Modulated neural processing of Western harmony in folk musicians

ELVIRA BRATTICO,a,b,c TIINA TUPALA,a ENRICO GLEREAN,b,c and MARI TERVANIEMIa,b

aCognitive Brain Research Unit, Institute of Behavioral Sciences, University of Helsinki, Helsinki, FinlandbCenter of Excellence in Interdisciplinary Music Research, University of Jyväskylä, Jyväskylä, FinlandcBrain and Mind Laboratory, Biomedical Engineering and Computational Science (BECS), Aalto University School of Science, Espoo, Finland

Abstract

A chord deviating from the conventions of Western tonal music elicits an early right anterior negativity (ERAN) ininferofrontal brain regions. Here, we tested whether the ERAN is modulated by expertise in more than one music culture,as typical of folk musicians. Finnish folk musicians and nonmusicians participated in electroencephalography record-ings. The cadences consisted of seven chords. In incongruous cadences, the third, fifth, or seventh chord was aNeapolitan. The ERAN to the Neapolitans was enhanced in folk musicians compared to nonmusicians. Folk musiciansshowed an enhanced P3a for the ending Neapolitan. The Neapolitan at the fifth position was perceived differently andelicited a late enhanced ERAN in folk musicians. Hence, expertise in more than one music culture seems to modify chordprocessing by enhancing the ERAN to ambivalent chords and the P3a to incongruous chords, and by altering theirperceptual attributes.

Descriptors: Cognition, Learning/memory, EEG/ERP, Individual differences, Music

In Western tonal music, the stylistic conventions of tonal harmony(Piston, 1962) constitute reference schemata, which provide guide-lines for building and listening to musical pieces. The rules ofWestern tonal music are based on subsets of twelve pitchesincluded in the chromatic equal-tempered scale, where the intervalbetween consecutive pitches is a semitone (100 cents; e.g.,Krumhansl, 2000). These subsets, namely, the diatonic scalesincluding only seven out of twelve pitches, display the hierarchicalrelations between sounds by means of the rules of tonality. Whenseveral sounds are played simultaneously as chords, harmonic rulesof the Western tonal system determine their permissible succession,voice leading, and the hierarchical importance of those chordswithin a sequence or cadence (e.g., the authentic cadence containsthe following chords: tonic-subdominant-dominant-tonic; Piston,1962).

In subjects possessing an implicit or explicit knowledge ofWestern tonal music, violations of its harmonic conventions elicit

an event-related potential (ERP) response called early right anteriornegativity (ERAN), which occurs at 150–250 ms after stimulusonset over anterior regions of the scalp (Koelsch, 2005, 2009;Koelsch, Gunter, Friederici, & Schröger, 2000; Koelsch & Mulder,2002). The ERAN is elicited by incongruous, unexpected chordsinserted in a musical sequence, such as by the Neapolitan subdomi-nant (Koelsch et al., 2000; Loui, Grent-‘t-Jong, Torpey, &Woldorff, 2005) or double dominant chords (Koelsch, Jentschke,Sammler, & Mietchen, 2007; Sammler, Koelsch, & Friederici,2011) when a tonic or a dominant chord is expected (GarzaVillarreal, Brattico, Leino, Ostergaard, & Vuust, 2011; Leino,Brattico, Tervaniemi, & Vuust, 2007). The latency and amplitudeof the ERAN sensitively reflect the degree of harmonic appropri-ateness of the chord (Garza Villarreal et al., 2011; Koelsch &Sammler, 2008). Even chord modulations (Koelsch, Gunter,Schröger, & Friederici, 2003) and violations of expected keypitches in melodic lines (Brattico, Tervaniemi, Näätänen, & Peretz,2006; Koelsch & Jentschke, 2010) elicit an ERAN-like response.Although in most ERAN investigations attention is diverted fromthe harmonically incongruous chords and focused on chords with adifferent timbre (e.g., organ or harpsichord; Koelsch et al., 2000),attentive listening to the music is required for elicitation of theERAN, especially when the incongruous chord is inserted withinthe cadence (Loui et al., 2005; Maidhof & Koelsch, 2011). Whenthe incongruous chord is salient and highly unexpected, such as aNeapolitan at the end of a chorale by Bach (Steinbeis, Koelsch, &Sloboda, 2006), the ERAN is followed at around 250–400 ms by apositive frontocentral deflection, namely the P3a (e.g., Koelschet al., 2007; Steinbeis et al., 2006). Originally, the P3a wasobserved in attentive stimulation paradigms including novel, thatis, rarely occurring nontarget sounds. Subsequently, it has beenidentified also with inattentive paradigms where the subject was

This study was financially supported by the BrainTuning project of theEuropean Commission (FP6-2004-NEST-PATH-028570) and the Academyof Finland (project number 133673). We thank Mr. Tommi Makkonen,M.Sci., for his technical help, and Benjamin Gold, B.Sc., and UmbertoTrivella, M.Sc., for their assistance in behavioral data collection. A specialthanks to Dr. Juha Ojala for his invaluable contribution to the musicologicalaspects of this paper. We also wish to express our gratitude to all subjectsand especially those who agreed to participate at a second behavioralsession. Parts of this study were presented at the conference “The Neuro-sciences and Music IV: Learning and Memory,” and a summary of theresults appeared in the conference proceedings published by the Annals ofthe New York Academy of Sciences.

Address correspondence to: Elvira Brattico, Cognitive Brain ResearchUnit, Institute of Behavioral Sciences, P. O. Box 9 (Siltavuorenpenger 1 B),00014 University of Helsinki, Helsinki, Finland. E-mail: elvira.brattico@helsinki.fi

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Psychophysiology, 50 (2013), 653–663. Wiley Periodicals, Inc. Printed in the USA.Copyright © 2013 Society for Psychophysiological ResearchDOI: 10.1111/psyp.12049

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asked to ignore the auditory stimulation, and even with infrequentbut non-novel changes in a stream of predictable sounds (Polich,2007). The P3a is hypothesized to index the recruitment ofinvoluntary attention (independently of the required behavioralresponse) towards a surprising, unexpected sound event followingthe automatic comparison between frequently presented soundsand the novel ones. P3a responses can hence be observed also withhighly unexpected sounds inserted in a musical context.

