rHuman Brain Mapping 30:588–601 (2009) r

Language and Music: Differential Hemispheric
Dominance in Detecting Unexpected Errors in the
Lyrics and Melody of Memorized Songs
Takuya Yasui,1,2,3 Kimitaka Kaga,2,4 and Kuniyoshi L. Sakai1,3*
1

Department of Basic Science, Graduate School of Arts and Sciences, The University of Tokyo, Komaba,
3-8-1 Komaba, Meguro-ku, Tokyo 153-8902, Japan2
Department of Otolaryngology, Graduate School of Medicine, The University of Tokyo, 7-3-1 Hongo,
Bunkyo-ku, Tokyo 113-8655, Japan
3
CREST, Japan Science and Technology Corporation, Kawaguchi-shi 332-0012, Japan
4
National Institute of Sensory Organs, 2-5-1 Higashigaoka, Meguro-ku, Tokyo 152-0021, Japan

Abstract: Using magnetoencephalography (MEG), we report here the hemispheric dominance of the auditory cortex that is selectively modulated by unexpected errors in the lyrics and melody of songs (lyrics andmelody deviants), thereby elucidating under which conditions the lateralization of auditory processingchanges. In experiment 1 using familiar songs, we found that the dipole strength of responses to the lyrics
deviants was left-dominant at 140 ms (M140), whereas that of responses to the melody deviants was rightdominant at 130 ms (M130). In experiment 2 using familiar songs with a constant syllable or pitch, thedipole strength of frequency mismatch negativity elicited by oddballs was left-dominant. There were signiﬁcant main effects of experiment (1 and 2) for the peak latencies and for the coordinates of the dipoles,
indicating that the M140 and M130 were not the frequency mismatch negativity. In experiment 3 using
newly memorized songs, the right-dominant M130 was observed only when the presented note was unexpected one, independent of perceiving unnatural pitch transitions (i.e., perceptual saliency) and of selective
attention to the melody of songs. The consistent right-dominance of the M130 between experiments 1 and 3
suggests that the M130 in experiment 1 is due to unexpected notes deviating from well-memorized songs.
On the other hand, the left-dominant M140 was elicited by lyrics deviants, suggesting the inﬂuence of
top-down linguistic information and the memory of the familiar songs. We thus conclude that the leftlateralized M140 and right-lateralized M130 reﬂect the expectation based on top-down information of
language and music, respectively. Hum Brain Mapp 30:588–601, 2009. V2008 Wiley-Liss, Inc.
C

Key words: hemispheric dominance; auditory processing; temporal cortex; magnetoencephalography;
mismatch negativity

Contract grant sponsor: Ministry of Education, Culture, Sports, Science, and Technology of Japan; Contract grant numbers: 17022013;
Contract grant sponsor: Japan Science and Technology Agency (JST).
*Correspondence to: Kuniyoshi L. Sakai, Department of Basic Science, Graduate School of Arts and Sciences, The University ofTokyo, Komaba, 3-8-1 Komaba, Meguro-ku, Tokyo 153-8902, Japan.
E-mail: sakai@mind.c.u-tokyo.ac.jp
Received for publication 17 July 2007; Revised 14 November 2007;
Accepted 15 November 2007
DOI: 10.1002/hbm.20529
Published online 2 January 2008 in Wiley InterScience (www.
interscience.wiley.com).
C 2008 Wiley-Liss, Inc.
V

INTRODUCTION
The higher brain functions related to language andmusic are thought to involve uniquely human abilities[Lerdahl and Jackendoff, 1983; Patel, 2003], and they are
known to have a strong tendency for hemispheric dominance in the brain. Syntax, semantics, and phonology are
generally lateralized with left hemispheric dominance[Gazzaniga, 2000; Geschwind, 1979; Price, 2000; Sakai,2005], whereas the right hemispheric dominance in processing prosodic information has been reported in frontal

r Lateralization in Monitoring Lyrics and Melody r

and temporal regions [Hesling et al., 2005; Nicholson et al.,
2003; Plante et al., 2002; Ross, 1981]. On the other hand,right temporal lesions are reported to cause amusia, ordeﬁcits in the discrimination of melodies [Ayotte et al.,
2000; Dennis and Hopyan, 2001; Grifﬁths et al., 1997;
Liégeois-Chauvel et al., 1998; Murayama et al., 2004; Nicholson et al., 2003]. A positron emission tomography study
on healthy participants has also reported that activation of
the right superior temporal cortex is enhanced during passive listening to melodies [Zatorre et al., 1994]. Active listening to instrumental music activates both hemispheres,with right-hemispheric weighting [Koelsch et al., 2002;
Kraemer et al., 2005], and previous lesion studies haveindicated that long-term memory of musical information,in contrast to verbal information, is not clearly lateralized
in one hemisphere [Samson, 1999; Stewart et al., 2006]. To
establish the right hemispheric dominance of auditory
processing more conclusively, cortical activation should befurther examined with sufﬁcient temporal resolution.
Recent magnetoencephalography (MEG) studies withspeech sounds have reported that phoneme or voice oddballs induce the left-dominant mismatch negativity(MMNm) [Knösche et al., 2002; Näätänen et al., 1997]. This
type of ‘‘frequency MMNm’’ is generated by an automatic
detection of frequency errors (deviants) occurring in a series of standard stimuli, and the frequency MMNm can be
elicited even by auditory oddballs eliciting no N1 response[Näätänen et al., 2005]. An MEG study has reported thatthe frequency MMNm for chord oddballs was larger thanthat for phoneme oddballs in the right hemisphere, but theright dominance of the MMNm for chord oddballs was
not signiﬁcant [Tervaniemi et al., 1999]. In these previous
studies, speech sounds were used for language stimuli,
while the instrumental sounds or chords were used formusic stimuli. Thus the bottom-up processes for the acoustic features of stimuli can explain the contrasting hemispheric dominance between language and music processing. Speech sounds are highly dependent on rapidlychanging sounds whereas tonal music patterns tend to be
slower; thus, hemispheric dominance of the superior temporal cortex may depend on the temporal and spectral
properties of the acoustic stimuli and associated bottomup processes [Boemio et al., 2005; Zatorre et al., 2002].
However, it remains unclear how the hemispheric dominance is inﬂuenced by top-down processes including attention and memory.
A previous event-related brain potential (ERP) study
examined the effect of familiar melodies with a regular but
unexpected ending, or with an irregular tone, and reported
that a late positive component with the peak latency of
300 ms was elicited by these wrong notes [Besson andFaı̈ta, 1995]. Another ERP study reported that the N400
component was elicited by memory violations, whereas an
early right anterior negativity (ERAN) with the peaklatency of 200 ms was elicited by out-of-key violations[Miranda and Ullman, 2007]. The ERAN was also associated with expectancy based on musical regularity [Koelschret al., 2000; Maess et al., 2001]. It remains to be elucidated
how the expectancy of melody itself based on stored memory affects early components. If the exact timing of the ﬁrst
component reﬂecting the top-down processes based on
stored memory could be clariﬁed, this would further elucidate basic auditory processing.
To examine such top-down processes, experiments inwhich participants listen actively to song stimuli with bothproperties of lyrics and melody would be ideal, because
the temporal and spectral properties of the song stimuli
can be equated as much as possible. Figure 1A shows one
example of song stimuli used in the present study, which
was made with a speech synthesis program that can be
conﬁgured to produce songs by assigning musical notes
and lyrics. Using this program, the duration, pitch, andpower of each note can be held constant. We used aforced-choice error-detection paradigm, which has been
established in previous studies [Embick et al., 2000]; ineach trial, there were always one or two unexpected errors
in the lyrics and melody of songs (lyrics and melody deviants). In experiment 1 using familiar songs, we ﬁrst compared the hemispheric dominance between language andmusic tasks, where identical song stimuli were used without any instrumental accompaniment or chords. We
detected event-related MEG responses to either the lyrics
or melody deviants in two tasks performed separately: alyrics and a melody task (Fig. 1B). Here the stored memoryof songs was required, since there was no cue in the
acoustic features per se to discriminate deviants from thenormal reference stimuli. In experiment 2 using familiar
songs with a constant syllable or pitch, which did not
require stored memory of songs, we examined the effect of
oddballs in two tasks performed separately: a syllable anda pitch task (Fig. 1C). We tried to clarify whether the deviant-induced ﬁelds observed in experiment 1 were different
from the frequency MMNm. In experiment 3 using unfamiliar and newly memorized songs, we further tried toseparate the top-down expectation processes from the bottom-up perception of unnatural pitch transitions in the melody task (Fig. 1D), thereby conﬁrming the consistency of thedeviant-induced ﬁelds between experiments 1 and 3.

