2.39, po.05,Z2 5 0.25) but not for LE (t(11)5 1.91, po.05, Z2 5 0.05) par-ticipants.

Heart Rate

There were 12 LE and 11 HE participants left in the HR

analyses. RRI was extracted by slices of 500 ms and averaged by

musical condition and experimental group in order to provide

average response curves for the dissonant and consonant ex-

cerpts and LE and HE participants (see Figure 3D). Inspection

of the mean RRI curves revealed a typical triphasic pattern

(Bradley et al., 2001) comprising an initial increase of RRI (i.e.,

deceleration of heart rate), followed by a decrease of RRI (i.e.,

acceleration of heart rate), and then by a final increase of RRI.

The amplitude of the first increase and of the following decrease

were then extracted and subtracted from a 1-s baseline preceding

the onset of the musical excerpts. An ANOVA with Dissonance

and Phase of response was carried out on the RRI for the two

groups of participants. The amplitude of the first increase and

secondary decrease in RRI for the HE and LE groups are dis-

played in Table 3. No significant effect of Dissonance

(F(1,21)5 0.001, p5 n.s., Z2 5 0.00) or of Musical experience(F(1,21)5 0.01, p5 n.s., Z2 5 0.00) was observed. Only the ex-pected effect of Phase was observed (F(1,21)5 162.53, po.05,Z2 5 0.87), confirming that RRI were higher during the first in-crease than during the following decrease.

Facial Electromyography

The corrugator and zygomatic EMG responses to musical

excerpts were calculated as the difference between the raw signal

during each musical excerpts and the mean activity observed in

an 800-ms baseline preceding the onset of the excerpts. The re-

sulting response curves were then averaged by slices of 100ms for

the dissonant and consonant excerpts and for LE andHE groups

(see Figures 3B and 3C). Visual inspection of the resulting graphs

indicated that facial muscles appeared to show two phases of

Psychophysiology of musical emotion 7

ACorrugator EMG

BSkin Conductance Responses

0.6000.150

0.1252nd

0.50

2nd

window 21st0.100 0.40window 21

window0.100

0 301st window0.075

0 .30so

lts

μvo

0.0500.20

ns

me

nsi

eμ

0.100.025

00 1 2 3 4 5 6 7

0

1 2 3 4 5 6 7 8 9 −0.10−0.025

Time in seconds Time in seconds

DCZygomatic EMG R-R interval

101.20

8

2nd2nd window1.00 6 2 window2nd window

4

nds0.80

0

2

eco

n1st0.60

0lis

e1 2 3 4 5 6 7

window0.40 −2

mil

l

1 window1 2 3 4 5 6 7o

ltsvoμ

0.40

−4 1 window0.20

−8

−60

−101 2 3 4 5 6 7−0.20−12

Time in secondsTime in seconds

windowwindownd

Figure 3. Average time-courses of the physiological responses during the presentation of consonant and dissonant excerpts for low and high musical

experience groups. Each physiologicalmeasure was divided in two phases of responding. (A) For skin conductance responses, a first orientation response

(0–6 s) was followed by a second slower response (6–8 s). Note that the averaging of slight inter-trials and inter-subjects differences in the onsets of the

second non-orientation response smoothes the shape of themean response curves, compared to individual trials, fromwhich the analyzed responses were

extracted. (B) Corrugator EMG started to diverge between consonant and dissonant excerpts after the 2nd second. (C) Zygomatic EMG started to

diverge between consonant and dissonant excerpts after the 2nd second. (D) RR Intervals first increase between 0–3.5 s (slowing of heart rate) and

decrease between 3.5–7 s (speeding of heart rate).

responding: a first phase ranging from 0 to 2 s, where there were

no differences between the consonant and dissonant trials, and a

second phase from 2 to 7 s, where responses to consonant and

dissonant musical excerpts started to diverge. The area under the

curve was extracted within these two time windows and was

averaged for consonant and dissonant versions for each partic-

ipant. The averaged area under the curve for the LE and HE

groups are displayed in Table 3. An ANOVA with two repeated

measures, Dissonance and Time window, was carried out on the

averaged area under the curve for the LE and HE groups.