The neural representations of harmonic conventions are formedeven with only a few years of passive exposure to Western tonalmusic, as indexed by the ERAN elicitation to Neapolitan chords atthe end of an authentic cadence in 5- and 6-year-old childrenwithout formal music education (Koelsch et al., 2003). However,neural representations of harmonic rules strengthen with years ofexposure, and particularly with active exposure and training. Inmusicians, the amplitude of the ERAN is enhanced compared tolaypersons and can be elicited even when the violation of chordexpectation appears in a nonsalient position, such as in the middleof the cadence (Koelsch, Schmidt, & Kansok, 2002). Similarly,James, Britz, Vuilleumier, Hauert, and Michel (2008) found anERAN-like response peaking at around 200 ms from the onset of aslightly incongruous chord at the end of a musical piece only inexperts and not in laypersons. These findings indicate that musi-cians possess stronger representations of Western harmonic princi-ples of chord succession and react more strongly to their violations,including mild or nonsalient ones. Hence, they expect certainupcoming musical events more strongly than nonmusicians due toactive, decades-long exposure to Western tonal music and toexplicit knowledge of its underlying conventions.

The studies mentioned so far have focused on exposure toWestern tonal music, and musical expertise has been identified onlyas academic training in Western classical music even though it isrelevant for only a minority of the population. ERAN investigationshave also adopted mainly chord chorales composed during theBaroque period of classical music (e.g., by J. S. Bach; Koelsch et al.,2000, 2007; Koelsch & Jentschke, 2010) or actual expressive musicof the Romantic period (Koelsch & Mulder, 2002). Here, we wantedto test whether exposure to other music systems in addition to theWestern tonal one (omnipresent in the modern world) would modu-late the ERAN response to unexpected chords. To this aim, werecruited musicians trained in folk music. To be clear, the folk musicreferred to in this study is not the primarily NorthAmerican genre offolk music that has close ties to pop/rock music (e.g., WoodyGuthrie, Pete Seeger; Mitchell, 2007) but the aggregate of thetraditional, ethnic music of indigenous and nonindigenous peoplesworldwide. Our folk musicians were Finnish and trained in Finland.In the following, we will specify the content of local folk music.

Inherently, each music culture has its own aesthetics, contextsfor music, repertories, ways of transmitting music from generationto another, genres of music, and styles, including elements of pitchorganization such as scales used for melodies and possibly har-monic conventions (Slobin & Titon, 1992). Similar to folk music inmany other parts of the world, Finnish folk music has traditionallybeen aurally transmitted. It also makes use of improvisation andmelodic variations. It includes two major traditions. The older, lessWestern tradition exhibits genres such as the epic runo songs, thesinging of the Kalevalaic poems (in octosyllabic, trochaic tetram-eter, resulting in common practice meters such as 5/4, 4/4, or 9/4),which exhibits scales with three to five structural pitches: systemsof pitch organization that since the early collection of the tunes(beginning from the 18th century) have been compared to subsetsof major and minor modes of Western tonal music (Asplund &

Hako, 1981; Kolehmainen, 2007; Pekkilä, 1990). The other genreof the older Finnish folk tradition includes itkuvirsi laments, whichare improvisatory melodic, rhythmically complex recitations per-formed by (often hired specialist) women, expressing grief or otherpowerful emotions particularly in rituals of change (such as wed-dings or funerals). The pitch organization in laments mostly has abasic structure of three to five diatonic pitches, which might consistof the first pitches of the major mode or of the minor mode (butwith the neutral third, which is an interval smaller than major butlarger than the minor third). In this subgenre, microintervals, melis-mas, and ornamentation are used extensively (Asplund & Hako,1981; Kolehmainen, 2007; Pekkilä, 1990). Alternatively, theyounger pelimanni tradition exhibits genres such as the rekilaulusong and other unaccompanied songs, such as ballads, and a varietyof dance forms with regular meters originating from or influencedby Western tonal music. Although the transition to the newer tra-dition also includes a gradual transition to Western tonality, somemodal features persist, such as the occasional use of diatonic modesother than major or minor (e.g., the Dorian mode with a raised sixthcompared to the minor, or the lack of raised sixth and seventhdegrees in the minor mode, i.e., the use of natural minor). Thedance genres (e.g., polka, quadrille, waltz, etc.) incorporate newerinstruments (e.g., clarinet, harmonium instead of the box zitherkantele or the bowed lyre jouhikko), and use the diatonic scale andits major and minor modes but usually without the chromaticextensions thereof. Hence, not even in this newer tradition areNeapolitan chords encountered, although occasional secondarydominants may occur (Asplund & Hako, 1981; Kolehmainen,2007; Pekkilä, 1990). Nowadays, folk musicians are involved inwell-structured musical training offered by public music schoolsand academies, where they familiarize themselves with the above-introduced Finnish folk traditions and contemporary world music(Hill, 2009). Finnish folk music is an active field where the oldlocal traditions have been rejuvenated and often mixed with othergenres (Hill, 2007, 2009). Thus, Finnish folk musicians offer anexcellent possibility to investigate auditory neurocognition, whichin their special case is a result of the interplay between variousmusic systems from both the Western tonal tradition and othernon-Western music cultures.

By employing the paradigm developed by Leino et al. (2007)with Neapolitan chords replacing either a subdominant or a tonicchord in three different positions in the authentic cadence, wecompared harmonic processing in Finnish folk musicians andmusically untrained control participants. Contrary to most previousERP studies with musicians in which the music systems to whichthey were acculturated were not specified nor any behavioral meas-ures of their auditory skills or music cognition were obtained (e.g.,Brattico, Tervaniemi, & Näätänen, 2001; Brattico et al., 2009;Koelsch et al., 2002), here we carefully controlled for the musicalbackground and skills of the musically trained subjects with aquestionnaire, a musicality test, and a perceptual rating test. Ourhypothesis was that folk musicians would express the sensitivity tomusical chords typical of trained music experts due to their long-term daily musical practice and education in Western tonal musicalong with other music systems. In addition, we expected that folkmusicians would show a divergent neural and behavioral process-ing of Neapolitan chords, mildly or strongly violating the princi-ples of Western harmony depending on their role within thecadence. Particularly, Neapolitan chords substituting the subdomi-nant were hypothesized to be perceived as acceptable by nonmu-sicians but less so by Finnish folk musicians. The divergence inneural and behavioral processing of harmonic successions by folk

654 E. Brattico et al.

musicians would result from the interplay between neural sub-strates for Western harmonic system with those for the Finnish folkmusic, which does not include Neapolitan chords.