MATERIALS AND METHODS
Participants
Sixteen native Japanese speakers participated in thepresent study (aged 21–32 years, mean 27 years; eightfemales). Fourteen participants participated in experiment1, nine of whom also participated in experiment 2 alongwith an additional participant (a total of 10 participants).
Eleven participants participated in experiment 3, eight ofwhom also participated in both experiments 1 and 2 but oneof them was discarded due to low performance at the training stage (a total of 10 participants), and another two ofwhom had participated in either experiment 1 or 2. All participants showed right-handedness by the Edinburgh inventory589 r

r Yasui et al. r

Figure 1.
The experimental paradigm with songs to examine the differen- preceding the ﬁrst appearing deviant was used as the referenceces between lyrics and melody processing. A: An example of 12 for deviant-induced ﬁelds (gray rectangles). C: Experiment 2,
original songs used in experiments 1 and 2 (e.g., a French folk employing syllable and pitch tasks on familiar songs with a consong ‘‘Twinkle, twinkle, little star’’ with typical Japanese lyrics). stant syllable or pitch. In the syllable task, lyrics were replaced
Each song has seven notes, and is sung without any instrumental with the constant syllable of ‘‘na.’’ A blue circle denotes a syllable
accompaniment and chords. The time line of one trial is shown oddball. In the pitch task, the melody was replaced with the conbelow the musical score. In experiment 3, a different set of stant note of A4 (440 Hz). A red circle denotes a pitch oddball.
songs was used with the same time sequence. B: Experiment 1, The note preceding the ﬁrst appearing oddball was used as theemploying lyrics and melody tasks on familiar songs. There were reference (gray rectangles). D: Experiment 3, employing the melone or two deviants in lyrics (a blue circle) or melody (a red ody task on newly memorized songs. See Materials and Methods
circle) among the third to the seventh notes. The normal note for full explanation.

[Oldﬁeld, 1971], and had normal hearing abilities without
professional singing or instrument training besides school
education. The study was approved by the institutional
review board of The University of Tokyo, Komaba.

Stimuli and Tasks
The song stimuli without accompaniment were produced with SMARTTALK software (Oki Electric Industry,
Tokyo, Japan). For each trial in all experiments, a songstimulus with seven notes (600 ms each) was presented ata sound pressure level of about 85 dB (Fig. 1A). There
were always one or two deviants among the third to theseventh notes, and the second deviant appeared two orrthree sounds apart from the ﬁrst one, but in a pseudorandomized position, so that the melody after the ﬁrst
deviant clearly returned to the original one. After the songstimulus, a pure tone of 1,000 Hz was presented for 50 ms
to initiate the button press within the following 2 s. The
participants were informed that there would always be
one or two deviants in each trial, and they indicated thenumber of deviants by pushing one of two buttons. In
experiments 1 and 2, MEG responses to the ﬁrst appearing
deviant/oddball and those to a reference note just before it
were used for the analysis; the responses to the second
deviant/oddball were not used because they might reﬂect
priming or habituation effects. In experiment 3, MEG responses to the third note and those to the second reference

590 r

r Lateralization in Monitoring Lyrics and Melody rnote were used for the analysis. The trial interval was
randomized within the range of 62% at 6.8 s to reduceany periodical noises. The stimuli were delivered binaurally to participants via plastic tubes (length, 6.5 m), whichwere connected to insert earphones (Etymotic ER-30; Etymotic Research, Elk Grove Village, IL). Stimulus presentation and behavioral data collection were controlled using
the LabVIEW software and interface (National Instruments, Austin, TX).

Experiment 1
We prepared for a sequence of 12 familiar songs (original songs), each consisting of seven quarter notes. The
order of these 12 song stimuli was always ﬁxed, so thatthe participants could expect lyrics and melody even fromthe beginning of each song. For the lyrics task, lyrics deviants were made by changing both consonants and vowels
in two ways for each syllable (e.g., ‘‘wa’’ and ‘‘bu’’ from‘‘ki’’), while retaining the original melody (Fig. 1B). The
songs with lyrics deviants were phonotactically legal, butthey were syntactically and semantically anomalous. On
the basis of such top-down linguistic information and thememory of familiar songs, the participants responded to
one or two lyrics deviants. For the melody task, melody
deviants were made by raising or lowering the original
note by ﬁve semitones, while retaining the original lyrics(Fig. 1B). On the basis of the memory of familiar songs,
the participants responded to one or two melody deviants.
For each task, 228 trials were conducted for about 25 min; thelast three trials were excluded from the analysis because oftechnical problems with the program. Behavioral data were
not successfully collected from three participants due to technical problems with the buttons. The order of the two tasks
was counterbalanced among the participants.