Corrugator activity differed between consonant and disso-

nant excerpts as a function of Time window, as revealed by the

significant interaction between Dissonance and Time window

(F(1,25)5 4.09, p5 .05, Z2 5 0.14). Decomposition of the inter-action revealed that corrugator activity was higher for dissonant

than for consonant excerpts in the second window

(F(1,25)5 4.06, p5 .05, Z2 5 0.14), but not in the first window(F(1,25)5 0.40, p5 n.s., Z2 5 0.02). Zygomatic activity differedas a function of Dissonance, Time window, and Musical expe-

rience, as revealed by the significant interaction between the three

factors (F(1,25)5 4.09, po.05, Z2 5 0.14). Decomposition ofthe interaction showed that there was no difference between

consonant and dissonant excerpts in the first time window

(F(1,25)5 0.01, p5 n.s., Z2 5 0.00) whereas zygomatic activitywas higher for dissonant than for consonant excerpts in the sec-

ond window for the HE (F(1,11)5 4.84, p5 .05, Z2 5 0.31), butnot for the LE group (F(1,14)5 0.01, p5 n.s., Z2 5 0.00).

Regression Analyses

Stepwise Regressions

In order to assess the relationship between personal variables,

valence ratings, and physiological responses, we performed two

stepwise regressions with the effects of dissonance on valence

ratings as the dependent variable and physiological responses

(SCR, facial EMGs, and RRI within all time windows) and per-

sonal variables (Musical experience and STAI and POMS sub-

scales) as the independent variables. The goal of these regressions

was to identify the physiological responses andpersonal variables

that best predicted the effects of dissonance on valence ratings.

For all the variables included in the analysis, the difference be-

tween the mean value for the dissonant and consonant excerpts

(dissonant–consonant) was used as an index of the effects of

dissonance on these variables. Because the regression models in-

cluded all the physiological measurements, we only kept the

participants for which SCR and HR data were available (9 LE

and 11 HE).

The results of the first stepwise regression revealed that

zygomatic activity in the second time window was the best pre-

dictor of the decrease in valence in reaction to dissonant excerpts

(r5 0.61, po.05). None of the other variables (SCRs, corrugatoractivity, RRI, and zygomatic activity in the first time window)

significantly improved the proportion of variance explained by

the model once the variance explained by the zygomatic activity

in the second time window was taken into consideration. The

results of the second stepwise regression showed that musical

experience was the best predictor of the decrease in valence in

reaction to dissonant excerpts (r5 0.53, po.05). Again, allother personal variables (STAI and POMS subscales) did not

significantly improve the proportion of variance explained by the

model. Thus, the participants who scored higher on the musical

experience questionnaire or who had the largest increases in

zygomatic activity in reaction to dissonant excerpts were the ones

showing the largest decreases in valence ratings in response to

dissonance. A final correlational analysis indicated that musical

experience was also correlated to the amount of zygomatic

activity in response to dissonance (r(26)5 0.59, po.05),suggesting that musical experience predicted zygomatic

responses to dissonance, which in turn strongly predicted the

decreases in valence ratings in response to dissonance.

Discussion

The purpose of our study was to determine the effect of musical

experience on emotional reaction to dissonance by recording

emotional self-reports as well as their psychophysiological corre-

lates. For this purpose, we compared the emotional responses of

two categories of listeners. One consisted of nonmusicians with

low musical experience (LE). The other one was composed of

musicians with high musical experience (HE) who had received at

least 3 years of formal training or who were still practicing a mu-

sical instrument although they did not reach professional levels.

Therefore, we were able to investigate the interaction between

musical experience and emotional pleasantness judgments on au-

tonomic and somatic responses by measuring SCR, HR, and fa-

cial EMG for the zygomatic and corrugatormuscles in response to

consonant and dissonant musical excerpts. Before we discuss the

results in further detail, we should first point out that the two

groups did not differ in terms of age, sex, education, or mood.

Moreover, given that the only difference between the stimuli con-

cerned the pitch shift of the melodic line, tempo differences could

not have contaminated physiological measurements, in contrast to

most previous studies onmusical emotionswhere differentmusical

excerpts were used to test different emotions.

Behavioral Measures

As predicted, the behavioral results confirm previous find-

ings, indicating that listening to dissonance induces a more neg-

ative affect than listening to consonance (Blood et al., 1999;

Gosselin et al., 2006; Koelsch et al., 2006; Pallesen et al., 2005;

Passynkova et al., 2007; Peretz et al., 2001; Sammler et al., 2007).

Arousal ratings, which were used as a proxy for motivational

activation (Ito, Larsen, et al., 1998) did not differ between con-

sonant and dissonant excerpts. In agreement with Pallesen et al.

