Neuroscience Research 52 (2005) 323–329

www.elsevier.com/locate/neures

Brain activation during music listening in individuals

with or without prior music training
Yunhee Seung c, Jeong-Sug Kyong b, Sung-Ho Woo b,
Byeong-Taek Lee a, Kyoung-Min Lee a,b,*
a
Department of Neurology, Seoul National University Hospital, 28 Yongon-Dong, Chongno-Gu, Seoul 110-744, Korea
b
Program of Cognitive Sciences, Seoul National University, 28 Yongon-Dong, Chongno-Gu, Seoul 110-744, Korea
c
Department of Music Education, Gwangju National University of Education, Korea
Received 1 October 2004; accepted 12 April 2005
Available online 6 June 2005

Abstract

The present study investigated activation during listening to music with and without a task in female musicians and non-musicians. Five
subjects with long musical training for a mean period of 19  1 years (musician group) and five subjects with no training in musical
instruments (non-musician group) were imaged in a 1.5 T scanner, while they simply listened to short segments of piano pieces (LIS), and
while they performed a distorted tune test, designed using the same pieces (DTT). A significant group effect with higher signals in the
musician group was observed in the right superior and middle temporal gyri, the right inferior frontal gyrus, and the left supramarginal gyrus.
A task effect with higher signals during DTT was observed in the left sensorimotor cortex, where the interaction between the task and group
effects was also significant. Thus, the pattern of brain activation differed depending on tasks when identical music stimuli were used, and more
importantly, comparable music tasks activated the brain differently depending on prior musical training of subjects.
# 2005 Elsevier Ireland Ltd and the Japan Neuroscience Society. All rights reserved.

Keywords: Music processing; Brain activation; Functional MRI; Musical training; Temporal cortex; Distorted tune test

1. Introduction left-sided activations were reported in a task where pitch is

attended to, relative to timbre or rhythm, during melody
An influential view that arose from neuropsychological listening (Platel et al., 1997). Though these studies aimed at
(Liegeois-Chauvel et al., 1998) and electrophysiological localizing the same pitch module in the brain (Peretz and
(Maess et al., 2001) investigations on music processing in Coltheart, 2003), they obtained different results presumably
the brain is that various aspects of the processing are due to differences in musical materials used, task conditions
subserved by modules that are individually lateralized and compared, etc.
localized (Peretz and Coltheart, 2003). However, this view While this kind of inconsistency may indicate that brain
of modular localization of music processing has not always localization of music processing is task-dependent and more
been supported consistently by studies using functional complicated than the modular localization views might
neuroimaging. A supposedly same module may be found to imply, two concerns came to our attention in comparing
be in different parts, or even on different sides of the brain, in studies in the literature: first, studies often used tasks that
different studies. For example, right-sided activations were differed in music stimuli as well as cognitive processes
observed in pitch judgments with low and high memory involved in performing the music tasks. For example,
loads during listening to melodies (Zatorre et al., 1994), but subjects heard one set of stimuli for one task, and listened to
another for another task (Mazziotta et al., 1982; Zatorre
* Corresponding author. Tel.: +82 2 2072 2985; fax: +82 2 3674 7553. et al., 1994; Platel et al., 1997; Janata et al., 2002b).
E-mail address: [email protected] (K.-M. Lee). Obviously, this created a potential ambiguity in identifying

0168-0102/$ – see front matter # 2005 Elsevier Ireland Ltd and the Japan Neuroscience Society. All rights reserved.
doi:10.1016/j.neures.2005.04.011
324 Y. Seung et al. / Neuroscience Research 52 (2005) 323–329