Experiment 2
In the syllable task, all lyrics were replaced with theconstant syllable of ‘‘na’’ while keeping the original melody
used in experiment 1. The participants responded to one
or two oddballs consisting of the syllables ‘‘ki’’ or ‘‘so’’ ineach song stimulus (Fig. 1C). In the pitch task, all melodies
were replaced with the constant note of A4 (440 Hz) while
keeping the original lyrics used in experiment 1. The participants responded to one or two oddballs, which weremade by raising or lowering the constant note by ﬁve
semitones. The location and number of oddballs in thisexperiment were the same as those of the deviants in experiment 1. For each task, 228 trials were conducted for about
25 min; the last three trials were excluded from the analysis
due to technical problems with the program. The order of thetwo tasks was counterbalanced among the participants.

Experiment 3
As shown in Figure 1D (orange column), the participants memorized a sequence of the eight unfamiliar songsr(a, b’, a’, . . .) just before the MEG recordings. The order ofthese eight songs was always ﬁxed, so that the participants
could expect lyrics and melody even from the beginningof each song. On the basis of the memorized sequence, theparticipants responded to one or two melody deviants.
Here, we deﬁne the expected note condition as presenting
a note from the memorized sequence (the third note in theorange column), and the unexpected note condition aspresenting a melody deviant (the third note in the purple
column). If the expected note condition consisted of theoriginally composed notes alone whereas the unexpected
note condition included the experimentally modiﬁed notes,
it was not possible to separate the top-down expectation
processes from the bottom-up perception of unnatural
pitch transitions. To equate the melody stimuli between
these conditions, each condition included both originallycomposed notes (green circles) and experimentally modiﬁed notes (blue circles), whereas the second notes wereused as the reference (gray rectangles). From an original
song (a, b, c, and d), each modiﬁed song (a’, b’, c’, and d’)
was made by raising or lowering the third note alone by
ﬁve semitones. This procedure was identical with that formaking melody deviants, but the presence of modiﬁed
songs did not hamper memorization, since the original
songs were totally unfamiliar to the participants before theexperiment. Moreover, a melody deviant at the third noteof a modiﬁed song (e.g., a’) was made always identicalwith the third note of the corresponding original song (afor a’), thereby strictly equating the stimulus properties.
Melody deviants were also present in the fourth to the seventh notes as in experiment 1, but they were not used forthe analysis.
After an initial self-paced memorization block for 10
min, a conﬁrmation test consisting of eight trials of the
melody task was performed twice. If participants failed toscore seven out of eight, another memorization block wasadministered. All but one participant successfully scoredseven out of eight in two consecutive tests, within two or
three memorization blocks (20–30 min in total). The MEG
recordings were then started for the successfully passed
participants. During the MEG recordings, only the melody
task was performed, using the same design as in experiment 1. In each of three separate blocks, 128 trials were
performed for about 15 min each.

MEG Recordings and Analyses
The MEG data were acquired with a 160-channel wholehead recording system (MEGvision; Yokogawa Electric
Corporation, Kanazawa-city, Japan). Signals were digitizedon-line with a bandwidth of 0.3–1,000 Hz at a sampling
rate of 2,000 Hz. From the MEG data, only artifact-free trials (peak-to-peak amplitude <3,000 fT and without eye
movement) were selected and averaged separately for thedifferent types of deviant/oddball and reference stimuli.
In experiments 1 and 2, both correct and incorrect trials
were analyzed, whereas in experiment 3, only correct trials

591 r

r Yasui et al. rwere used for analyses because the accuracies were lower.
The averaged data were further ﬁltered with an off-line
band-pass ﬁlter of 1–20 Hz [Sinkkonen and Tervaniemi,
2000].
The MEG responses from 2100 ms to 600 ms (0 ms at
the onset of each note) were analyzed with a time windowof 0.5 ms, and those from 2100 ms to 0 ms were regarded
as a baseline. For each MEG component (e.g., P1m), a temporal peak latency and a spatial peak channel (the posterior peak of a source/sink pair) were simultaneously
searched and determined in each hemisphere of a participant. The selected peak channels did not necessarily coincide among the participants. With a software package(MEG Laboratory; Yokogawa Electric Corporation, Kanazawa-city, Japan) based on previously established procedures [Sarvas, 1987], an equivalent current dipole (ECD) atthe latency of each component was calculated for artifactfree channels corresponding to a source/sink pair of MEG
responses in each hemisphere, and a spherical model wasused to determine ECDs without the use of seed-points orother constraints. In experiments 1 and 2, the criterion for
an acceptable dipole solution was a goodness-of-ﬁt of atleast 90% for each participant, whereas in experiment 3, a
goodness-of-ﬁt of at least 85% was required. Each ECD
location was plotted as a point on individual MR images
using MRIcro software (http://www.mricro.com). Individual MR images were then normalized with SPM2 software(Welcome Department of Imaging Neuroscience, London,
UK), and the ECD points were transformed with the same
normalization parameters using the MRIcro subroutine(lesionmask.m). In the present study, we used the MNI(Montreal Neurological Institute) coordinate system. The
MNI coordinates of each ECD were plotted onto the standard brain, and their positions relative to Heschl’s sulcus
were determined on the standard brain. For the statisticalanalysis within each experiment, repeated measures analysis of variance (ANOVA) was performed. If an interaction
among several factors was signiﬁcant, further analyses
with one-way ANOVAs were performed. Regarding the
ECD location in each hemisphere, the unsigned x coordinates (more medial or lateral within the hemisphere),as well as the signed y and z coordinates, were used forcomparisons.

RESULTS
Experiment 1
The accuracies of the lyrics and melody tasks for the 11
participants were (94 6 1.0)% (mean 6 SEM) and (94 61.3)%, respectively, and these two values were not signiﬁcantly different (F(1, 10) 5 0.066, P 5 0.8). Figure 2A
shows event-related MEG responses to the reference stimuli, lyrics deviants, and melody deviants; these responses
were taken from the same posterior peak channel in eachhemisphere of one representative participant. The ﬁrst detectable peak, a P1m, was observed with the latency of 50r