(2005) and Schoen et al. (2005), we also found that the effect of

dissonance on emotional valence judgments ismore salient inHE

than in LE participants. Given that arousal judgments obtained

for consonant and dissonant stimuli did not differ between the

two groups of listeners, we can therefore suggest that musical

experience modulates valence judgments. Moreover, regression

analyses showed that musical experience was a very good pre-

dictor of valence ratings in the present study, suggesting that

musical experience may have enhanced the participants’ sensi-

tivity to dissonance. However, since almost all HE participants

were trained in classical music, it remains to be determined if the

observed results can be generalized to training in other musical

genres more marked by dissonance, such as free jazz or contem-

porary music. Nevertheless, this result appears to be consistent

with data of another study (Bigand, Parncutt, & Lerdahl, 1996)

showing that dissonant chords induce stronger ratings of musical

tension than consonant chords, this effect being more pro-

nounced in musicians than in nonmusicians. We can therefore

conclude that musical experience can modulate emotional judg-

ment of music even if the role of the specific background remains

to be clarified.

8 D. Dellacherie et al.

Physiological Responses

The analysis of physiological measures revealed that disso-

nance elicited stronger responses than consonance in the two

groups of participants. Given that dissonant excerpts were

judged by all participants as more unpleasant than consonant

excerpts but equally arousing, we interpret this reaction to dis-

sonance as resulting from the emotional negativity bias, defined

as the tendency for the negative motivational system to respond

more intensely than the positive motivational system to compa-

rable amounts of activation (Cacioppo et al., 2000; Taylor, 1991).

The main finding of our study confirmed that autonomic and

somatomotor responses to musical dissonance are influenced by

musical experience, with HE participants presenting stronger

physiological responses to dissonance in comparison to conso-

nance than LE participants. Since HE participants judge disso-

nance as even more unpleasant than consonance as compared to

LE participants, the differential reactivity to dissonance between

the two groups seems to be related to the subjective ratings of

emotional valence.

HR Measures

HR is normally responsive to the affective valence of the

stimuli. Emotional stimuli generally prompt a triphasic HR re-

sponse characterized by an initial brief deceleration, followed by

a short acceleration and a latemoderate deceleration (Lang et al.,

1997; Sanchez-Navarro et al., 2006). Although we found the

predicted triphasic response when listening to the musical ex-

cerpts, it was not modulated by dissonance. This result does not

seem to be consistent with previous findings obtained with un-

pleasant pictures (Bradley et al., 2001). However, these authors

found an effect of valence on HR response only with high (e.g.,

erotic pictures or mutilation scenes) but not with low arousal

pictures from International Affective Picture System (Lang,

Bradley, & Cuthbert, 1999). Similarly, Sammler et al. (2007)

found an effect of dissonance on HR deceleration with real but

not synthetic excerpts that might have been more arousing than

our computer-synthesized versions of the stimuli. It might be

therefore possible that the musical stimuli used in the present

study do not induce a sufficiently high emotional response to

affect HR. Notwithstanding the fact that we did not observe any

significant difference inHR responses between the two emotional

conditions, the successive phases of decelerative and accelerative

cardiac responses clearly argue in favor of the two-step defensive

cascade model (Bradley et al., 2001; Lang et al., 1997).

SCR

The analysis of phasic SCR revealed two different peaks of

response to emotional musical excerpts in the 2–6 s and in the

6–8 s windows. As the late response could not be ascribed to the

onset of the musical excerpt, this second bump indicates a sec-

ond, superimposed skin conductance response. Indeed, SCRs are

monophasic, and a secondary increase indicates that another

SCR has been triggered after the first one (Dawson, 2007). These

two successive responses match our hypothesis based on a two-

step cascade model and are consistent with the HR response

pattern (Bradley et al., 2001). The initial response, which occurs

immediately after the onset of the stimulus (early time window)

