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Neuropsychologia 119 (2018) 118–127

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Neuropsychologia
journal homepage: www.elsevier.com/locate/neuropsychologia

Roles of posterior parietal and dorsal premotor cortices in relative pitch T


processing: Comparing musical intervals to lexical tones

Chen-Gia Tsaia,b, Tai-Li Choub,c,d,e, Chia-Wei Lif,
a
Graduate Institute of Musicology, National Taiwan University, Taipei, Taiwan
b
Neurobiology and Cognitive Science Center, National Taiwan University, Taipei, Taiwan
c
Department of Psychology, National Taiwan University, Taipei, Taiwan
d
Graduate Institute of Brain and Mind Sciences, National Taiwan University, Taipei, Taiwan
e
Graduate Institute of Linguistics, National Taiwan University, Taipei, Taiwan
f
Department of Radiology, Wan Fang Hospital, Taipei Medical University, Taipei, Taiwan

A R T I C LE I N FO A B S T R A C T

Keywords: Humans use time-varying pitch patterns to convey information in music and speech. Recognition of musical
Attention melodies and lexical tones relies on relative pitch (RP), the ability to identify intervals between two pitches. RP
Pitch processing in music is usually more fine-grained than that in tonal languages. In Western music, there are twelve
Lexical tone pitch categories within an octave, whereas there are only three level (non-glide) lexical tones in Taiwanese (or
Music
Taiwanese Hokkien, a tonal language). The present study aimed at comparing the neural substrates underlying
FMRI
RP processing of musical melodic intervals with that of level lexical tones in Taiwanese. Functional magnetic
resonance imaging data from fourteen participants with good RP were analyzed. The results showed that ima-
gining the sounds of visually presented musical intervals was associated with enhanced activity in the central
subregion of the right dorsal premotor cortex (dPMC), right posterior parietal cortex (PPC), and right dorsal
precuneus compared to auditory imagery of visually presented Taiwanese bi-character words with level lexical
tones. During the sound-congruence-judgement task (auditory imagery of musical intervals or bi-character
words, and subsequently judging if the imagined sounds were melodically congruent with heard sounds), the
contrast of the musical minus linguistic conditions yielded activity in the bilateral dPMC-PPC network and dorsal
precuneus, with the dPMC activated in the rostral subregion. The central dPMC and PPC may mediate the
attention-based maintenance of pitch intervals, whereas the dorsal precuneus may support attention control and
the spatial/sensorimotor processing of the fine-grained pitch structures of music. When judging the congruence
between the imagined and heard musical intervals, the bilateral rostral dPMC may play a role in attention
control, working memory, evaluation of motor activities, and monitoring mechanisms. Based on the findings of
this study and recent studies of amusia, we suggest that higher order cognitive operations are critical to the more
fine-grained pitch processing of musical melodies compared to lexical tones.

1. Introduction short musical melodies (Nan and Friederici, 2013) or melodic intervals
in music (Bradley, 2016).
One prominent feature shared between music and language is the While the global pitch perception is essential for both tonal and non-
deliberate use of pitch variation in conveying information. Parallels of tonal languages, the local pitch perception for lexical tone categoriza-
pitch perception in speech and music can be noted in the global and tion seems to play a more important role for tonal languages relative to
local pitch patterns (Nan and Friederici, 2013; Xie and Myers, 2015). non-tonal languages. This raises a question about how experience with
Sentence intonation is analogous to the global pitch contour of a mu- a tone language affects the local pitch perception for music. Deutsch
sical melody, which tracks the direction of musical pitch movement et al. (2006) hypothesized that an acquired tonal language might have a
over time (Stalinski et al., 2008). At the lexical level, local pitch var- considerable impact on the development of absolute pitch (AP), a rare
iations determine word meanings in tonal languages. The latter form of ability to identify musical notes without a reference pitch. This hy-
pitch information for lexical tone categorization may be comparable to pothesis, however, was not supported by the experimental results that


Corresponding author.
E-mail address: 799032@w.tmu.edu.tw (C.-W. Li).

https://doi.org/10.1016/j.neuropsychologia.2018.07.028
Received 22 March 2018; Received in revised form 24 July 2018; Accepted 25 July 2018
Available online 26 July 2018
0028-3932/ © 2018 Elsevier Ltd. All rights reserved.
C.-G. Tsai et al. Neuropsychologia 119 (2018) 118–127