ms for all types of stimuli, whereas the second peak, an
N1m, was observed with the latency of 100 ms for reference stimuli (Fig. 2A, black lines). From all 28 hemispheres, we obtained the peak latencies of the P1m
induced by reference stimuli (52 6 2.4 ms), as well as
those induced by both types of deviants (lyrics deviants,
55 6 1.6 ms; melody deviants, 56 6 1.1 ms). According toa two-way repeated measures ANOVA, neither the main
effects of task and hemisphere nor their interaction weresigniﬁcant for the peak latencies (Table I). The grand-average of the ECD locations for the P1m is shown in Figure
2B. The ECDs in the lyrics task [MNI coordinates, left: (x,y, z) 5 (247 6 3.0, 225 6 2.9, 11 6 3.1); right: (50 6 1.8,
220 6 2.6, 8.4 6 1.5)] and those in the melody task [left:(252 6 2.7, 223 6 3.1, 16 6 2.7); right: (52 6 1.9, 219 6
2.6, 13 6 2.3)] were located in the posterior region ofHeschl’s gyrus. The signiﬁcant main effects of task for the
ECD locations were observed, such that the ECDs of the
P1m in the lyrics task were signiﬁcantly superior to thosein the melody task (Table I). Neither the main effects oftask and hemisphere nor their interaction were signiﬁcant
for the ECD strength (Fig. 2C, Table I). The N1m for reference stimuli (peak latency, 111 6 4.3 ms) was not always
induced (nine hemispheres), presumably due to the rapid
succession of notes [Hari et al., 1982]. The P1m and N1m
are also known as P50m and N100m [McEvoy et al., 1994].
In contrast, prominent deviant-induced ﬁelds with thelatency of 130–150 ms were elicited by both the lyrics andmelody deviants (Fig. 2A, blue and red lines). The distribution of these magnetic ﬁelds with an inverse polarity ineach hemisphere suggests the existence of symmetrical
dipoles that point to the infero-posterior direction (Fig. 2A,
upper panel). To extract pure deviant-induced ﬁelds, theevent-related responses to reference stimuli were subtracted from the responses to deviants (Fig. 2D, blue andred lines), which resulted in a similar distribution patternof magnetic ﬁelds for this representative participant (Fig.
2D, upper panel for melody deviants). Because the reference notes include both features of syllable and pitch usedin the deviants, the subtracted MEG responses are selective
to the temporal properties of either lyrics or melody,
which are independent from particular languages, songs,
and constant acoustic features. In all 14 participants tested,these deviant-induced ﬁelds were consistently observed. In
the lyrics task, their temporal peak latencies in the left andright hemispheres, simultaneously determined with spatial
peak channels from the subtracted data, were 149 6 4.4 ms
and 146 6 3.0 ms, respectively. In the melody task, the peaklatencies in the left and right hemispheres were 132 6 4.6 ms
and 130 6 3.6 ms, respectively. The signiﬁcant main effects oftask for the peak latencies were observed, such that the peaklatency in the lyrics task was signiﬁcantly longer than that in
the melody task (Table I). We thus named the deviantsinduced ﬁelds in the lyrics and melody tasks the M140 and
M130, respectively.
To evaluate the hemispheric dominance of deviantinduced ﬁelds in the lyrics and melody tasks, we calculated

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r Lateralization in Monitoring Lyrics and Melody r

ECDs of the M140 and M130 from the subtracted data. The
ECDs of the M140 [left: (249 6 3.3, 218 6 3.0, 19 6 4);
right: (49 6 2.3, 213 6 3.4, 16 6 2.6)] and those of the
M130 [left: (250 6 2.9, 210 6 4.2, 7.8 6 2.0); right: (39 6
1.7, 26.8 6 5.4, 14 6 3.3)] were located anterior to Heschl’s
gyrus (Fig. 2E). According to a two-way repeated measures
ANOVA, the signiﬁcant main effects of task for the z coordinate were signiﬁcant, such that the ECDs of the M140
were signiﬁcantly superior to those of the M130 (Table I).
On the other hand, the main effects of task and an interaction between task and hemisphere were marginally signiﬁcant for the x coordinate. The ECDs of the M140 were signiﬁcantly more lateral to those of the M130 in the righthemisphere (F(1, 13) 5 16, P 5 0.014), while there wasno such difference in the left hemisphere (F(1, 13) 5 0.009,
P 5 0.9).
Next, we compared the ECD locations between the P1m
and M140/M130. According to a three-way repeated measures ANOVA for the y coordinate (Table I), the signiﬁcantmain effects of both component and hemisphere wereobserved. The ECDs of the M140/M130 were signiﬁcantly
more anterior to those of the P1m, while those of theM140/M130 and P1m were signiﬁcantly more anterior in
the right hemisphere to those in the left hemisphere. A signiﬁcant interaction between component and task for the x
and z coordinates, as well as that between task and hemisphere for the x coordinate, was also observed. The ECDsof the M140 were signiﬁcantly more dorsal to those of the

Figure 2.
The results of experiment 1, showing the left-dominant M140
and right-dominant M130. A: MEG responses to the lyrics deviants (blue lines), melody deviants (red lines), and reference stimuli (black lines), taken from one representative participant. The
upper panel shows the magnetic ﬁeld distribution at 130 msfrom the onset of the ﬁrst melody deviants (source: red lines;
sink: green lines). The lower left and lower right panels show
the responses at the same posterior channels in the left andright hemispheres, respectively. B: The grand-averaged ECD
locations of the P1m in the lyrics (blue circles) and melody tasks(red circles). The ECDs were superimposed on a structural
image of the standard brain (Montreal Neurological Institute)showing a horizontal slice at z 5 9. C: The ECD strength (mean
6 SE) of the P1m in the lyrics and melody tasks. D: Timecourses of the M140 (blue lines) and M130 (red lines), takenfrom one representative participant. The data were calculated bysubtracting event-related responses to reference stimuli fromthose to the ﬁrst appearing deviants. The upper panel is themagnetic ﬁeld distribution of the M130 (after subtraction) at 130
ms from the onset of the ﬁrst melody deviants. E: The grandaveraged ECD locations of the M140 (blue circles) and M130
(red circles). F: The ECD strength of the M140 and M130. The
M140 was signiﬁcantly larger in the left than in the right hemisphere (*P < 0.05). In contrast, the M130 was signiﬁcantly larger
in the right than in the left hemisphere.

r

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r Yasui et al. r

TABLE I. Repeated measures ANOVAs for the P1m and M140/M130 in experiment 1x

P1m
Task (lyrics, melody)
Hemisphere (L, R)
Task 3 Hemisphere

df

F

1, 13
1, 13
1, 13

2.2
0.23
0.39

y
P

0.2
0.6
0.5
Latency

z

F

P

0.16
3.1
0.10

0.7
0.1
0.8

P

6.5
1.4
0.001

0.024*
0.3
1

5.4
0.37
0.15

0.037*
0.6
0.2

0.71
0.28
0.17
18
1.6
1.2
0.63

0.4
0.6
0.7
0.0009**
0.2
0.3
0.4

ECD strength

Task (lyrics, melody)
Hemisphere (L, R)
Task 3 Hemisphere

1, 13
1, 13
1, 13

0.57
0.43
0.44

0.5
0.5
0.5

0.060
0.22
0.41

0.8
0.6
0.5

M140/M130
Task (lyrics, melody)
Hemisphere (L, R)
Task 3 Hemisphere

1, 13
1, 13
1, 13

3.9
3.9
4.2

0.068
0.070
0.062

1.5
1.8
0.097

0.2
0.2
0.8

LatencyF

ECD strength

Task (lyrics, melody)
Hemisphere (L, R)
Task 3 Hemisphere

1, 13
1, 13
1, 13

12
1.3
0.17

0.0043**
0.3
0.7

2.8
0.027
1.4

0.1
0.9
0.0022**Component (P1m, M140/M130)
Task (lyrics, melody)
Hemisphere (L, R)
Component 3 Task
Component 3 Hemisphere
Task 3 Hemisphere
Hemisphere 3 Task 3 Component

1, 13
1, 13
1, 13
1, 13
1, 13
1, 13
1, 13

2.8
0.2
1.4
5.6
3.1
4.9
2.8

0.1
0.7
0.3
0.034*
0.1
0.046*
0.1

14
2.0
6.2
0.67
0.001
0.19
0.006

0.0027**0.2
0.027*
0.4
1
0.7
0.9

df, degree of freedom; L, left; R, right; *, P < 0.05; **, P < 0.01.