factors responsible for differences in the pattern of brain

activation. Second, the amount of prior music experience of
subjects differed across studies that investigated localization
of different aspects in music processing. For example, some
studies used non-musicians (Zatorre et al., 1994; Platel et al.,
1997; Koelsch et al., 2002; Doeller et al., 2003), and others
used musically experienced individuals (Mazziotta et al.,
1982; Satoh et al., 2001; Tillmann et al., 2003). This lack of
consistency in subjects’ musical experience makes it
difficult to compare studies claiming different localization
of music processes, because a variation in musical expertise
of subjects, rather than a variation in musical processes
involved in the tasks, may account for the differences.
It is possible then that these confounds, namely, co-
variation of tasks with stimuli, and co-variation of subjects’
musical experiences with tasks, contributed in some degree to
the discrepancy among studies on lateralization or localiza-
tion of music processing. In this study, we examined this
possibility by comparing brain activations: (1) between task
Fig. 1. (A) An example of a music piece is shown in original and altered
conditions that require different musical processes using the versions. Arrows indicate the altered parts. (B) A schematic depiction of
same musical materials; and (2) between individuals with and activation and rest periods during scan runs. The upper drawing is for the
without previous musical training, given identical musical music listening condition (LIS) and the lower for the distorted tune test task
materials and instructions to perform the tasks. (DTT). Only original versions were used in LIS, while altered versions were
randomly mixed with original versions in DTT. Note that comparison of
brain activation between LIS and DTT was performed only for the activation
periods with unaltered versions. DTT was twice as long as LIS in order to
2. Materials and methods match the number of compared activation periods.

2.1. Subjects abnormal. In some of the randomly chosen activation

periods, the altered versions of the pieces were played
Two groups of subjects were studied according to the (Fig. 1B). The unaltered, original versions were played in the
guidelines of the Institutional Review Board of Seoul other activation periods in DTT, and during all activation
National University Hospital and the Declaration of periods of the LIS condition. In order to ensure that brain
Helsinki. One group consisted of five graduate students activations with identical auditory stimuli were compared
majoring in music, and the other of five graduate students between LIS and DTT, we analyzed only those scans
majoring in other disciplines who had no previous training in acquired during activation periods with unaltered versions.
musical instruments. The student musicians began piano The number of activation periods to be analyzed was
instruction when they were 3–7-year-old, and practiced for a matched by having two runs of DTT and one run of LIS per
mean period of 19  1 years. All subjects were right-handed subject. The sequence of LIS and DTT was randomized
women, of similar age (average 23.8 and the range of 21–26 across subjects.
years old), normal in hearing, and without a history of
neurological disorders. Visual acuity, corrected if needed, 2.3. Music stimuli
was 20/40 or better in all subjects.
Stimuli were twenty short segments from popular piano
2.2. Behavioral tasks works by composers in the Classical and Romantic periods
(Mozart, Beethoven, Schubert, Chopin, and Schumann).
While being scanned, subjects listened to short musical Four pieces were chosen from each composer. In most cases,
pieces (Fig. 1A, and see Music Stimuli below) in two task the segments were taken from the first few phrases
conditions. In a music listening condition (LIS), no specific consisting of 10–25 notes (not counting notes in the
task requirement was given, other than to simply listen to the accompaniment).
music. Music stimuli were presented during activation For use in the DTT condition, an altered version was
periods 21 s long, which alternated with rest periods with no made for each piece in a manner similar to the Distorted
stimuli. In a distorted tune test condition (DTT), subjects Tunes Test (Drayna et al., 2001). Two to six notes per piece
were instructed to detect abnormalities in the tune while were displaced en masse either up or down by one or two
listening to the music and to press a button on an MR- semitones. The number of notes to be changed was decided
compatible keypad at the end of each activation period, by the musical tempo and the total number of notes of a
indicating whether the music they just heard was normal or piece.
 Y. Seung et al. / Neuroscience Research 52 (2005) 323–329 325

A professional pianist played the original version of the

music segments on a digital piano (Yamaha Clavinova CLP-
155). MIDI signals were recorded using a MIDI sequencing/
editing program (Cakewalk Pro Audio 9.0), which was also
used to produce altered versions. Both the original and
altered versions were then played back on the digital piano,
and saved into a Windows wave-sound format (.wav)
through a digital converter (Roland U-8 Digital Mixer).