Method

Participants

For the ERP experiment, we measured 22 healthy, right-handedyoung adults with normal hearing aged between 20 and 35 years old.They included 11 Finnish folk musicians who were studying in amusic academy or university in Finland (4 males; M = 27.7 years)and 11 nonmusicians who had studied or practiced music for notmore than 2 years and had finished lessons over 10 years before theexperiment (2 males; M = 25.3 years). Nonmusicians were chosenas control subjects because of their perceptual competence inWestern tonal music, as a consequence of their passive daily expo-sure and musical education in the basic school system (cf. Margulis,2008). All folk musicians had completed at a music conservatorybasic exams of ear training and performance (level 3/3 in the Finnishsystem, equivalent to grade 5 of the Associated Board of the RoyalSchool of Music [ABRSM] British system). Music education inFinnish conservatories is centered on the classical genre andincludes traditional solfège, Western tonal harmonic theory, andpractice with an instrument mainly based on score reading. Hence,all our folk musicians had received basic formal education includingactive and passive exposure to Western classical music for at least 5years, plus informal learning, that is, exposure outside of formaleducation in Finnish daily life. After basic studies, our participantswent to study Finnish folk music. Two of our folk musicians hadobtained Master’s degrees in Folk Music from Sibelius Academy(Helsinki, Finland), two had Bachelor’s degrees in Music Pedagogywith specializations in folk music, and one had a Master’s degree inclassical music performance.

In the context of another ongoing study on neural auditorydiscrimination, we also measured the musical skills of folk musi-cians by administering the Advanced Measure of Musical Audition

(AMMA) test (Gordon, 1998). In AMMA, for about 15 min sub-jects have to decide whether two melodies played by a digitizedpiano are the same or different based on the discrimination of tonalor rhythmic changes. The raw scores for the tonal test section were31.9 � 3.4 SD, range 26–37 (scale 0–40). The raw scores for therhythm test section were 33.8 � 2.9 SD, range 29–37 (scale from0–40). These results testify that the musical skills of folk musicianscorresponded to those of (classical, jazz, or rock) professionalmusicians previously studied in the neuroimaging literature(Habermeyer et al., 2009; Schneider et al., 2002; Seppänen,Brattico, & Tervaniemi, 2007; Vuust, Brattico, Seppanen,Näätänen, & Tervaniemi, 2012). The instruments studied by thefolk musicians are illustrated in Table 1. Subjects’ participation inthe experiment was compensated with movie tickets.

Stimuli

Digitally produced piano chords were used to create the seven-chord cadences transposed over twelve keys (see Figure 1). Thethird, fifth, or seventh chord was a Neapolitan in 75% of the cases.Thus, standard cadences, in which there was no Neapolitan, madeup only 25% of all cadences. The frequency of the Neapolitans wasapproximately equal in all three positions. In the standard condi-tion, the chords formed an authentic cadence and there was noNeapolitan. The Neapolitan was used to replace the tonic chord atthe third position, the subdominant chord at the fifth position, or theending tonic chord at the seventh position. The substitution of thesubdominant chord at the fifth position with the Neapolitan createsthe least change, and perceptually the mildest violation of expec-tation. The Neapolitan is, namely, a chromatic variant of the sub-dominant of the minor mode. In fact, in a chromatic context of,especially, Western music of the Romantic era, the substitutionwould likely not be considered a violation at all. Instead, thesubstitution of the tonic chord at the third position yields a

Table 1. Main and Side Instruments of the Folk Musicians

Main instruments # Side instruments #

Violin 6 Piano 8Accordion with two rows 1 Vocals 6Folk wind instruments 1 Harmonium 5Guitar 1 Percussions 4Vocals 1 Mandolin 3Accordion 1 Guitar 3

Kantele 3Jouhikko 2Keyboards 2Nyckelharpa 2Accordion with two rows 2Estonian bagpipes 1Didgeridoo 1Viola 1Drums 1Saxophone 1Banjo 1Violin 1Bass guitar 1Bouzouki 1Double bass 1

Note. The number indicates how many of the folk musicians reported theinstrument as main or side instrument.

Figure 1. Experimental stimuli. Top: Musical notation of the chords usedfor the stimulus, only in the key of “C” for illustration purposes. Bottom:Illustration of the chord cadence. The left column shows the type ofcondition. The right columns show the positions of the cadence andthe key deviant chords at third, fifth, and seventh positions. T =tonic; S = subdominant; Sn = Neapolitan subdominant; D = dominant;T3 = inverted tonic.

Modulated processing of harmony in folk musicians 655

moderate violation in which the resolution of the preceding domi-nant chord, assumed necessary in the tradition of Western tonalmusic, is rendered unconventional. Finally, the substitution of thelast chord, another tonic chord preceded by a dominant chord, waspredicted to result in the strongest violation in terms of tonalexpectations, since the Neapolitan chord concludes the cadentialsequence after an unresolved dominant.

For the ERP experiment, the cadences also included acousti-cally deviating chords with an organ-like timbre. The subjects wereinstructed to press a button whenever they heard this deviant chord.These chords were included in about 8% of the cadences, and theywere similar to the other chords in all aspects except timbre. Thistask was created to ensure that the subjects were listening to thesounds but did not pay any special attention to the Neapolitanchords. These deviant chords were never presented at the third,fifth, or seventh position.

The durations and volumes of the sounds were equalized usingSound Forge 7.0 and Cool Edit 2000 software. The first six chordsof the cadence were 600 ms in duration and the last one was1,200 ms. The experimental sounds were faded out graduallyduring the last 50 ms, and there was a 5-ms silence between allchords. The gap between the cadences was kept to only 5 ms inorder to give the listener an experience of continuous music.

Procedure

The sounds were presented with Presentation software (v. 12, Neu-robehavioral Systems) at 50dB above individual hearing thresholdsas determined in the beginning of the experimental session. Thesounds were delivered with Sony MDR-7506 headphones while thesubject was sitting in a comfortable chair in an electronicallyshielded room. The complete duration of the ERP experiment was40 min, divided into three blocks lasting about 13 min each. Par-ticipants were instructed to look at a fixation cross. This ERPexperimental session always followed another one lasting 60 min,which included passive listening to musical patterns and watchingsilenced movies (data to be reported elsewhere).

Subsequently, in a separate session, we conducted a behavioralexperiment utilizing the same stimulation as the ERP study. For thisexperiment, 8 out of a total of 11 folk musicians and 4 out of 11nonmusicians from the ERP experiment agreed to participate. Toreach appropriate statistical power, we recruited additional newsubjects, reaching a total of 12 folk musicians (4 males; M = 29.7years) and 14 nonmusicians (4 males; M = 29 years). In the behav-

ioral experiment, subjects were asked to rate the cadences contain-ing the Neapolitan chords at the three different positions (henceexcluding the cadences containing the organ sounds). Eachcadence was repeated 12 times. Subjects gave ratings first on ascale of expectedness with 1 = completely unexpected and 5 = veryexpected, and then on a scale of fittingness with 1 = not fitting at allwith the preceding context and 5 = highly fitting with the previouscontext. Subjects had 3 s of silence between cadences in which togive their ratings and were offered three trials of training before thestart of the actual experiment.