Table 1
The five conditions in the fMRI study.
Task Stimuli Cognitive processes

Visual presentation Auditory Auditory imagery via Comparing the imagined and Fine-grained pitch
presentation subvocal rehearsal heard pitch intervals processing

Auditory-imagery task Number-pseudoword* – – – –


Bi-character word – O – –
Musical score – O – O
Sound-congruence- Bi-character word Tone pair O O –
judgement task Musical score Tone pair O O O

Note: The auditory-imagery task was to ask the participant to imagine the sound of a written number-pseudoword combination, a written bi-character Mandarin
word (in Taiwanese), or written musical scores (with numerical notations) via covertly speaking/singing. The participant was instructed to judge whether the visual
stimulus could be spoken/sung. Moreover, the sound-congruence-judgment task was to ask the participant to imagine the sounds of visually-presented musical
intervals or bi-character words. The participant was instructed to judge if the imagined sounds were melodically congruent with the auditory presentation of a tone
pair.
* Baseline condition. O The involved cognitive process(es).

Fig. 1. Examples of the stimuli for the lin-


guistic conditions, music conditions, and
baseline. There were three types of visual
presentation: Taiwanese bi-character words,
tone pairs notated using numerical notation,
and number-pseudoword combinations. In this
example of numerical notation, “5” and “1”
correspond to sol and do, respectively. A vi-
sually presented bi-character word was fol-
lowed by the sound of a tone pair, which was
either congruent or incongruent with the lex-
ical tones of the word with regard of melody. A
visually presented number pair was followed
by the sound of a tone pair, which was either
congruent or incongruent with the numerical
musical notation. We used the incongruent
tone pair with the pitch interval of an adjacent
category of the congruent tone pair. For the
interval category of Mandarin words, the pitch
interval between F#4 and B3 (seven semi-
tones) was adjacent to the pitch interval be-
tween F#4 and E4 (two semitones). For the
interval category of Western music, the pitch
interval between F#4 and B3 was adjacent to
the pitch interval between G4 and B3 (eight semitones). The fact that there was a smaller difference between the congruent and incongruent musical pitch intervals
than the difference between the congruent and incongruent linguistic pitch intervals reflects the more fine-grained pitch structures of music compared to language.

no correlation between AP and lexical tone identification was observed neuroanatomical features, musicians with RP exhibited greater cortical
in musicians (Lee and Lee, 2010; Lee et al., 2011). AP should not be thickness or gray matter concentration in the IPL, IPS, SPL, and dorsal
confused with another important musical ability, relative pitch (RP), premotor cortex (dPMC) than musicians with AP (Bermudez et al.,
which allows to identify musical pitch intervals or relations between 2009). This may be linked to RP musicians’ superior ability of tonal
several pitches. RP possessors are able to categorize pitch intervals working memory than AP musicians. It has been proposed that the
while singing and perceiving pitch intervals. When a familiar melody is processing of lexical tones was closely related to RP and working
transposed, RP possessors are able to recognize it even though the memory (Nan et al., 2010; Wong et al., 2012; Bidelman et al., 2013).
fundamental frequencies of this melody are changed. Since lexical tones During RP tasks, working memory may be involved with (1) retrieving
are mainly determined by contextual pitch cues, it is believed that RP is pitch intervals from long-term memory, (2) maintaining and/or at-
more essential than AP for comprehension of a tonal language (Zhang tending to the retrieved and/or heard pitch intervals, and (3) com-
et al., 2012, 2017; Zhang and Chen, 2016). paring these pitch intervals. The PPC and dPMC may be part of a do-
Evidence has accumulated showing that the RP processing of lexical main-general network subserving RP processing in music and speech.
tones and musical melodic intervals rely on overlapping neural cir- Although there are shared neural substrates for the RP processing of
cuitry. Several regions in the posterior parietal cortex (PPC) were in- lexical tones and musical melodic intervals, very few neuroimaging
volved in the processing of lexical tones (Klein et al., 2001; Gandour studies directly compared them. This is partially due to the fact that the
et al., 2004) and musical melodies in RP possessors (Schulze et al., majority of lexical tones are characterized by distinct patterns of gliding
2009), including the superior parietal lobule (SPL), intraparietal sulcus pitch, whereas musical pitches are usually steady. Specifically, three of
(IPS), and inferior parietal lobule (IPL). Li et al. (2010) showed in- the four Mandarin tones are characterized by pitch glides and thus not
creased activation in the right IPL for the processing of lexical tones exactly analogous to the scale tones in Western music (Lee and Lee,
compared to the processing of consonants and rhymes. Foster and 2010). To circumvent this problem, Zhang et al. (2017) used Cantonese
Zatorre (2010) found that activity within the right IPS predicted task words to examined the neural substrates of RP processing of lexical
performance in recognition of transposed melody, a task requiring lis- tones and musical melodic intervals. The reason to use Cantonese in-
teners to use pitch interval information. In a study on musicians’ stead of Mandarin was that Cantonese has three level tones with steady