2.4. Image acquisition and data analysis

Whole-brain scans were acquired on a 1.5 T Signa (GE,

Madison, Wisconsin). Twenty AC-PC parallel slices were
obtained with T1-weighting for an anatomy scan and with
T2*-weighting for functional scans. An interleaved EPI
Fig. 2. Brain areas that showed a significant signal increase in the task
gradient echo sequence was used with the following conditions compared to the baseline are shown for the musician group (A)
parameters: TR 3 s, TE 60 ms, flip angle 908, FOV and for non-musicians (B) ( p < 0.01, corrected for multiple comparison,
24  24 cm, matrix size 64  64 (voxel resolution of with a cluster size threshold of 30 voxels). Voxels with the LIS activation are
3.75 mm  3.75 mm), and slice thickness 5 mm with no colored green, while those with the DTT activation red. The overlap is
separation. Functional scans were aligned, smoothed, and shown in yellow, indicating an activation for both tasks. The base image is a
template T1-image provided in SPM99.
normalized onto the Talairach coordinates using SPM99
(Wellcome Department of Cognitive Neurology, University
College London). LIS condition were color-coded in green, while those
Overall patterns of activations were obtained by a fixed- activated during DTT in red. Overlaps were shown in yellow,
effect analysis on data pooled over all subjects. Statistical indicating areas that were activated in both tasks.
inference on task and group differences was then performed During LIS, musicians activated bilateral primary
with a random effect model to render the results of the auditory cortices (A1, Brodmann area 41/42) and an
inference generalizable to the population. First, a summary adjacent posterior portion of BA 22 in the superior temporal
volume was obtained for each subject and task condition by gyrus (STG), as shown in green and yellow in Fig. 2A. This
contrasting task and rest epochs, using a general linear activation extended ventrally into the middle temporal gyrus
model of SPM99. For inferences using the individual (MTG, BA21) in the right hemisphere, during DTT (shown
summary volumes, thresholds were set at the level of in red). Non-musicians showed similar activations in
p < 0.05, corrected for multiple comparisons based on the bilateral A1 and adjacent STG regions for both LIS and
Gaussian random field theory. Second, the 20 summary DTT conditions (Fig. 2B). Fig. 3 shows the degree of
volumes (two volumes per each of the 10 subjects) were individual variation of these temporal activations.
submitted to the following statistical procedures that tested Under the DTT condition, both groups showed additional
for a signal change against inter-subject variability: (1) a activations in the frontal (superior BA 44/45 and medial BA
voxel-wise t-test to see if the summary values were 8) and parietal (BA 40) cortices, as indicated by red areas in
significantly different from zero for each group (musicians Fig. 2. The frontal activations in the prefrontal regions were
or non-musicians) and task (LIS or DTT), and (2) a two-way right-lateralized, while an activation of the left sensori-
ANOVA for the main effects (musicians versus non- motor cortex was observed, obviously related to the
musicians, or LIS versus DTT) and interaction ((musicians preparation and execution of a motor response at the end
versus non-musicians) X (LIS versus DTT)). Resulting t- or of task periods in DTT.
F-score maps were thresholded at p < 0.01 (uncorrected) Difference in activity between musicians and non-
and at a cluster size of 10 or more. musicians was evaluated as a group effect in two-way
ANOVA using a random effect model. Two regions that
showed higher signals in musicians than non-musicians
3. Results stood out in this analysis (Fig. 4A, Table 1): one in the right
STG/MTG, and the other in the left supramarginal gyrus
Both groups of subjects performed the DTT task well: no (BA 40). Lower signals were observed in the musicians in
error was made in the musician group, and only two non- the posterior cingulate gyrus and the medial occipital region.
musicians made an error each. All of these regions showed a group effect because the task-
Areas that exhibited significant signal increase during related modulation of signal was in the same direction but
task periods in comparison to the baseline periods are shown stronger in musicians than in non-musicians. In contrast,
in Fig. 2A and B for musicians and non-musicians, group effects observed in the right inferior frontal gyrus and
respectively. In these figures, areas activated during the the left insular/opercular region were due to modulations of
326 Y. Seung et al. / Neuroscience Research 52 (2005) 323–329

Fig. 3. An activity map at the level of the superior and middle temporal gyri (z = +12 mm) is shown for each subject, with the t-statistic thresholded at p < 0.05,
corrected for multiple comparison, and color-coded as indicated at the bottom. Warm colors represent activation during task with respect to rest periods, and cool
colors deactivation. M1–M5 refer to individual subjects in the musician group, while N1–N5 those in the non-musician group. The upper panels are for the LIS
condition and the lower for DTT.

the opposite polarity in the two groups, namely, activation in interaction was found between the group and task effects
musicians and de-activation in non-musicians. (Table 1). A signal plot from this area indicates that the
Clusters with significant task effect were found at the interaction was such that the task effect was much stronger
following areas (Table 1): the left sensorimotor cortex and in musicians than in non-musicians, with the polarity of
the anterior cingulate gyrus, where the activity was higher modulation opposite between the two tasks (Fig. 5).
during DTT than during LIS, and the left angular gyrus (BA An additional analysis was performed comparing
39) and medial occipital regions, where it was lower during activation epochs with unaltered and altered musical pieces
DTT than during LIS. Among these areas, the left during DTT. No significant differences were observed in this
sensorimotor cortex was the only one where a significant analysis for both musicians and non-musicians.