Before the experiments, participants gave written informedconsent. The experiment was approved by the Ethics Committee ofthe former Department of Psychology, University of Helsinki, andconducted in accordance with the Declaration of Helsinki.

Data Recording and Analysis

The electroencephalography (EEG; sampling rate 4096 Hz down-sampled to 512 Hz offline; online band-pass filter 0.16–100 Hz)was recorded with a 64 AgCl active electrode cap using BioSemiActiView equipment. Additional electrodes were attached to themastoids, under the right eye, and on the nose. The data wereanalyzed in the MATLAB environment (MathWorks) either withfunctions adapted from EEGLAB (Swartz Center for Computa-tional Neuroscience) or FieldTrip (Centre for Cognitive Neuroim-aging of the Donders Institute for Brain, Cognition and Behaviour),or with custom-made functions.

The data were filtered offline using a 1–30 Hz band-passfilter and a baseline set to the mean 100-ms prestimulusinterval. The analysis epoch lasted from 100 ms prestimulus to600 ms poststimulus. Any EEG epoch containing a deflectionexceeding � 100 mV on at least eight channels was excluded fromfurther processing. The number of artifact-free trials and their ratioto the total number of trials is reported in Table 2.

To allow for comparison with previous ERAN literature, thedata were rereferenced to the mean value recorded from the twomastoid channels. Based on visual inspection of the ERPresponses, we set up two time windows for analysis, the first at100–230 ms and the second at 230–350 ms. Peak latencies in thesetwo time windows were computed from the vertex electrode (Cz).Mean amplitudes were instead calculated averaging overa � 20-ms window around the grand-average peaks at Cz anddefined for each group and stimulus condition. The mean ampli-tudes were extracted from the following electrodes: F1, F3, F5,

Table 2. Number of Artifact-Free Trials and Their Ratio with Number of Total Trials

Mean of artifact-free trials SD Ratio with total trials SD

Folk musiciansStandard for Neapolitan in third position 113.1 13.4 0.96 0.02Standard for Neapolitan in fifth position 111.9 15.3 0.95 0.03Standard for Neapolitan in seventh position 111.5 15 0.95 0.03Neapolitan in third position 113.7 12 0.95 0.03Neapolitan in fifth position 112.3 12.6 0.95 0.03Neapolitan in seventh position 117.6 13.6 0.95 0.03

NonmusiciansStandard for Neapolitan in third position 116.5 5.9 0.94 0.05Standard for Neapolitan in fifth position 115.8 4.7 0.94 0.04Standard for Neapolitan in seventh position 115 6.9 0.93 0.05Neapolitan in third position 117.8 7.8 0.94 0.04Neapolitan in fifth position 115.8 5.5 0.94 0.04Neapolitan in seventh position 115 5.4 0.94 0.04

656 E. Brattico et al.

FC1, FC3, FC5, C1, C3, C5, CP1, CP3, CP5, P1, P3, P5, F2, F4,F6, FC2, FC4, FC6, C2, C4, C6, CP2, CP4, CP6, P2, P4, and P6.

To test for the presence of differences between groups andstimulus conditions in neural responses or in behavioral ratings, weperformed repeated measures analyses of variance (ANOVAs) withgroup (folk musicians, nonmusicians) as the between-subjectsfactor and position (third, fifth, seventh) as the within-subjectfactor. For the ERP responses, we also included the followingwithin-subject electrode factors: laterality (left, right), frontality(frontal—including the electrodes F1, F3, F5, F2, F4, F5;frontocentral—FC1, FC3, FC5, FC2, FC4, FC6; central—C1, C3,C5, C2, C4, C6; parietocentral—CP1, CP3, CP5, CP2, CP4, CP6;parietal—P1, P3, P5, P2, P4, P6), and the electrodes included ineach region of interest (ROI; three levels including the electrodes atline 1 or 2, line 3 or 4, and line 5 or 6), when appropriate. Theequality of variances was measured using Mauchly’s test ofsphericity. The important variances for the study turned out to beequal. Thus, the results of the ANOVA could be used withoutstatistical corrections. For testing the directions of the significantmain effects, we performed LSD post hoc tests or with plannedpairwise t tests. Effect sizes for all the significant findings areindexed by the partial eta-squared ηp

2.

Results

ERP Responses at 100–230 ms

As visible from Figure 2, within the time window of the ERAN(100–230 ms), the Neapolitan chords at the three cadence positionselicited an enhanced frontocentral negative response peaking ataround 170 ms (this latency not differing between chord positions

or groups) in folk musicians as compared with nonmusicians (maineffect of group: F(1,20) = 10.96, p = .003; ηp

2 35= . ; for a summaryof the ANOVA results, see Table 3). No interactions includinggroup as a factor were significant (F < 1).

Across groups and electrodes, the position of the Neapolitanchord did not have a significant effect, F(2,40) = 2.43, p =

CzC3 C4

ERP responses of folk musicians

ERP responses of nonmusicians

CzC3 C4

Neapolitan at third positionNeapolitan at fifth position

Neapolitan at seventh position

–4 µV

–100 ms 400 ms

–100 ms 400 ms

+4 µV

–4 µV

+4 µV

Figure 2. Group difference waves shown for three central electrode sites (C3, Cz, and C4). These waves were obtained by subtracting the ERP responsesto Neapolitan chords at the three different cadence positions from the ERP responses to the corresponding congruous chords separately for folk musiciansand nonmusicians.