and the following response beginning a few seconds later (late

time window) are both larger for dissonant than for consonant

excerpts. Based on the cascade defense model (Bradley et al.,

2001; Lang et al., 1997; Ohman & Wiens, 2003), these two suc-

cessive responses could correspond to the (automatic) orienting

response in the first time window and the (more controlled) de-

fense response in the second one. Curiously, such a biphasic re-

sponse is rarely seen in most psychophysiological studies of

emotion, where visual stimuli are used to induce emotions. For

instance, in Bradley and Lang’s study (2000), although the affec-

tive pictures had a 6-s duration, no second phasic response after

the initial orienting response was observed. Classically, a ‘‘more

sustained’’ electrodermal response is observed as defense re-

sponse (Norris et al., 2007; Ohman & Wiens, 2003). One expla-

nation for this original finding could be that musical emotions

unfold in time (Grewe et al., 2007) whereas for pictures, all the

relevant emotional information is available immediately. Inter-

estingly, musical experience seems to have contributed to the

enhancement of this second, more controlled response. Indeed,

after verifying that baseline electrodermal reactivity did not differ

between the two groups of participants, we showed that the SCR

difference between dissonance and consonance is more pro-

nounced in HE than in LE group, as predicted. Subsequent SCR

analysis showed that musical experience interacted with disso-

nance in the second but not in the first time window, suggesting

that these orienting and defense responses were differently mod-

ulated by musical experience.

The orienting response is known to be processed at a pre-

attentive level and without implication of cortical association

areas (Ohman &Wiens, 2003). The orienting SCR to dissonance

that we observed in the present study could be interpreted as an

automatic response reflecting an emotional unconscious pro-

cessing. Given that this orienting SCR was higher for dissonance

than for consonance in HE and LE participants, we suggest that

nonmusicians can discriminate dissonance from consonance on

the basis of emotional feeling as musicians do. We were not able

to demonstrate an interaction between musical experience and

dissonance in this first time window, although larger SCRs were

recorded for HE than for LE participants. This finding indicates

that LE as well as HE participants showed stronger orienting

responses to dissonance than to consonance. We note that this

result was observed despite the lack of formal musical training in

LE participants.

At a more evaluative level (defense response in the second

time window), we also observed larger responses to dissonance

(unpleasant) than to consonance (pleasant) but only in HE par-

ticipants. This response seems to be enhanced in musicians. The

influence of musical valence on the SCR in the second time win-

dow is compatible with previous results demonstrated by re-

cording the tonic (and not phasic) SCRs when listening to

musical stimuli for more than 1 min (Baumgartner et al., 2006;

Nater et al., 2006). The fact that a sustained and higher response

was obtained for dissonance than for consonance suggests that

dissonance affects a more elaborate processing in addition to the

automatic response. This later processing of musical stimuli

could be specific to musicians. An interpretation of this result is

that formal musical learning led musicians to consciously reject

dissonant stimuli in a controlled defense response that follows the

pre-attentive orienting one. Taken together, analysis of SCR

suggests that musical experience could modulate musical disso-

nance processing, particularly in the late SCR. This result indi-

cates that musical education and training enhance emotional

response to dissonance.

An alternative interpretation of these results could be that

stronger SCRs to dissonance than to consonance may be related

to an effect of unfamiliarity of dissonance. It is clear that con-

sonant excerpts are more frequent or familiar than dissonant

Psychophysiology of musical emotion 9

ones because we are immersed in a consonant musical environ-

ment. Moreover, Terhard (1984) hypothesized that prenatal ex-

posure to the overtone structure of maternal speech could serve

as the basis for developing preferences for consonance. It is also

clear that musical experience contributes to the fact that some

people are more exposed to consonance than others are, even

more so if the formation is classical as it is in the present study. It

is therefore impossible to differentiate the pleasantness induced

by consonance from the frequency or exposure effects (Zajonc,

1980) and to dissociate the unpleasantness to dissonance from a

novelty or incongruity effect. However, the aim of the study was

not to explain the origin of aversion to dissonance, which is still a

hotly debated topic (Hauser &McDermott, 2003), but rather to

explore the relationships between physiological and self-re-

ported measures of emotional responses to dissonance (Bradley

et al., 2001; Ohman & Wiens, 2003). Because subjective emo-

tional ratings and physiological responses converge, the inter-

pretation of physiological responses as reflecting emotional

reactions to dissonance appears highly probable, even if this

emotion might be at least partly explained by the unfamiliarity

of dissonance.