Fig. 4. Brain areas that showed a significant group effect (musicians vs. non-musicians, in A) and task effect (LIS vs. DTT, in B) by two-way ANOVA over
twenty individual summary volumes. The probability maps were thresholded at p < 0.01 (uncorrected) and at a cluster size of 10 or more. Color-codes are
indicated below each panel. Warm colors in (A) represent higher signal in musicians than in non-musicians, and cold colors vice versa, and warm colors in (B)
represent higher activity under the LIS condition than DTT, and cold colors vice versa.
 Y. Seung et al. / Neuroscience Research 52 (2005) 323–329 327

Table 1 to consider the LIS condition not in isolation but in contrast

Areas with significant effects in voxelwise two-way ANOVA to the DTT. In the performance of the latter, attention and
kE p at maximum x y z BA working memory are obligatorily employed. For the former,
Group effect in contrast, cognitive processes involved are presumably
Musicians < 88 0.000032 2 44 46 5.23 more stimulus-driven and automatic. In this regard, the
non-musicians 21 0.001008 2 62 4 17.18
difference between LIS and DTT is similar to distinctions
10 0.000658 14 66 23 18.23
Musicians > 81 0.000152 50 39 41 2.40 often made in cognitive psychology literature, such as
non-musicians 66 0.000028 55 47 0 21.22 automatic versus controlled, attention-free versus attention-
32 0.000075 51 9 10 6.44 demanding, and bottom-up versus top-down processes.
22 0.000493 36 1 5 Insular Given the limited number of tasks that could be tested during
cortex
a scanning session, we selected tasks that were as different as
20 0.000001 48 16 15 20
14 0.000451 24 2 0 BG possible from each other. Task conditions similar to our LIS
10 0.000114 4 21 44 8.32 were also found in the literature, the ‘‘passive melodies’’
Task effect
condition of the Zatorre et al. (1994) study, the ‘‘listen
LIS < DTT 50 0.000002 50 16 46 3.4 condition’’ of the Janata et al. (2002b) study, and the
22 0.001643 7 24 34 24.32 instructions given in the Doeller et al. (2003) study.
14 0.000625 46 31 56 2.3 Previous studies have reported functional (Elbert et al.,
LIS > DTT 32 0.000689 41 70 40 19.39 1995; Pantev et al., 1998; Hirata et al., 1999; Schneider et al.,
23 0.000746 4 58 12 18.30
15 0.00064 8 32 69 4
2002) and structural (Schlaug et al., 1995; Keenan et al.,
2001; Ozturk et al., 2002; Schneider et al., 2002; Gaser and
Interaction
Schlaug, 2003) differences in the brain between individuals
20 0.000241 49 16 50 4
with musical training and those without. We have added
kE: number of voxels in a cluster. Each voxel was of 2 mm 
2 mm  2 mm; p: the probability value at a local maximum activation
several functional differences to the list, by comparing the
within each cluster; x, y, z: Talairach coordinates (in mm) of the local two groups in the same experimental setting with compar-
maximum activation in each cluster; BA: Brodmann area. able tasks. We also compared brain activity of these groups
between two levels of task demands, which allowed us to
examine not only the effects of task demands and musical
4. Discussion training separately, but also the interaction between the two
factors. However, it should be noted that our findings and the
The main aim of the current study was to compare brain following discussion may not be applicable to men because
activations of individuals with and without musical training only women subjects participated in the study and that the
while they processed identical musical materials. Since the observations need confirmation by future study before firmly
effect of music training may differ in tasks that require accepted as findings generalizable to the population, given
different levels of cognitive processing, we have instructed the small size of our sample.
subjects to perform the LIS and DTT tasks. In the LIS task, First, we found that activation of temporal cortices, i.e.
since no specific task responses were requested, it is the MTG and STG, was stronger and more extensive in
uncertain what mental processes were involved and whether musicians than in non-musicians. The difference in MTG
they were the same across subjects. However, it is important activation in particular could have been predicted by a meta-
analysis of previous studies. Activations in bilateral or
predominantly the right MTG were reported in musicians in
tonal tasks (Zatorre et al., 1998; Janata et al., 2002a),
whereas other studies found that the same area was only
weakly, if at all, activated in non-musicians (Zatorre et al.,
1994; Platel et al., 1997). Since this region is part of the
network devoted to multimodal processing (Mesulam,
2000), its stronger bilateral activation in musicians may
reflect multimodal integration of cognitive aspects in music
processing, presumably attainable only through a long-term
musical training. It is also remarkable that these areas are
close to the region where activity related to linguistic or
musical semantic processing was detected using electro-
Fig. 5. Signals at the left sensori-motor cortex, where a significant inter- encephalography (Koelsch et al., 2004). Musicians may
action between the task and group effects was found (Table 1), are plotted have accessed more musical meanings of presented stimuli
for LIS and DTT tasks for individual subjects of the musician and non-
musician groups. The signals were an average over nine neighboring voxels than non-musicians, leading to higher activity in the areas.
centered at (x: 49, y: 16, z: 50). Values on the ordinate are in an arbitrary Our study is limited, however, in characterizing the
unit. differences in this regard any further.
328 Y. Seung et al. / Neuroscience Research 52 (2005) 323–329