P1m in the right hemisphere (F(1, 13) 5 5.6, P 5 0.034).
The ECDs of the M130 were marginally more ventral tothose of the P1m in the left hemisphere (F(1, 13) 5 4.3, P
5 0.058), and those of the M130 were signiﬁcantly medialin the right hemisphere (F(1, 13) 5 28, P 5 0.0001), while
there was no signiﬁcant difference for the other cases (P >0.1). Therefore, the ECD locations of the M140 and M130
were signiﬁcantly different from those of the P1m.
To evaluate the hemispheric dominance of M140 and
M130, we compared the ECD strength of these two components between hemispheres. We found a signiﬁcant interaction between task (i.e., component) and hemisphere for
the ECD strength of the M140/M130 (Fig. 2F, Table I). In
the lyrics task, the ECD strength of the M140 was signiﬁcantly larger in the left than in the right hemisphere (F(1, 13)
5 6.4, P 5 0.025), whereas that of the M130 was signiﬁcantly
larger in the right than in the left hemisphere in the melody
task (F(1, 13) 5 8.8, P 5 0.011). These results demonstrate theevidence of the left-dominant M140 in the lyrics task and theright-dominant M130 in the melody task.

Experiment 2
The latencies of M140 and M130 were within the rangeof those of the previously reported frequency MMNmr

(100–200 ms), and the deviants might be regarded as oddballs from the natural ﬂow of songs. Moreover, it has beenreported that the frequency MMNm was elicited even by
oddballs at the second position (i.e., different from the ﬁrst
stimulus) [Jääskeläinen et al., 2004]. It is thus necessary to
examine whether or not the M140 and M130 were identicalwith the frequency MMNm. For this purpose, we tested
syllable and pitch tasks with an oddball paradigm, which
did not require judgment based on the stored memory ofsongs. In the syllable task, all lyrics were replaced with aconstant syllable, whereas in the pitch task, all notes were
replaced with a constant pitch (Fig. 1C). Participants
detected the oddballs by monitoring the constant syllable
or pitch, where other features of the songs were identicalwith those tested in experiment 1. If the right dominanceobserved in the melody task in experiment 1 were due tothe characteristics of the frequency MMNm, then the rightdominance would be observed in the pitch task in experiment 2 as well, because the frequency MMNm is more
likely to be produced by the pitch oddballs.
The accuracies in the syllable and pitch tasks for the 10participants were (97 6 1.7)% and (91 6 1.6)%, respectively, and these two values were signiﬁcantly different(F(1, 9) 5 5.1, P 5 0.050). Figure 3A shows event-related
MEG responses to pitch oddballs and reference stimuli in

594 r

r Lateralization in Monitoring Lyrics and Melody r

the pitch task (Fig. 3A, red and black lines, respectively);
these responses were taken from the same posterior channel in each hemisphere of one representative participant.
At 120 ms, the oddballs at the positions of the third/fourthand ﬁfth-seventh notes elicited larger responses than thereference stimuli. The distribution of these magnetic ﬁelds
with an inverse polarity in each hemisphere suggests theexistence of symmetrical dipoles that point to the inferoposterior direction (Fig. 3A, upper panel). To extract the
MMNm, event-related responses to the reference stimuli,
which were the notes just before the ﬁrst appearing oddballs at all stimulus positions, were subtracted from theevent-related responses to the oddballs (Fig. 3B, blue andred lines), which resulted in a similar distribution patternof magnetic ﬁelds for this representative participant (Fig.
3B, upper panel for pitch oddballs).
In all 10 participants, both syllable and pitch oddballs
elicited larger magnetic responses than the reference stimuli in the left and right temporal cortices. In the syllabletask, the temporal peak latencies of the MMNm in the left
and right hemispheres, simultaneously determined withspatial peak channels from the subtracted data, were 136
6 11 ms and 124 6 12 ms, respectively. In the pitch task,
the peak latencies in the left and right hemispheres were
123 6 5.8 ms and 122 6 6.0 ms, respectively. Regarding
the peak latencies of the MMNm, neither the main effectsof the task and hemisphere nor their interaction were signiﬁcant (Table II). With these individually determined temporal peak latencies and spatial peak channels, the wave
amplitudes of MEG responses were calculated for oddballand reference stimuli, as well as for stimulus positions(ﬁrst and second halves of the ﬁve notes used for analyses), by using the data before the subtraction. In both tasks
and hemispheres, the signiﬁcant main effects of stimulus
type for the wave amplitudes were observed, such thatthe wave amplitudes for oddballs were larger than thosefor reference stimuli. On the other hand, the main effectsof stimulus position were not signiﬁcant (Table II). An
interaction between stimulus type and stimulus position

Figure 3.
The results of experiment 2, showing the MMNm in the syllable
and pitch tasks. A: MEG responses to pitch oddballs at the thirdand fourth notes (solid red lines), pitch oddballs from the ﬁfth
to seventh notes (dashed red lines), reference stimuli at the second and third notes (solid black lines), and reference stimuli
from the fourth to sixth notes (dashed black lines), all takenfrom one representative participant in the pitch task. The upper
panel shows the magnetic ﬁeld distribution at 130 ms from theonset of the pitch oddballs at the third and fourth notes. The
lower left and lower right panels show the responses at thesame posterior channels in the left and right hemispheres,
respectively. B: Time courses of the MMNm in the syllable (bluelines) and pitch tasks (red lines), taken from one representative
participant. The data were calculated by subtracting eventrelated responses to reference stimuli from those to pitch oddballs. The upper panel shows the magnetic ﬁeld distribution of
the MMNm (after subtraction) at 130 ms from the onset of theﬁrst pitch oddballs. C: The grand-averaged ECD locations of the
MMNm in the syllable (blue circles) and pitch tasks (red circles).
The ECDs were superimposed on a structural image showing ahorizontal slice at z 5 9. D: The ECD strength of the MMNmin the syllable (blue bars) and pitch tasks (red bars).

r

595 r

r Yasui et al. r

TABLE II. Repeated-measures ANOVAs for the MMNm in experiment 2
Left

Right

df

F

P

F

P

Syllable task
Stimulus type (oddball, reference)
Stimulus position (ﬁrst, second half)
Stimulus type 3 Stimulus position