Table 3. Summary of ANOVA Results for the ERAN and P3aAmplitudes in Folk Musicians and Nonmusicians

F and p values and partial h2

ERP responses at 100–230 msGroup F(1,20) = 10.96, p = .003, h2 = 2.83Position ¥ Laterality F(2,40) = 3.8, p = .03, h2 = 6.36For the right hemisphere: Position F(2,40) = 4.2, p = .02, h2 = 5.88

ERP responses at 230–350 msPosition F(2,40) = 36.6, p < .001, h2 = 1.65For the third position: Laterality F(1,20) = 7.9, p = .01, h2 = 3.89For the third position: Frontality F(4,80) = 6.4, p = .001, h2 = 4.61For the seventh position:

FrontalityF(4,80) = 5.4, p = .02, h2 = 5.35

Group ¥ Position F(2,40) = 3.7, p = .03, h2 = 16.34For the fifth position: Group F(1,20) = 3.9, p = .06For the seventh position: Group F(1,40) = 5.7, p = .03, h2 = 4.5Group ¥ Frontality F(4,80) = 4.4, p = .003, h2 = 7.65Group ¥ Condition ¥

Frontality ¥ LateralityF(8,160) = 2.9, p = .005, h2 = 8.31

For frontal electrodes:Group ¥ Position

F(2,40) = 4.2, p = .02, h2 = 10.57

For frontocentral electrodes:Group ¥ Position

F(2,40) = 4.6, p = .02, h2 = 12.56

For central electrodes:Group ¥ Position

F(2,40) = 3.4, p = .04, h2 = 20.33

Modulated processing of harmony in folk musicians 657

.1. However, when focusing the analyses above the anterior andcentral electrodes, the effect of position approached significance,F(2,40) = 2.69, p = .08, indicating a tendency for the ERAN to thechord at the fifth position to be smaller than that at the seventhposition (p = .04). Moreover, since we found significant interactionPosition ¥ Laterality, F(2,40) = 3.8, p = .03, ηp

2 16= . ; e = .3, weseparately investigated the ERAN registered above the left andright hemispheres. While the ERAN recorded at left hemisphericelectrodes did not differentiate groups or chord positions (F � 1),we obtained a significant main effect for position from theright hemispheric electrodes, F(2,40) = 4.2, p = .02, ηp

2 17= . (seeFigure 3). This effect resulted from the larger ERAN amplitudeelicited by the chord at the seventh position as compared with theERAN amplitudes elicited by chords at the two other positions(third vs. seventh, p = .02 and fifth vs. seventh, p = .03). The ERANamplitudes to chords in the third and fifth positions, though, did notdiffer between each other.

ERP Responses at 230–350 ms

As illustrated in Figure 2, the amplitudes of the frontocentral ERPresponses peaking at around 285 ms (latencies not differing

between groups or chord positions) were modulated by groups andchord positions. Interestingly, the Neapolitans at the third andseventh positions of the cadence elicited a P3a-like responsewhereas the Neapolitan at the fifth position elicited a secondERAN-resembling response, especially in folk musicians.

The presence of a modulation by chord positions was con-firmed by the significant main effect of position, F(2,40) = 36.6,p < .0001, ηp

2 65= . , showing that the ERP in response to the chordat the fifth position had a more negative amplitude than the ERP atthe two other positions (p < .0001), hence confirming the elicitationof an ERAN-like response by this Neapolitan. Separate ANOVAsfor the position factor showed that the positive P3a-like responseelicited by the Neapolitan at the third position was right-lateralizedand maximal at the frontocentral ROI, as indicated in a separateANOVA by the significant main effects of laterality, F(1,20) = 7.9,p = .01, ηp

2 28= . ; right hemisphere: M = 1.6 mV � 0.3 SEM;left hemisphere: M = 1.2 mV � 0.3 SEM, and of frontality,F(4,80) = 6.4, p = .001, ηp

2 24= . ; frontal ROI: M = 1.5 mV � 0.4SEM: frontocentral ROI: M = 1.7 mV � 0.3 SEM; central ROI:M = 1.7 mV � 0.3 SEM; centroparietal ROI: M = 1.3 mV � 0.3SEM; parietal ROI: M = 1 mV � 0.3 SEM. At the fifth position, thenegative ERAN-like response was bilaterally distributed over the

Figure 3. Scalp maps of the early ERAN responses. Top: Voltage maps computed from a � 20-ms window around the most negative peak measured at Czwithin the 100–230 ms time interval, per each group and condition. Bottom: Scalp current density maps computed from a � 20-ms window around the mostnegative peak measured at Cz within the 100–230 ms time interval, per each group and condition.

658 E. Brattico et al.

scalp (main effect of laterality: F(1,20) = 1.6, p = .2; righthemisphere: M = -1.1 mV � 0.3 SEM; left hemisphere: M =-1.3 mV � 0.3 SEM, and main effect of frontality: F < 1). Finally,the positive P3a to chords at the ending seventh position was alsobilaterally distributed (main effect of laterality: F(1,20) = 3.4,p = .08; right hemisphere: M = 1.4 mV � 0.2 SEM; left hemi-sphere: M = 1.1 mV � 0.2 SEM) and maximal at frontocentralROI (as testified by the significant main effect of frontality:F(4,80) = 5.4, p = .02, e = .3, ηp

2 21= . ; frontal ROI: M =1.5 mV � 0.3 SEM: frontocentral ROI: M = 1.6 mV � 0.3 SEM;central ROI: M = 1.4 mV � 0.2 SEM; centroparietal ROI: M=1 mV � 0.2 SEM; parietal ROI: M 0.6 = mV � 0.2 SEM).

Importantly, while the groups did not differ in the overall ERPamplitudes across chord positions (F < 1 for the main effect ofgroup), they did differ in how they processed chords based on theirpositions. This was reflected by the P3a to the Neapolitan at thethird and seventh positions and by the ERAN elicited by the Nea-politan at the fifth position. Statistically, this was evidenced by thesignificant interaction Group ¥ Position, F(2,40) = 3.7, p = .03,ηp

2 16= . . Separate ANOVAs for the position factor revealed a sig-nificant main effect of group for the seventh position,

F(1,40) = 5.7, p = .03, ηp2 22= . , deriving from an enhanced posi-

tive ERP response in folk musicians compared with nonmusicians(musicians: M = 1.7 mV � 0.3 SEM; nonmusicians: M = 0.8 mV �0.3 SEM). A marginally significant main effect of group wasalso found for the fifth position, F(1,20) = 3.9, p = .06, ηp

2 16= . ,this time resulting from a larger negative ERP deflec-tion in folk musicians versus nonmusicians (musicians: M =-1.7 mV � 0.4 SEM; nonmusicians: M = -0.8 mV, � 0.4 SEM).The chord at the third position elicited a positive P3a-like response,which did not differentiate folk musicians from nonmusicians(F < 1; musicians: M = 1.6 mV � 0.4 SEM; nonmusicians:M = 1.3 mV � 0.4 SEM).