Second, while in non-musicians the activity in the during selective attention to one auditory stream out of two
anterior STG was enhanced during DTT compared to LIS in comparison to passive listening (Janata et al., 2002b).
(thus, red in this region in Fig. 2B), it was approximately the Thus, the group effect may have resulted from stronger
same for both tasks in musicians (thus, yellow in Fig. 2A), a activation of the area in musicians who may be more
finding consistent with a previous report (Krumhansl and proficient in attending selectively to musical materials. The
Zatorre, 2003). This difference suggests that musicians may effect might also reflect a difference between the two groups
have responded to incoming musical stimuli differently than in activating working memory system for performing the
non-musicians when no specific task was required. tasks. Gaab et al. (2003) observed significant positive
Musicians might have paid more attention to the music, correlations with the performance of a pitch memory task in
at a similar level as when they did during DTT. Or, since it is the supramarginal gyrus (mainly on the left).
highly likely that musicians were familiar with the musical Finally, the group effect observed in the right inferior
pieces prior to this study, memory retrieval and recognition frontal gyrus (BA44) may pertain to musical syntactic
might have added more activity in musicians than in non- processing by which incoming harmonic sequences are
musicians, leading to similar levels of activity in both LIS analyzed. Maess et al. (2001) found that electrical activity in
and DTT in the former group. It is also possible that in Broca’s area and its right-hemisphere homologue was
musicians, but not in musically naı̈ve individuals, even elicited by harmonically inappropriate chords occurring
passive listening to music automatically engages the within a major–minor tonal context. Since our effect was
network of relevant temporal cortices for a full analysis observed in comparison of epochs where no abnormality
of incoming musical stimuli. was present, the group difference in activity of the area likely
Third, musically-trained subjects as a group showed a reflected the difference in recruiting this region for
more balanced activation of the STG/MTG in both performing the DTT task, rather than the difference in the
hemispheres than non-musicians (Figs. 2 and 3). This activation due to actual detection of harmonic abnormality.
finding, while consistent with previous observations made in In conclusion, we confirmed that the pattern of brain
listeners with some musical training (Zatorre et al., 1998; activation changed depending on the task when identical
Tillmann et al., 2003) or non-musicians (Zatorre et al., 1994; music stimuli were used, and more importantly, that prior
Platel et al., 1997; Koelsch et al., 2002; Doeller et al., 2003), musical training of subjects influenced the activation evoked
may not be limited to musical tasks. In fact, hemispheres in by comparable tasks.
musicians’ brain are anatomically more connected to each
other, as evidenced by increased size of the corpus callosum
(Schlaug et al., 1995; Ozturk et al., 2002). Although it is Acknowledgements
unknown whether the part of the corpus callosum by which
bilateral STG/MTG are connected is in fact enlarged in We thank Dr. Eunshik Choi for preparing the musical
musicians, it is quite tempting to speculate that musical materials. Supported by Korea Science and Engineering
training activates both hemispheres in concert, so that inter- Foundation (R01-2000-000-00167-0).
hemispheric connectivity is enhanced, which may as well be
functional in the performance of tasks in other domains.
Fourth, activation of the left sensorimotor cortex showed
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