1, 9
1, 9
1, 9

41
3.2
0.16

0.0001**
0.1
0.7

28
0.97
3.9

0.0005**0.4
0.08

Pitch task
Stimulus type (oddball, reference)
Stimulus position (ﬁrst, second half)
Stimulus type 3 Stimulus position

1, 9
1, 9
1, 9

40
2.3
2.2

0.0001**
0.2
0.2

51
1.5
4.8

<0.0001**
0.3
0.056

x
Task (syllable, pitch)
Hemisphere (L, R)
Task 3 Hemisphere

1, 9
1, 9
1, 9

1.5
0.010
0.003

y
0.2
0.9
1

1.4
0.51
0.47

Latency
Task (syllable, pitch)
Hemisphere (L, R)
Task 3 Hemisphere

1, 9
1, 9
1, 9

0.36
1.7
1.5

F

0.6
0.2
0.2

P

z
0.3
0.5
0.5

0.001
0.036
0.015

1
0.9
0.9

ECD strength
2.8
8.7
2.8

0.1
0.016*
0.1

Regarding the stimulus position, oddballs at the third and fourth notes correspond to the ﬁrst half of the ﬁve notes used for analyses(there were no oddballs at the ﬁrst and two notes), and those from the ﬁfth to the seventh notes correspond to the second half. Reference stimuli at the second and third notes correspond to the ﬁrst half, and those from the fourth to the sixth notes correspond to the second half (reference notes just before the ﬁrst appearing oddballs were used for analyses). *, P < 0.05; **, P < 0.01.

was marginally signiﬁcant in the right hemisphere for the
pitch task; the wave amplitudes for the oddballs on thesecond half were signiﬁcantly smaller than that on the ﬁrst
half (F(1, 9) 5 11, P 5 0.0089), whereas no signiﬁcant effectof stimulus position was observed for the reference stimuli(F(1, 9) 5 0.12, P 5 0.7). Therefore, the effect of stimulus
position cannot account for the larger wave amplitudes ofthe oddballs that appeared later than the reference stimuli;
moreover, this result was observed in the two tasks andboth hemispheres (Table II). These results conﬁrm thatthe observed MMNm was indeed due to the presence of
oddballs.
Next, we calculated ECDs of the MMNm from the subtracted data, and the grand-average of the ECD locations
for the MMNm is shown in Figure 3C. The ECDs of the
MMNm in the syllable task [left: (256 6 3.7, 221 6 3.9, 10
6 2.3); right: (56 6 3.6, 221 6 6.1, 9.1 6 2.9)] and those in
the pitch task [left: (251 6 2.6, 219 6 3.5, 9.4 6 4.6), right:(51 6 4.1, 214 6 3.4, 9.4 6 4.7)] were located in the posterior region of Heschl’s gyrus. Neither the main effects oftask and hemisphere nor their interaction were signiﬁcant
for all coordinates (Table II). Regarding the ECD strengthof the MMNm, the signiﬁcant main effects of hemisphere
were observed, showing that the ECD strength was signiﬁcantly larger in the left hemisphere than in the right hemisphere (Fig. 3D, Table II).
To compare the results of experiments 1 and 2, threeway ANOVAs of experiment (1 and 2) 3 task 3 hemisphere for the peak latency, dipole coordinates, and ECD
strength were performed (Table III). By comparing therM140/M130 and MMNm, the signiﬁcant main effects ofexperiment were observed, such that the M140/M130
showed signiﬁcantly longer peak latencies, and the ECD
locations of the M140/M130 were signiﬁcantly more
medial, anterior, and marginally dorsal to those of the
MMNm. An interaction between task and experiment, as
well as that between task and hemisphere, were signiﬁcant
for the ECD strength. The ECD strength of the MMNm in
the pitch task was signiﬁcantly smaller than that of the
M130 in the right hemisphere (F(1, 9) 5 5.7, P 5 0.026),
while there was no signiﬁcant difference between experiments 1 and 2 for the other cases (P > 0.2). These results
strongly suggest that the M140 and M130 were unique andindependent of the property of the frequency MMNm.

Experiment 3
To examine which factors modulated the M130 observedin experiment 1, we introduced newly memorized songs in
the melody task (Fig. 1D). It is possible that the M130 was
elicited by unexpected notes deviating from well-memorized songs. An alternative possibility is that the M130
was induced by perceiving unnatural pitch transitions, independent from the stored memory of songs. In experiment 3, the participants were trained with a ﬁxed
sequence of original and modiﬁed songs before the MEG
recordings, and performed the melody task based on these
newly memorized songs. If the M130 was elicited by unexpected notes deviating from well-memorized songs, aright-dominant M130 would be observed under the unex-

596 r

r Lateralization in Monitoring Lyrics and Melody r

TABLE III. ANOVAs for the comparison between the M140/M130 in experiment 1 and the MMNm in experiment 2xExperiment (1, 2)
Task (lyrics/syllable, melody/pitch)
Hemisphere (L, R)
Experiment 3 Task
Experiment 3 Hemisphere
Task 3 Hemisphere
Experiment 3 Task 3 Hemisphere

y

df

F

P

F

P

F

P

1, 88
1, 88
1, 88
1, 88
1, 88
1, 88
1, 88

9.2
5.2
1.5
<0.0012.0
1.4
1.7

0.0032**0.025*
0.2
1
0.2
0.2
0.2

5.0
3.2
1.3
0.14
0.083
0.048
0.25

0.028*
0.08
0.3
0.7
0.8
0.8
0.6

3.6
2.0
0.043
0.88
0.16
1.0
0.70

0.064
0.2
0.8
0.4
0.7
0.3
0.4

Latency
Experiment (1, 2)
Task (lyrics/syllable, melody/pitch)
Hemisphere (L, R)
Experiment 3 Task
Experiment 3 Hemisphere
Task 3 Hemisphere
Experiment 3 Task 3 Hemisphere

z

1, 88
1, 88
1, 88
1, 88
1, 88
1, 88
1, 88

9.1
7.5
1.1
1.0
0.19
0.53
0.25

ECD strength0.033*
0.0074**
0.3
0.3
0.7
0.5
0.6

0.065
0.68
2.1
7.0
1.7
11
0.60

0.8
0.4
0.1
0.0095**0.2
0.0013**
0.4*, P < 0.05; **, P < 0.01.