Several lines of evidence in non-musical domains showed that

the relevance of the task for a given subject plays a role in mod-

ulating the SCR (Dindo & Fowles, 2008; Epstein & Fenz, 1962;

Fenz & Epstein, 1962; Lang et al., 1998; Ohman & Soares, 1994;

Perpina, Leonard, Bond, Bond, & Banos, 1998; Stormark, La-

berg, Nordby, & Hugdahl, 2000). These studies, carried out with

psychiatric patients and normal volunteers, revealed that higher

SCRs specific to a given object were dependent on individual ex-

perience. For example, studies examining phobic individuals

(Lang et al., 1998; Ohman & Soares, 1994) showed hyperactivity

to phobic objects, by expressing an intense and sustained fear in

response to stimuli thatwere not necessarily dangerous.According

to Mineka and Ohman (2002), the learning of fear could arise

froma classic Pavlovian conditioning: phobic patients would learn

to be frightened by a neutral stimulus because of an association

between such a stimulus and an aversive event. Because formal

musical education trains listeners to detect and reject dissonance

when it is not integrated in an appropriate musical context, we can

hypothesize by analogy that musical experience leads to a hyper-

sensitivity to dissonance. Consequently, HE participants could

produce larger SCRs to dissonance than do LE participants in the

same way as phobic subjects respond to phobia-relevant stimuli

(Lang et al., 1998). In other terms, even if aversion for dissonance

is natural (Schellenberg & Trainor, 1996; Zentner &Kagan, 1996)

or influenced by an exposure effect (Witvliet & Vrana, 2007), HE

participants could be seen as conditioned to react to dissonance,

resulting in an enhanced autonomic reactivity.

EMG Measures

Another index of the emotional appraisal of the stimuli was

the EMG responses. Corrugator and zygomatic muscles are

known to accompany unpleasant and pleasant emotions, re-

spectively. In the present study, we found a response of these

muscles that arose late (during the second time window), which

probably reflected a controlled mechanism. We can clearly see

that there is no zygomatic activity before 2 s after the onset of the

stimulus. Thus, the zygomatic response to musical excerpts is

particularly late. This cannot be explained by the characteristics

of EMG latencies. Indeed, unlike skin conductance responses,

which generally take around 2 s to develop because of sweat

gland physiology, EMG responses can be triggered within 100

ms. Therefore, this late response cannot be interpreted as an

orienting automatic response. Following the two-step cascade

model and given the coherent HR and SCR responses in regard

to this model, we thus propose to interpret a posteriori the late

EMG reactivity (more than 2 s) as reflecting more controlled

psychological processes related to conscious emotional evalua-

tion, which is characteristic of the defense response (Ohman &

Wiens, 2003). This interpretation is confirmed by the regression

analysis, which shows a strong relation between EMG responses

and valence ratings. Indeed, the zygomatic activity in the second

window was the best predictor of valence ratings.

Moreover, 2 s after the onset of the aversive stimulus, co-

rrugator activity was higher for dissonant than for consonant

excerpts, confirming that such a measure is relevant to the study

of emotional reaction to music (Roy et al., 2008; Witvliet &

Vrana, 2007). One probable reason why this facial response

seems to take more time to be initiated in the case of music

compared to pictures (Bradley et al., 2001) is the fact that the

musical emotions unfold in time whereas for pictures, all the

relevant emotional information is immediately available. More

studies on the dynamics of the response to musical emotion are

needed to clarify the specificity of emotional responses to music

in comparison to other types of stimuli, such as faces or brief

sounds.

In addition, we discovered that the late increase of zygomatic

contraction 2 s after the stimulus onset was specific to experienced

participants when they were listening to dissonant excerpts. This

unexpected result seems to be in contradiction with the responses

observed in other studies showing such a response with pleasant

stimuli (Ellis & Simons, 2005; Lang et al., 1998; Witvliet & Vrana,

1995). This zygomatic activity might be interpreted as an ironic

smile or as a grimace related to the displeasure induced by disso-

nance. Since we witnessed that some participants paradoxically

laughed (Ansfield, 2007; Craig & Patrick, 1985; Keltner & Bon-

anno, 1997; Papa&Bonanno, 2008; Prkachin&Solomon, 2008) at

the dissonant excerpts, we tend to favor the former interpretation,

although we lack objective empirical data to confirm it. Regression

analysis revealed that more unpleasant dissonances were associated

with stronger zygomatic activities and that this smile/grimace is the

best predictor of the subjective responses to dissonance. The

zygomatic response could therefore constitute not only a controlled

but also a communicative part of the emotional response to dis-

sonance, and this reaction could be more developed in HE partic-

ipants than in LE participants. Mimics to dissonance would have a

communicative function that musical learning might have contrib-

uted to create in HE participants. We could hypothesize that ex-

perience enhances the ability to communicate musically induced

negative emotions. Taken together, these results show that musical

experience plays a role in physiological and communicative re-

sponses induced by dissonance that are linked to emotional valence.