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C.-G. Tsai et al. Neuropsychologia 119 (2018) 118–127

Fig. 2. Schematic description of the functional scanning runs. Each trial began with a silence and warning tone. Then, numbers or characters were presented on the
screen. For the baseline condition, musical auditory-imagery condition, and linguistic auditory-imagery condition, the participants were instructed to imagine sounds
via subvocal rehearsal according to the visual presentation. At the end of a trial the participant answered whether the visual presentation could be sung or spoken by
pressing a button. For the musical and linguistic sound-congruence-judgement conditions, the participant was instructed to covertly sing/speak the visual pre-
sentation with the heard tone pair, which was consecutively presented for three times within 1.8 s. If the visually presented musical interval or bi-character word
could be correctly sung/spoken with the heard pitch interval, the visually presented pitch interval and the heard pitch interval were judged as melodically congruent.
Otherwise, they were judged as melodically incongruent. At the end of a trial the participant answered whether the pitch interval of the visual presentation was
melodically congruent with the heard tone pair by pressing a button.

Fig. 3. Behavioral results of the correct response rate of the two sound-congruence-judgment conditions. (a) Bar graph with mean and standard error of the mean
(SEM). (b) Stripcharts of linked observations. (c) Stripcharts. The correct rate for the musical sound-congruence-judgment condition was significant lower than that of
the linguistic sound-congruence-judgment condition (* p < 0.05).

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C.-G. Tsai et al. Neuropsychologia 119 (2018) 118–127