pected note condition (the purple column in Fig. 1D), but
not under the expected note condition (the orange
column). On the other hand, if the M130 was induced byperceiving unnatural pitch transitions, independent fromthe stored memory of songs, a right-dominant M130
would be observed for the modiﬁed songs (blue circles),but not for the original songs (green circles).
The accuracies under the expected note and unexpected
note conditions for the 10 participants were (87 6 2.2)%and (83 6 2.9)%, respectively, and these two values were
not signiﬁcantly different (F(1, 9) 5 2.3, P 5 0.2). To
extract pure deviant-induced ﬁelds, the event-related
responses to the second notes were subtracted from theresponses to the third notes. Regarding the peak latenciesof the M130, neither the main effects of the expectationand hemisphere nor their interaction were signiﬁcant (Table IV). Next, the ECDs were calculated from the subtracted data. The grand-average of the ECD locations for
the M130 is shown in Figure 4A. The ECDs of the M130
under the unexpected note condition [left: (246 6 6.2, 217
6 4.4, 19 6 5.2); right: (45 6 3.6, 29.7 6 4.4, 9.9 6 6.0)]
were located anterior to Heschl’s gyrus. The ECDs of the
M130 under the expected note condition [left: (245 6 5.7,
218 6 8.5, 6.3 6 6.3); right: (52 6 4.3, 227 6 5.4, 8.3 68.9)] were located anterior to Heschl’s gyrus in the left
hemisphere, and located in the posterior region of Heschl’s

TABLE IV. Repeated measures ANOVAs for the effect of expectation or melody on the M130 in experiment 3x

Effect of expectation
Expectation (expected note, unexpected note)
Hemisphere (L, R)
Expectation 3 Hemisphere

y

df

F

P

F

P

F

P

1, 9
1, 9
1, 9

0.56
0.15
2.4

0.5
0.7
0.2

2.9
0.007
1.1

0.1
0.9
0.3

1.0
0.30
1.6

0.3
0.6
0.2

3.0
0.25
0.47

0.1
0.6
0.5

Latency
Expectation (expected note, unexpected note)
Hemisphere (L, R)
Expectation 3 Hemisphere
Effect of melody
Melody (original, modiﬁed)
Hemisphere (L, R)
Melody 3 Hemisphere

ECD strength

1, 9
1, 9
1, 9

0.025
0.017
0.10

0.9
0.9
0.8

5.4
3.4
4.6

0.045*
0.1
0.060

1, 9
1, 9
1, 9

0.71
0.45
0.054

0.4
0.5
0.8

0.002
1.2
<0.0011
0.3
1Latency
Melody (original, modiﬁed)
Hemisphere (L, R)
Melody 3 Hemisphere

z

1, 9
1, 9
1, 9

0.035
0.043
0.50

*, P < 0.05; **, P < 0.01.

r

597 r

ECD strength
0.9
0.8
0.5

0.069
0.33
0.021

0.8
0.6
0.9

r Yasui et al. r

Figure 4.
The results of experiment 3, showing the right-dominant M130
under the unexpected note condition alone. A: The grand-averaged ECD locations of the M130 under the expected note (orange circles) and unexpected note conditions (purple circles).
The ECDs were superimposed on a structural image showing ahorizontal slice at z 5 9. B: The ECD strength of the M130
under the expected note (orange bars) and unexpected noteconditions (purple bars). Under the unexpected note condition,
the M130 was signiﬁcantly larger in the right than in the left
hemisphere. C: The grand-averaged ECD locations of the M130
for the original (green circles) and modiﬁed songs (blue circles).
D: The ECD strength of the M130 for the original (green bars)
and modiﬁed songs (blue bars).

gyrus in the right hemisphere. Neither the main effects of
expectation and hemisphere nor their interaction weresigniﬁcant for any of the coordinates (Table IV). Regarding
the ECD strength, the main effects of expectation (expected
note and unexpected note) were signiﬁcant, while an interaction between expectation and hemisphere was marginally signiﬁcant (Fig. 4B, Table IV). Under the unexpected
note condition, the ECD strength of the M130 was signiﬁcantly larger in the right than in the left hemisphere (F(1,9) 5 5.8, P 5 0.039), whereas there was no hemispheric
dominance under the expected note condition (F(1, 9) 5
0.042, P 5 0.8). In the right hemisphere, the ECD strength
was signiﬁcantly larger under the unexpected note thanunder the expected note condition (F(1, 9) 5 9.1, P 5

r0.014), whereas there was no such difference in the left
hemisphere (F(1, 9) 5 0.82, P 5 0.4).
Next, we examined whether or not the right-dominant
M130 was elicited automatically by the modiﬁed songs.
The accuracies for the original and modiﬁed songs for the
10 participants were (87 6 2.2)% and (84 6 2.6)%, respectively, and these two values were not signiﬁcantly different (F(1, 9) 5 0.94, P 5 0.4). Neither the main effects of
melody and hemisphere nor their interaction were signiﬁcant for the peak latency, any of the ECD coordinates, orthe ECD strength (Fig. 4C,D, Table IV).
To clarify whether the M130 observed under the unexpected note condition was consistent with the M130 inexperiment 1, in which the same melody task was
employed, we performed two-way ANOVAs of experiment (1 and 3) 3 hemisphere for the peak latency, dipole
coordinates, and ECD strength (Table V). We conﬁrmedthat the peak latency of the M130 were consistent between
experiments 1 and 3. Regarding the ECD locations, aninteraction between experiment and hemisphere was marginally signiﬁcant for the z coordinate alone. In the left
hemisphere, the ECDs in experiment 3 were signiﬁcantly
more superior to those in experiment 1 (F(1, 22) 5 5.3, P 5
0.031); no signiﬁcant difference was observed for the z
coordinate in the right hemisphere (F(1, 22) 5 0.34, P 50.6). We also examined whether the ECD strength of the
M130 was consistent between experiments 1 and 3.
Regarding the main effects of task (P 5 0.1), the difference
between experiments 1 and 3 for the ECD strength in theright hemisphere was 4.2 6 6.8 nAm covering zero (withthe conﬁdence level of 95%). The main effects of hemisphere were signiﬁcant for the ECD strength (P 5 0.016),
as expected from the right-dominant M130. The results of
the right-dominant M130, as well as its matched peak latency, ECD locations, and ECD strength in the right hemisphere, suggest that the M130 in experiment 1 is consistent
with that in experiment 3. Since the right-dominant M130
was clearly observed only under the unexpected note condiTABLE V. ANOVAs for the M130 under the unexpected
note condition in experiment 3 and the M130 in the
melody task in experiment 1
x

Experiment (1, 3)
Hemisphere (L, R)
Experiment 3
Hemisphere

y

df

F

P

F

P

F

P

1, 44
1, 44
1, 44

0.12
3.1
1.8

0.7
0.09
0.2

1.0
1.3
0.16

0.3
0.3
0.7

0.91
0.18
3.5

0.3
0.7
0.067

Latency
Experiment (1, 3)
Hemisphere (L, R)
Experiment 3
Hemisphere

ECD
strength

1, 44
1, 44

0.69
0.024

0.4
0.9

2.3
6.3

0.1
0.016*

1, 44

0.17

0.7

0.17

0.7

*, P < 0.05; **, P < 0.01.

598 r

z

r Lateralization in Monitoring Lyrics and Melody rtion in experiment 3, we conclude that the M130 in experiment 1 was indeed modulated by memory-related factors.