Further, as evidenced by the voltage and current source distri-bution maps in Figure 4, the folk musicians also differed fromnonmusicians in the scalp distribution of their ERP responses. Thiswas confirmed by the significant interactions Group ¥ Fronta-lity, F(4,80) = 4.4, p = .003, ηp

2 18= . , and Group ¥ Condition ¥Frontality ¥ Laterality, F(8,160) = 2.9, p = .005, ηp

2 13= . .A separate ANOVA for amplitudes measured at the frontal ROI

showed that the ERP responses of folk musicians differed fromthose of nonmusicians according to the position of the deviating

Figure 4. Scalp maps of the P3a and late ERAN responses. Top: Scalp maps of the early ERAN responses. Voltage maps computed from a � 20-ms windowaround the (negative or positive) peak measured at Cz within the 230–350 ms time interval, per each group and condition. Bottom: Scalp current density mapscomputed from a � 20-ms window around the (negative or positive) peak measured at Cz within the 230–350 ms time interval, per each group and condition.

Modulated processing of harmony in folk musicians 659

chords in the cadence (Group ¥ Position interaction: F(2,40) = 4.2,p = .02, ηp

2 17= . ) but did not differ across chord conditions (maineffect of group: F < 1). Importantly, post hoc LSD tests indicatedthat the ERAN-like response to the chord at the fifth position infolk musicians was more negative than that in nonmusicians(p = .04; M = -1.9 mV � 0.3 SEM for folk musicians, M =-0.6 mV � 0.2 SEM for nonmusicians) whereas the P3a-likeresponse to the chord in the seventh position in folk musicianswas more positive than that in nonmusicians (p = .03; M =2.3 mV � 0.4 SEM for folk musicians, M = .07 mV � 0.3 SEM fornonmusicians). No group difference at the frontal ROI wasobserved for the P3a-like responses elicited by the chord at thethird position (p = .5; M = 1.9 mV � 0.3 SEM for folk musicians,M = 1.3 mV � 0.3 SEM for nonmusicians). At frontocentral andcentral ROIs, the group factor also significantly interacted with theposition factor, F(2,40) = 4.6, p = .02, ηp

2 19= . , for frontocentralROI, and F(2,40) = 3.4, p = .04, ηp

2 3= . for central ROI. This inter-action derived from the larger P3a to the ending chord in folkmusicians compared with nonmusicians (in post hoc LSD tests:p = .01 for frontocentral ROI and p = .03 for central ROI). Nodifferences between groups in ERP amplitudes were instead foundfor the other chords (p > .09). The ERP responses at centroparietaland parietal ROIs did not yield any significant Group ¥ Positioninteraction, F(2,40) = 2, p = .2 and F < 1, respectively.

Behavioral Ratings

As illustrated by Figure 5, the ANOVA for the ratings of fittingnessrevealed a main effect of chord, F(3,72) = 50.3, p < .0001,ηp

2 68= . ; e = .67, resulting from the highest ratings of fittingnessfor the standard cadences (in post hoc LSD test: p < .0001) and thelowest for the cadences containing the Neapolitan in the endingposition (p < .0001), with the cadence having the Neapolitan at thefifth position marginally rated as more fitting than the one with theNeapolitan at the third position (in paired samples t tests: p = .06).Furthermore, folk musicians gave overall higher ratings of fitting-ness (M = 4 � 0.2 SEM) to the cadences than nonmusicians(M = 3.4 � 0.1 SEM), as testified by a significant main effect ofgroup, F(1,24) = 7.9, p = .01, ηp

2 25= . . However, pairwise groupcomparisons evidenced that the main effect of group derived fromthe higher ratings of folk musicians for all cadences, t(24 > 2.8,p < .02 for the cadences containing the ending tonic chord and theNeapolitans at the third and fifth positions) except for those havingthe Neapolitan at the seventh position, t(24) = 1.1, p = .3. Notably,folk musicians rated the cadence having the Neapolitan at the thirdposition (M = 4.1 � 0.2 SEM) similarly to the cadence containingthe Neapolitan at the fifth position (M = 4.2 � 0.2 SEM; t(11) = .9,p = .4, whereas nonmusicians tended to judge the cadence with theNeapolitan at the fifth position as more fitting with the context(M = 3.5 � 0.2 SEM) compared with the cadence with the Neapoli-tan at the third position (M = 3.2 � 0.2 SEM; t(13) = -2, p = .07).

The ANOVA for the ratings of expectedness similarly revealeda significant main effect of chord, F(3,72) = 73.7, p < .0001,ηp

2 73= . ; e = .74, derived from the highest ratings for standardcadences and the lowest for cadences ending with a Neapolitan,and from the nondiffering ratings between the other two cadences(p = .9). We also obtained a tendency for significance for the maineffect of group, F(1,24) = 4.1, p = .05, ηp

2 15= . , with higher ratingsof expectedness by folk musicians (M = 3.7 � 0.1 SEM) than bynonmusicians (M = 3.3 � 0.1 SEM). Independent samples t testsfor the expectedness ratings showed that folk musicians rated thestandard cadences, t(24) = 2.3, p = .03; M = 4.9 � 0.1 SEM for

folk musicians and M = 4.6 � 0.1 SEM for nonmusicians, andtended to rate the cadences with the Neapolitan at the third posi-tion, t(24) = 2, p = .05; M = 3.8 � 0.2 SEM for folk musicians andM = 3.2 � 0.2 SEM for nonmusicians, as more expected comparedto nonmusicians. Ratings of expectedness for the rest of thecadences did not vary between experimental groups (p > .1).

Correlations between ERP Responses and Behavioral Ratings

We conducted Spearman’s nonparametric correlation tests to checkany link between the neural responses and music perception ofthose subjects who participated in both the ERP and behavioralexperiments. We opted for nonparametric tests to account for thesmall N of this sample. Interestingly, the only significant correla-tion was a negative one in folk musicians between ratings of fit-tingness for the Neapolitan at the fifth position and the early ERANresponse to the same chord, rho(8) = -.8, p = .01, indicating thatthe more folk musicians perceived the cadence containing the mild

Ratings of Expectedness

Ratings of Fittingness

Standard Third position Fifth position Seventh position

Ra

tin

gs

1

2

3

4

5

Folk musicians

Non-musicians

1

2

3

4

5

Standard Third position Fifth position Seventh position

Ra

tin

gs

Figure 5. Means of the behavioral ratings of expectedness (top) andfittingness (bottom) for each experimental group and chord cadence.

660 E. Brattico et al.

violation at the fifth position as fitting (i.e., less violating) with thecontext the larger the early ERAN elicited by it was.