Table 2 patterns of bi-character words with level tones are called linguistic pitch
Contrast results for the auditory-imagery task. The MNI coordinates identify the intervals.
locations of cluster maxima and local maxima (> 4 mm apart within each Compared to the RP processing for tonal languages, the RP pro-
cluster). cessing for music requires more fine-grained pitch resolution. In
X Y Z Volume Information t-value Cluster Western music, for example, there are twelve pitch categories within an
(voxels) octave, whereas there are only three level lexical tones in Taiwanese.
The perfect fourth (five semitones), augmented fourth (six semitones),
Linguistic auditory-imagery condition > Baseline
12 − 84 −2 Lingual gyrus, paraippocampal 10.24 5840
and perfect fifth (seven semiones) can be used to speak the same
gyrus Taiwanese bi-charater word. In music domain, however, they are three
10 − 76 − 14 Cerebellar lobule V-VI 8.41 different pitch intervals. The aim of this study was to identify the neural
36 − 72 − 20 Fusiform gyrus 8.02 regions showing increased activity for the more fine-grained RP pro-
18 − 92 20 Superior/middle occipital gyrus 7.47
cessing of musical pitch intervals compared to linguistic pitch intervals.
6 − 44 −2 Cerebellar lobule IV 7.05
12 − 84 34 Cuneus 6.99 While undergoing fMRI, participants were asked to imagine the sounds
12 − 72 56 Precuneus 6.32 of visually presented musical intervals or Taiwanese words. Sounds of
34 − 76 48 Superior parietal lobule, 5.25 tone pairs were presented in half of the experimental trials, in which the
angular gyrus
participants were additionally asked to judge whether the imagined
− 20 12 −2 Putamen 9.79 364
4 − 16 10 Thalamus 6.07 86
sounds were melodically congruent with the heard sounds (sound-
12 16 2 Caudate 5.21 137 congruence-judgement task). As mentioned above, the PPC-dPMC net-
6 −6 16 Lentiform nucleus 4.94 192 work may be part of a domain-general working memory network for
− 42 4 − 12 Superior temporal gyrus 8.48 804 pitch processing. It seems plausible to predict that the PPC-dPMC net-
− 54 8 14 Rolandic operculum 5.93
work may show enhanced activity in both musical and linguistic RP-
− 44 12 −8 Anterior insula 5.48
− 64 − 40 22 Supramarginal gyrus, superior 4.37 50 related tasks relative to the baseline. Moreover, the PPC-dPMC network
temporal gyrus (left) may be observed in the contrast of the musical minus linguistic con-
58 − 40 22 Supramarginal gyrus, superior 3.96 52 ditions due to its putative role in the more fine-grained pitch processing
temporal gyrus (right) of music.
38 58 20 Middle frontal gyrus (right) 6.44 514
30 64 14 Superior frontal gyrus (right) 5.47
When comparing the musical to linguistic sound-congruence-jud-
− 30 40 28 Middle frontal gyrus (left) 5.59 215 gement conditions, brain regions related to attention, evaluation of
− 32 58 24 Superior frontal gyrus (left) 3.60 subvocal rehearsal and monitoring mechanisms may be activated. The
12 − 72 56 Precuneus 6.32 155 dorsolateral prefrontal cortex (dlPFC) has been found to play a role in
− 10 − 90 − 32 Cerebellar lobule Crus I-II 5.73 89
error-monitoring (Bengtsson et al., 2009), attention control (Vohn
58 14 − 10 Superior temporal pole 5.59 416
38 6 6 Insula, inferior frontal gyrus 5.48 et al., 2007; Natale et al., 2009; Wang et al., 2010; Braga et al., 2013),
64 −4 −8 Superior/middle temporal gyrus 4.80 and working memory (Veltman et al., 2003; Manoach et al., 2004;
4 2 62 Supplementary motor area 5.08 412 Mitchell, 2007; Takahama et al., 2010; Harding et al., 2016), and is
−6 24 30 Anterior cingulate gyrus 4.95 therefore a candidate region that supports attention and working
−2 20 38 Middle cingulate gyrus 3.76
2 16 42 Superior/medial frontal gyrus 3.56
memory in the fine-grained processing of musical pitch intervals during
− 28 − 100 10 Middle occipital gyrus 4.71 194 the sound-congruence-judgement task. Recent studies have indicated
− 12 − 94 14 Cuneus 3.97 that higher order cognitive operations may be critical to the perfor-
50 6 48 Precentral gyrus 4.62 102 mance of musical RP processing tasks. Albouy et al. (2013) reported a
48 6 54 Middle frontal gyrus 4.45
correlation between the impairment in maintaining musical melodic
(dorsolateral prefrontal cortex)
54 0 48 Premotor cortex (right) 3.63 information and decreased gamma oscillations within the right dlPFC.
− 50 − 2 52 Premotor cortex (left) 4.22 51 Zhang et al. (2017) observed that impaired attention to repeated pitch
8 − 36 −8 Cerebellar lobule III 4.61 33 stimuli or effortful encoding of repeated pitch stimuli may be reflected
Musical auditory-imagery condition > Linguistic auditory-imagery condition in abnormal activation in the middle frontal gyrus and precuneus. In
30 − 52 50 Inferior parietal lobule 7.84 303 the current study, we investigated the role of higher order cognitive
50 − 32 46 Supramarginal gyrus 7.15 operations in RP processing by comparing the musical versus linguistic
48 − 48 60 Intraparietal sulcus 5.46
conditions. Compared to the RP processing for lexical tones, the RP
28 −2 54 Dorsal premotor cortex 7.53 120
28 0 66 Superior frontal gyrus 5.78 processing for music may engage higher order cognitive operations
18 − 66 52 Superior parietal lobule, dorsal 4.19 33 such as working memory and attention control for more fine-grained
precuneus pitch discrimination.