DISCUSSION
In the present MEG study, we demonstrated the hemispheric dominance of the auditory cortex that is selectively
modulated by deviant types of songs consisting of both
lyrics and melody, thereby elucidating under which conditions the lateralization of auditory processing changes. In
experiment 1 using familiar songs, we found that the ECD
strength of the M140 to the lyrics deviants was left-dominant, whereas that of the M130 to the melody deviants
was right-dominant (see Fig. 2). Moreover, the ECDs of the
M130 were located signiﬁcantly anterior to the P1m foci.
In experiment 2 using familiar songs with a constant syllable or pitch, the ECD strength of the frequency MMNm
elicited by oddballs was left-dominant (see Fig. 3). There
were signiﬁcant main effects of experiment (1 and 2) forthe peak latencies and for the ECD coordinates, indicatingthat the M140 and M130 were not the frequency MMNm.
In experiment 3 using newly memorized songs, the rightdominant M130 was observed only under the unexpected
note condition (see Fig. 4), independent of perceiving
unnatural pitch transitions (i.e., perceptual saliency). Moreover, the effect of selective attention to the melody ofsongs can also be excluded, as discussed in detail below.
The consistent right-dominance of the M130, as well as its
matched peak latency, ECD strength, and ECD coordinates
between experiments 1 and 3, suggests that the M130
observed in experiment 1 is due to unexpected notes deviating from well-memorized songs, but not due to unnatural pitch transitions. On the other hand, the left-dominant
M140 was elicited by lyrics deviants, suggesting the inﬂuence of top-down linguistic information and the memoryof the familiar songs. We thus conclude that the left-lateralized M140 and right-lateralized M130 reﬂect the expectation based on top-down information of language andmusic, respectively.
The effect of selective attention has been examined byprevious neuroimaging studies, and it is known that selective attention to auditory stimuli enhances neural
responses in the auditory cortex [Alho et al., 1999; Hashimoto et al., 2000; Johnson and Zatorre, 2005; Neelon et al.,2006]. The differential modulation of hemispheric dominance in detecting the lyrics and melody deviants revealed
in experiment 1 might also be due to selective attention toparticular components of songs, although there have beenno previous report comparing language and music components of the same stimuli. However, the frequency
MMNm observed in the pitch task was comparable
between the hemispheres (Fig. 3D), even when the selective attention to the music component was required. Moreover, the selective attention to the melody was required
under all of the conditions in experiment 3, but nevertheless the right-dominant M130 was observed only under theunexpected note condition (Fig. 4B, D). Therefore, ourrresults clearly established that memory-related components, independent of selective attention, speciﬁcally modulate the right-dominant M130.
A recent MEG study has reported an activity in thebilateral superior temporal area within 100–200 ms of auditory stimulus onset, which was observed only when participants could predict an incoming sound and the presented sound was an unexpected one [Aoyama et al.,2006]. The authors in this previous study suggested that
this component was similar to the MMNm, whereas the
ECD strength was not signiﬁcantly different between
hemispheres. Frequency MMNm responses are known tooriginate in the temporal cortex, and they are automatically elicited by oddballs even when participants are asked
to ignore the stimuli [Alho et al., 1998; Imaizumi et al.,
1998; Knösche et al., 2002; Näätänen et al., 2005]. However,the unexpected notes in the present experiments 1 and 3
were not frequency deviants, and the peak latency of theM140/M130 in experiment 1 were signiﬁcantly longer thanthat of the MMNm in experiment 2. This difference might
be due to the fact that the deviants in experiment 1 were
harder to detect than the oddballs in experiment 2, but theright dominance of the M130 conﬁrmed by experiment 3
suggests the expectation based on top-down information
rather than bottom-up features like detectability of stimuli.
Indeed, there was no cue in the acoustic features per se todiscriminate deviants from the reference stimuli in the
melody task of experiments 1 and 3. The melody deviants
could be discriminated by using expectation based on
stored memory of the melody sequence, and this process
would probably elicit the M130. These results indicate thepresence of memory-related sources in the superior temporal cortex, which are functionally distinct from the frequency MMNm.
The ECDs of the M140/M130 were located signiﬁcantly
anterior to that of the P1m located in the posterior regionof Heschl’s gyrus (Fig. 2B, E). One putative auditory
region located anterior to the human primary auditory
area (A1) has been proposed as the medial area (MA),based on the pattern of cytochrome oxidase and acetylcholinesterase activity [Rivier and Clarke, 1997]. This previous
anatomical study regarded the MA as an upstream association area, deﬁned as a unimodal association cortex receiving direct input from A1, and as an intermediate level
between A1 and the superior temporal area. One might
speculate that the ECDs of the M140/M130 are located in
the MA, which subserves intermediate auditory processing
and combines both bottom-up information from A1 andtop-down information from the higher auditory areas. Furthermore, the M140/M130 reﬂecting top-down processes
occurs at the latency of 140/130 ms, which is clearly later
than the P1m. These results suggest that top-down information starts to inﬂuence auditory processing as early as
130 ms, which may be useful in a fast identiﬁcation anddetection of stimulus changes.
The right hemispheric dominance of frontal and temporal regions in processing prosodic information has been599 r

r Yasui et al. r

indicated by both lesion studies [Nicholson et al., 2003;
Ross, 1981] and functional magnetic resonance imaging
studies [Hesling et al., 2005; Plante et al., 2002]. Prosody isoften regarded as the melodic line of speech, including thevariation in pitch, rhythm, and accent in common, whereas
such semantic or emotional modulation is independent
from other speech processing, e.g., syntax that is clearlylateralized in the brain [Sakai, 2005]. Songs also consist of
two independent components, lyrics and melody, albeit
coordinated in an artistic way. It is thus intriguing to notethat these different aspects of speech processing (syntax
vs. prosody), or of music processing (lyrics vs. melody),
correspond to the dominance of left and right brains. Our
results reported here may shed new light on the similarityand difference of language and music processing.
Previous MEG studies have reported possible top-down
inﬂuences on auditory processing of music, such that amusical training or aptitude enhanced the ECD strength of
the MMNm and N1m, as well as N19m–P30m, when auditory stimuli were presented under an ignored condition[Fujioka et al., 2004; Pantev et al., 1998; Schneider et al.,2002]. The ECD strength of the N19m–P30m, the gray matter volume of the anteromedial portion of Heschl’s gyrus,
and the musical aptitude were also correlated each other[Schneider et al., 2002]. However, it is still unknown which
processes musical experiences actually affect in the auditory cortex. In the present study, we suggest that the expectation based on stored memory provides a top-down
inﬂuence on the right-dominant M130. Our results are consistent with the previous studies, in that the skills of expectation and thinking ahead are experience-dependentand critically involved in on-line monitoring of speech andmusic, thereby quickly detecting and correcting production
errors.

ACKNOWLEDGMENTS
We thank K. Iijima and N. Saeki for their technical assistance, and S. Matsukura for her administrative assistance.

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