Discussion

The present study investigated ERPs to Neapolitan chords violatingexpectations for chord resolution depending on their position in acadence. Here, for the first time, the musical practice of the subjectsand their behavioral responses to chord cadences were carefullyclarified by a background questionnaire, a musicality test, and byselecting musicians having, along with basic studies in Westerntonal music, also a special training in Finnish folk music. This lattermusical genre, melodically, harmonically, and rhythmically differ-ent from the Western major-minor music system, is a pertinentmodel for studying neuroplasticity to harmony processing sincenone of its music systems include a Neapolitan chord. Hence, bycomparing the ERPs to unexpected chords elicited in folk musi-cians and nonmusicians, we determined whether long-term expo-sure to and practice of Finnish folk music would modulate theERAN responses as well as the expectedness and fittingness ratingsof Neapolitan chords not present in Finnish folk music.

We found that, overall, Finnish folk musicians responded withan enhanced ERAN peaking at around 170 ms to violations ofchord expectations when compared to nonmusicians. The Nea-politan placed at the third and seventh positions of the cadenceelicited a P3a (subsequent to the ERAN) in both folk musiciansand nonmusicians, but only the P3a to the ending chord wasclearly enhanced in folk musicians compared with nonmusicians.We also observed a second slightly strengthened ERAN in folkmusicians compared with nonmusicians in response to the milderviolation of chord expectations when the Neapolitan worked as asubdominant in the fifth position of the cadence. We interpretthese findings as showing a culture-dependent neuroplasticity ofbrain responses to violations of musical expectations. In folkmusicians, the enhanced late ERAN to the Neapolitan at the fifthposition was accompanied by ratings of fittingness for thiscadence not differing from those for the Neapolitan at the thirdposition. Hence, interference from knowledge of other musicsystems in folk musicians resulted in additional inferofrontalneural activity to a chord succession, which would otherwise beperceived as quite conventional by listeners accustomed primarilyto Western tonal music. In turn, the enhanced P3a at the seventhposition might reflect the stronger salience of the incongruentsound mismatching with harmonic expectations.

According to the initial interpretation of Koelsch et al. (2000),a stronger expectation for a tonic chord at the end of an authenticcadence increases the ERAN response. However, we recently dem-onstrated that the preceding chord, rather than the position of agiven chord within a cadence as such, determines the responses tothe current Neapolitan chord (Garza Villarreal et al., 2011; Leinoet al., 2007). This was evidenced by a larger ERAN elicited by theNeapolitan when it followed a dominant chord than when it fol-lowed a tonic chord. This effect was obtained even by contrastingthe degree of violation of the conventions of chord succession withthe building of the tonal context: the Neapolitan following thedominant in the third position, hence replacing the expected tonicchord, elicited a larger ERAN than the Neapolitan following thetonic at a later (fifth) position in the cadence, thereby playing therole of a subdominant chord. In the present study, we found thatthe ERAN amplitudes to the Neapolitans at the third and fifthpositions did not differ from each other, whereas a pronouncedERAN was elicited by the Neapolitan at the last position of the

cadence. This suggests that the tonal context affected the ERANamplitude only at the ending chord, but did not cause an ERANenhancement dependent on the accumulation of chord presenta-tions when the violation of harmonic conventions was milder. Nev-ertheless, this ERAN effect of the chord position in the cadencewas most noticeable in the frontocentral regions of the right hemi-sphere. Such a finding replicates what has been observed in severalprevious ERAN studies indicating predominant ERAN generatorsin the right inferior frontal cortex and superior temporal gyrus(Garza Villarreal et al., 2011; Koelsch, Fritz, Schulze, Alsop, &Schlaug, 2005; Maess, Koelsch, Gunter, & Friederici, 2001). Tonote, a very recent ERP study with brain-lesioned patients eluci-dated a supportive role of the left pars opercularis in processingharmonic incongruity. This was evidenced by a decreased recog-nition of deviating chords and an enhanced ERAN in the righthemisphere in patients with lesions in that brain area as comparedto the ERAN in healthy subjects and in patients with left anteriortemporal lobe damage (Sammler et al., 2011). However, these find-ings do not rule out the contribution of right hemispheric structuresin music harmony processing.

The early ERAN was enhanced in folk musicians as comparedto nonmusicians irrespective of the position of the Neapolitanchord in the cadence. This finding replicates and extends what hasbeen observed in two separate studies (Koelsch et al., 2002; Müller,Höfel, Brattico, & Jacobsen, 2010) in classically trained musicians.We explain this finding by the deep roots of folk musicians inWestern tonal music, a consequence of continued implicit exposurein Finland and also formal education in music schools and con-servatories prior to entering their studies in folk music. It seemsthat, taking note of the idea that ERAN is “sensitive to culture-specific knowledge that is built up through exposure to a particularmusic system” (Trainor & Zatorre, 2009, p. 176), folk musicianshave developed sensitivity to the Western tonal music system, andpossibly to other music systems as well. The multimusicality offolk musicians instead of diminishing the ERAN due to their expo-sure to non-Western music, resulted in overall reinforced neuralrepresentations of chords representative of one of the musicsystems they are accustomed to, namely, Western tonal music. Thisinterpretation is supported by our finding of a significant negativecorrelation in folk musicians between the early ERAN to the Nea-politan at the fifth position and the fittingness rating for the corre-sponding cadence. When folk musicians considered this cadencecongruous, they complied with the theory-based prediction accord-ing to which the Neapolitan plays the role of a subdominant asencountered in Romantic chromatic music. In other words, the folkmusicians who rated the cadence as more fitting possessed a deeperacculturation with Western tonal harmony conventions. Thisresulted in an enhanced early ERAN, hence indexing a strongerneural representation of Western harmonic conventions in folkmusicians. In turn, folk musicians for whom the cadence succes-sions of Western tonal music were less well incorporated (as aconsequence either of their less formal instruction in Westernmusic or higher exposure and practice of folk music) displayedweaker ERAN to the Neapolitan at the fifth position. The relationbetween the amplitude of the early ERAN and fittingness ratingsthus provides further support for the dependence of the ERAN ondegrees of acculturation with music conventions. Although it mightbe challenging in our globalized world, future studies should aimfirst at targeting musicians exposed only (or mainly) to non-Western music systems, and second at utilizing a paradigm withnon-Western music sequences familiar to the subjects. In thesemusicians, a reduction of the early ERAN response in Western

Modulated processing of harmony in folk musicians 661

cadences might be observed as opposed to an ERAN increase innon-Western cadences.