Based on our results, it seems that dissonance produces larger

SCR and EMG responses in HE than LE listeners. The results

could be further explained by brain reorganization induced by

musical experience. Indeed, even in amateur musicians or in

children, musical training can produce functional and morpho-

logical brain changes in auditory areas (Gaser & Schlaug, 2003;

Schneider, Scherg, Dosch, Specht, Gutschalk, & Rupp, 2002) as

well as in emotional structures such as the amygdala and the

insula (James, Britz, Vuilleumier, Hauert, & Michel, 2008), the

anterior cingulate cortex (Foss, Altschuler, & James, 2007) and

the frontal-lobe areas (Koelsch, Fritz, Schulze, Alsop, & Schl-

aug, 2005; Minati et al., 2009).

10 D. Dellacherie et al.

Taken together, based on the obtained physiological results,

we argue for the existence of a first automatic response to dis-

sonance, which is mainly characterized by an orienting SCR.

This initial physiological reaction seems to be followed by a sec-

ond phase, which is less automatic and more controlled. This

second step is characterized by facial EMG responses and a sec-

ond non-orienting SCR. The first response suggests the existence

of an alarm system that can automatically interpret dissonant

music as an aversive stimulus and then evaluate this stimulus in

terms of displeasure. In the nonmusical domain, an alarm system

in response to fear was shown to involve the amygdala (Liddell,

Brown, Kemp, Barton, Das, et al., 2005). Given that responses

of the autonomic nervous system are linked to the amygdala and

other structures such as the cingulate cortex, the effect of dis-

sonance on SCR is compatible with neuroimaging studies sug-

gesting that affective responses to dissonance could mainly

involve mesio-temporal lobe structures (Blood et al., 1999; Ko-

elsch et al., 2006, 2007). The stronger late responses obtained in

HE than in LE participants (second phase response in SCR and

zygomatic response) seem consistent with the more important

involvement of cortical structures such as the cingulate cortex or

frontal lobe areas in musicians than in nonmusicians in response

to dissonance and consonance (Foss et al., 2007; Minati et al.,

2009). Considering that the cingulate cortex is known to con-

tribute in regulating autonomic functions as well as in attentional

control (Critchley, Mathias, Josephs, O’Doherty, Zanini, et al.,

2003; Devinsky, Morrell, & Vogt, 1995), we may hypothesize

that the SCR and the activity observed with fMRI in the cingu-

late cortex in musicians may reflect a similar mechanism, result-

ing in a hypersensitivity to dissonance. This hyperactivation may

result in a stronger negativity bias observed in autonomic reac-

tivity in HE than in LE participants in the second phase of pro-

cessing dissonance.

Conclusion

The data of the present study emphasize the effect of musical

experience on emotional responses to music and the role of emo-

tional valence in determining the autonomic and somatomotor

components of these responses. The results indicate that SCR and

EMG responses are sensitive to dissonance. These measures

therefore appear to be appropriate tools to assess emotional re-

action to unpleasant music. Stronger responses to dissonant than

to consonantmusic were observed in the orienting SCR, and there

were no differences between HE and LE participants in the am-

plitude of this response, suggesting that the first step of emotional

response to dissonance could be independent of musical experi-

ence. Late skin conductance and EMG responses (corrugator and

zygomatic) confirmed the presence of a subsequent, more con-

trolled, response to dissonance. Moreover, whereas corrugator

activity when listening to dissonancewas found in all participants,

late SCRs and zygomatic activity were found only in experienced

musicians, suggesting a specific emotional response in this group.

This specific reaction expressed by stronger physiological late re-

sponses to dissonance in HE than in LE participants was con-

firmed by self-report responses. This suggests that experience

could influence the negativity bias. An interpretation is that mu-

sical education could have reinforced the representation of dis-

sonance for musicians by a long and sustained associative

learning between dissonance and unpleasant emotions. These re-

sults add arguments in favor of both physiological and psycho-

logical origins of the feeling of dissonance. Moreover, the smile

observed in musicians represents an original finding indicating

that musical experience influences not only conscious appraisal of

emotional significance but also its means of communication. In

other terms, learned aesthetic preference might play a powerful

role in affective and expressive response to music.

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(Received April 20, 2009; Accepted April 13, 2010)

Psychophysiology of musical emotion 13