2. Materials and methods


pitches. The stimuli in Zhang et al.’s (2017) study were spoken Can-
tonese words contrasting the three level tones and musical stimuli that 2.1. Participants
were matched in pitch with these spoken Cantonese words. Although
Zhang et al. (2017) demonstrated the common and differential activa- Fifty-two individuals responded to an online advertisement of the
tion maps yielded by listening to spoken words and melodically mat- present study and completed an online RP pre-test, which was a shor-
ched musical stimuli, the participants were not instructed to explicitly tened version of the sound-congruence-judgement task in the fMRI
detect changes in the heard pitch intervals. Therefore, the neural me- experiment (see below). Eighteen of them were allowed to enter the
chanisms underlying explicit RP processing still remain unclear. To fMRI experiment because their correct response rates for the four ex-
characterize the explicit RP processing for musical melodic intervals perimental conditions were equal to or higher than 80%, but one was
and lexical tones, the present study used pairs of musical tones and not willing to participate in the fMRI experiment. The remaining 17
Taiwanese bi-character words with level tones as stimuli. Similar to participants completed the fMRI experiment. The data of one partici-
Cantonese, Taiwanese (a tonal language, also known as Taiwanese pant were discarded because of hardware glitches during the fMRI
Hokkien) has three level tones (Wu, 2000). The pitch variation patterns scans. The data of the other two participants were discarded because
of bi-character words with level tones bear resemblance to musical their correct rates were lower than 70% during the fMRI scans. The
melodic intervals. In the remaining part of this paper, pitch variation final sample consisted of 14 participants (11 female, age ranged 20–27

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Fig. 4. Contrast results for the auditory-imagery task. The threshold was set at FDR-corrected p < 0.05 with a minimum cluster size of 30 voxels. (A) The linguistic
auditory-imagery condition was associated with significant greater activation in the bilateral anterior insula, bilateral dorsal premotor cortices, bilateral caudate,
bilateral middle frontal gyrus, bilateral superior temporal gyrus, precuneus, supplementary motor area, and right angular gyrus relative to the baseline condition. (B)
Significantly increased activation in the right intraparietal sulcus, right angular gyrus, right superior parietal lobule, right precuneus, and right dorsal premotor
cortex (central subregion) was observed for the contrast of the musical minus linguistic auditory-imagery conditions.

years) with correct rates higher than 70% during the fMRI scans. None conditions. The stimuli for the baseline condition were visual pre-
of the participants reported neurological, mental, or hearing problems. sentations of number-pseudoword, which were not associated with any
This fMRI study was conducted in accordance with the Declaration of pitch interval because the lexical tones of pseudowords could not be
Helsinki, and all procedures were approved by the Internal Review determined. These five conditions involve different cognitive processes,
Board of National Taiwan University, Taiwan. Written informed con- as shown in Table 1. For the linguistic and musical auditory-imagery
sent was obtained from all participants, and written instructions were condition, working memory was involved in auditory imagery via
given to the participants before the fMRI experiment. subvocal rehearsal. For the linguistic and musical sound-congruence-
judgement conditions, working memory was involved in auditory
imagery via subvocal rehearsaland comparing the imagined pitch in-
2.2. Tasks terval with the heard pitch interval. In addition, fine-grained pitch
processing was involved in the musical auditory-imagery condition and
There were two tasks during the fMRI scans, and the participants musical sound-congruence-judgement condition.
performed one of them for each trial. The first task, auditory-imagery
task, was to attempt to imagine the sound of a visually presented
number-pseudoword combination (a number and a one-character 2.3. Stimuli and procedure
pseudoword), a bi-character Mandarin word (in Taiwanese), or musical
scores (with numerical notations) via covertly speaking/singing, and to Example stimuli are shown in Fig. 1. For the auditory-imagery task,
judge whether the visual presentation could be spoken/sung. The a visual stimulus was presented in a trial. For the sound-congruence-
second task, sound-congruence-judgement task, was to imagine the judgement task, a visual stimulus and an auditory stimulus were pre-
sounds of visually presented musical pitch intervals or bi-character sented in a trial. There were three types of visual stimuli: bi-character
words, and subsequently to judge if the imagined sounds were melo- words, musical scores with numerical notation, and number-pseudo-
dically congruent with the auditory presentation of a tone pair. In this word combinations. The auditory stimuli were tone pairs.
task, the participant attempted to covertly sing/speak the visual pre- We used numerical notation for the visual presentation of musical
sentation with the heard pitches. If the visually presented musical in- scores because numerical notation is commonplace in Taiwan. The
terval or bi-character word could be correctly sung/spoken with the numbers 1, 2, 3, ……, and 7 correspond to the solfège pitch names do,
heard pitch interval, the visually presented pitch interval and the heard re, mi, ……, and si, respectively. A musical score had two numbers that
pitch interval were judged as melodically congruent. Otherwise, they indicated a melodic interval. The auditory stimuli of tone pairs were
were judged as melodically incongruent. A mixed visual-auditory de- generated from the oboe instrument sound by Reason 7.0 (Reason
sign was used for the sound-congruence-judgement task because we Software Company Inc.). All oboe tones were within the pitch range
thought that if bi-character words were auditorily presented, the ma- from G#3 to A4 (fundamental frequency 207.7–440.0 Hz), which is
jority of participants may use fine-grained pitch processing to judge encompassed by the pitch range of human female adult speech.
whether the pitch interval of the auditorily presented word was con- For the sound-congruence-judgement task, there were congruent
gruent with the auditorily presented tone pair. and incongruent trials. A linguistic congruent trial included a visually
There were four experimental conditions and a baseline condition in presented Taiwanese word and an auditorily presented tone pair with
the present study. The four experimental conditions were linguistic the same linguistic pitch interval category. A musical congruent trial
auditory-imagery, musical auditory-imagery, linguistic sound-con- included visually presented musical scores and an auditorily presented
gruence-judgement, and musical sound-congruence-judgement tone pair with the same musical pitch interval category. For the