Interestingly, we also found that the subsequent ERP deflectionspeaking at about 285 ms were modulated by both group and chordposition in the cadence. More specifically, a frontocentral positivitywas elicited in all subjects to Neapolitan chords at the third andseventh positions, and enhanced in response to the Neapolitanchord at the seventh position only in folk musicians when com-pared with nonmusicians. Our interpretation is that, based on itslatency, polarity, and topography, this deflection can be labeled asP3a reflecting an involuntary switch of attention towards the Nea-politan chord particularly in folk musicians (Friedman, Cycowicz,& Gaeta, 2001; Polich, 2007). To note, the amplitude of the P3a inresponse to infrequent tones did not differentiate professionalmusicians from nonmusicians in prior studies by Tervaniemi, Just,Koelsch, Widmann, & Schröger (2005) and Nikjeh, Lister, andFrisch (2009; although the P3a latency can be modulated by groupmembership: see also Nikjeh et al., 2008). Recent evidence bySeppänen, Pesonen, and Tervaniemi (2012) points instead at anincrease of attentional sound-related skills in musicians, evidencedduring the perceptual learning of sound discrimination. Steinbeiset al. (2006) similarly found an effect of musical expertise on theP3a. They obtained a P3a following an ERAN to the highly unex-pected Neapolitan chords placed at the end of (modified) choralesby Bach. Notably, this P3a was enhanced in musicians trained atthe conservatory compared with nonmusicians (in nonmusicians, asmall P3a was obtained also in response to the mildly unexpectedchords originally composed by Bach). None of these studies,though, provide any details about the acculturation of these musi-cians with certain music cultures or genres or about their preferredmusical style, hence leaving unresolved the question on how bimu-sicalism or multimusicalism might affect auditory involuntaryattention skills. Additionally, in our study, the P3a elicitation maybe a consequence of the attention demands requested for by thepresent semiattended paradigm. Although subjects were asked tofocus on the infrequent organ timbre (as opposed to an MMNparadigm, which compels subjects to fully ignore auditory stimu-lation), they were attentively listening to the cadences, and theirattentional focus might have diverted to the salient Neapolitanchords, generating a P3a response. In the current context, the P3aelicitation is a novel finding and, due to group comparison designand P3a sensitivity to the expertise of folk musicians, a highlyinteresting one. Its elicitation can be interpreted as reflecting aneural sensitivity of folk musicians targeting their attention towardsthe ending chords that do not match their expectations for the tonic.

In response to the Neapolitan at the fifth position only, a fron-tocentral negativity peaking at around 285 ms was elicited. Thislate ERAN-resembling response was enhanced in folk musicians ascompared to nonmusicians (in which it approached the baseline).The Neapolitan at the fifth position, based on the conventions ofchromatic Romantic music style, is only mildly incongruous andworks as a subdominant within the cadence. The enhanced lateERAN in folk musicians seems to reflect their divergent neuralprocessing of Western tonal chord conventions. To support such aninterpretation and relate the neural data with chord perception, weconducted a behavioral experiment with most of the subjects whohad participated in the ERP experiment. The ratings of fittingnessand expectedness in folk musicians as compared to nonmusicianswere overall higher especially for the Neapolitan chords insertedinside the cadence. In particular, folk musicians judged the Nea-politans at the third and fifth position as quite fitting and they didnot differentiate those chords from each other, whereas nonmusi-

cians judged these chords as neutral (neither fitting nor unfitting)and tended to differentiate them. Pairwise comparisons, in contrast,did not show a significant difference between groups for the nega-tive fittingness and expectedness ratings of the Neapolitan termi-nating the cadence. The peculiar representation of Westernharmonic conventions in folk musicians is also substantiated by thelack of significant differences between their fittingness ratings forthe cadences having the Neapolitan at the third or fifth positions,contrasted with a trend for a difference in nonmusicians. Hence, theNeapolitan chords within the cadence, and particularly the one atthe fifth position, were perceived as challenging or interesting, butperhaps without the perception of the nuances in their fittingnessbased on Western harmonic conventions as a result of the interfer-ence with neural substrates for the Finnish folk music system.Summing up the ERAN and behavioral data, it is reasonable tospeculate that folk musicians, due to their training in several kindsof music systems, have developed a flexible and deviating way ofprocessing the conventions of the Western tonal system that doesnot coincide with the typical neural processing and behavior ofindividuals accustomed primarily to Western music culture. Futurestudies should compare musicians exclusively exposed to Westernclassical music and others exposed to various music cultures or,optimally, only to non-Western music cultures to further disentan-gle the general effects of musical expertise with those that areselective of a single music system.

Taken together, the current findings provide a novel compre-hensive approach to music neurocognition, ranging from basicauditory discriminatory abilities (assessed with the AMMA test) toperceptual (fittingness and expectedness) attributes, and neural(ERAN and P3a) correlates of harmony processing in a particulartype of musician possessing expert knowledge of both Finnish folkmusic and Western tonal culture. The findings indicate superiorneurocognitive ability for chord processing in Finnish folk musi-cians, possibly resulting from their active training in more than onemusic culture, which is activated when harmonic encoding isrequired. This general ability for chord processing is reflected inthe overall enhanced ERAN to all chord incongruities and the P3ato the strongest violation of chord expectations. In addition, folkmusicians showed divergent behavioral ratings of the cadencesincluding the Neapolitan chord (not existing in Finnish folk music),and particularly of the mildest violation of chord expectation, com-bined with a corresponding enhanced late ERAN. Thus, Finnishfolk musicians, less exposed than nonmusicians to the Neapolitanchord, and more exposed to other nondiatonic music systems,demonstrate a neural reactivity to musical rule deviations, whichseems to both qualitatively and quantitatively depart from the har-monic processing in the brain shown in individuals primarilyexposed to Western tonal music. The present study can be consid-ered solely as an initial step towards the definition of neuroplastic-ity in bimusical or multimusical individuals (see Wong, Roy, &Margulis, 2009), and is confined to one kind of folk musicians notexposed to one kind of chords in the genre they are primarilyperforming. Nevertheless, our initial findings do support theemerging view (e.g., Margulis, 2008; Tervaniemi, 2009; Vuustet al., 2012) of the importance of listening/playing biography fordiscerning the intersubject variability in auditory neurocognition.By adopting an approach similar to ours, namely, by obtaining acomprehensive overview of the perceptual and neural correlates ofthe basic and higher-order musical skills while controlling for themusic background variables, future studies with different kinds ofmusicians and stimuli from different music cultures will ultimatelyresolve the issue of neuroplasticity of music cognition.

662 E. Brattico et al.

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(Received January 5, 2012; Accepted, February 7, 2013)

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