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Table 3 interval differed by one semitone.


Contrast results for the sound-congruence-judgment task. The MNI coordinates The experimental procedure is schematically illustrated in Fig. 2.
identify the locations of cluster maxima and local maxima (> 4 mm apart There were four runs of fMRI scanning. Each run consisted of 30 trials.
within each cluster). The total 120 trials (24 trials per condition) were presented in a unique
X Y Z Volume Information t-value Cluster random order for each participant. Each trial began with a silence (2 s)
(voxels) and warning tone (2.8 kHz, 0.7 s). Then, numbers or characters were
presented on the screen. For the baseline condition (number-pseudo-
Linguistic sound-congruence-judgment condition > Baseline
56 0 − 10 Superior temporal gyrus, 14.08 67237
word), the musical auditory-imagery condition, and the linguistic au-
supramarginal gyrus ditory-imagery condition, the participants were instructed to attempt to
− 38 18 0 Anterior insula 13.51 imagine the sounds via subvocal rehearsal three times according to the
− 58 − 26 2 Middle temporal gyrus 13.12 visual presentation. The participants answered whether the visual
52 4 − 12 Superior temporal pole 11.99
presentation could be sung or spoken by pressing a button (a left button
10 10 54 Supplementary motor area, 11.35
anterior/middle cingulate gyrus in response to “can” and a right button for “cannot”). For the musical
8 − 20 10 Thalamus 11.35 sound-congruence-judgement condition and the linguistic sound-con-
8 − 74 6 Middle/inferior occipital gyrus, 9.15 gruence-judgement condition, the participants were instructed to cov-
lingual gyrus, precuneus, cuneus
ertly sing/speak the visual presentation with the heard tone pair, which
− 34 − 60 − 30 Cerebellar lobule IV-Crus I 8.97
50 4 48 Precentral gyrus, dorsal premotor 8.85
was consecutively presented for three times within 1.8 s. This tone pair
cortex (right) was either melodically congruent or incongruent (50% probability for
− 48 16 20 Inferior frontal gyrus 8.83 each, randomly presented) with the pitch interval of the visual pre-
50 30 24 Dorsolateral prefrontal cortex 8.71 sentation. Then, the participants answered whether the pitch interval of
−6 26 46 Superior/middle/inferior frontal 8.71
the visual presentation was congruent with the pitch interval of the
gyrus
− 52 2 48 Precentral gyrus, dorsal premotor 7.66 auditory presentation by button pressing (a left button in response to
cortex (left) “no” and a right button for “yes”).
42 − 46 46 Inferior parietal lobule 7.19
− 18 − 28 − 12 Hippocampus, parahippocampal 7.17 2.4. MRI data acquisition
gyrus
24 4 6 Putamen, caudate 6.02
− 10 − 26 − 14 Pons, midbrain 6.02 For imaging data collection, participants were scanned using a 3 T
− 38 − 48 40 Inferior parietal lobule 5.68 344 MR system (MAGNETOM Prisma, Siemens, Erlangen, Germany) and a
− 46 − 40 38 Supramarginal gyrus 3.57 20-channels array head coil at the Imaging Center for Integrated Body,
− 54 − 32 38 Intraparietal sulcus 3.28
Mind, and Culture Research, National Taiwan University. In the func-
− 12 − 68 36 Precuneus 4.36 83
− 48 − 52 52 Inferior parietal lobule 3.93 62 tional scanning, about 2.5 mm slices of axial images were acquired
52 − 38 − 20 Inferior temporal gyrus 3.93 55 using a gradient echo planar imaging (EPI) with the following para-
Musical sound-congruence-judgment condition > Linguistic sound-congruence- meters: time to repetition = 2500 ms, echo time = 30 ms, flip angle
judgment condition = 87°, in-plane field of view = 192 × 192 mm, and acquisition matrix
36 − 54 54 Postcentral gyrus, superior/ 8.57 535 = 78 × 78 × 45 to cover whole cerebral areas. For spatial individual-
inferior parietal lobule (right) to-template normalization in preprocessing, a Magnetization Prepared
30 − 70 30 Middle occipital gyrus 6.44
Rapid Gradient Echo T1-weighted imaging with spatial resolution of
30 − 66 38 Superior occipital gyrus 6.12
30 − 58 42 Angular gyrus 5.15 0.9 mm isotropic was acquired for each participant.
24 − 60 30 Precuneus 4.71
− 32 − 54 52 Superior/inferior parietal lobule 5.29 82 2.5. Data analysis
(left)
12 − 68 56 Dorsal precuneus (right) 6.20 184
− 10 − 62 52 Dorsal precuneus (left) 5.40 32
The correct response rates for all five conditions during the fMRI
24 6 54 Dorsal premotor cortex (right) 6.09 131 scan session were computed. Participants with a correct rate lower than
− 24 12 54 Dorsal premotor cortex (left) 5.21 40 70% were excluded from the fMRI data analysis. We performed a one-
36 − 38 34 Intraparietal sulcus 5.44 58 tailed t-test on participants’ correct rates for the musical versus lin-
40 − 44 36 Supramarginal gyrus 5.39
guistic sound-congruence-judgement conditions because of the a priori
− 34 − 84 36 Middle occipital gyrus 5.37 37
hypothesis that the musical sound-congruence-judgement condition
required more fine-grained pitch discrimination and thus was more
linguistic sound-congruence-judgement task, the tone pairs had the difficult than the linguistic sound-congruence-judgement condition.
melodic intervals of 0, ± 2, and ± 7 semitones, which approximately For preprocessing and statistical analysis of the fMRI data, we used
correspond to the pitch intervals of Taiwanese words of three level SPM12 statistical parametric mapping software (Wellcome Trust Center
tones. Therefore, the linguistic pitch intervals had five categories. A for Neuroimaging; http://www.fil.ion.ucl.ac.uk/spm) and the Artifact
linguistic incongruent trial consisted of a visually presented Taiwanese Detection Tools (ART) toolkit. The first four volumes of each run were
word and an auditorily presented tone pair with the pitch interval ad- discarded to allow for magnetic saturation effects. The remaining
jacent to the linguistic interval category of the Taiwanese word. For the functional images were corrected for head movement artifact and
musical sound-congruence-judgement condition, the tone pairs had the timing differences in slice acquisitions. Preprocessed functional images
melodic intervals of 0, ± 1, ± 2, ± 3, ± 7, and ± 8 semitones. A were then coregistered to the individual anatomical image and nor-
musical incongruent trial consisted of a visually presented number pair malised to the standard MNI brain template and resampled to a 2-mm
and an auditorily presented tone pair, with their pitch intervals dif- isotropic voxel size. Normalised images were spatially smoothed with a
fering by one semitone. The degree of pitch-interval incongruence Gaussian kernel of 4-mm full width at half maximum (FWHM) to ac-
varied across the linguistic incongruent trials and musical incongruent commodate any anatomical variability across participants. ART toolkit
trials. For linguistic incongruent trials, the auditorily presented pitch was used to check the spikes and motion of each dataset. All functional
interval and the visually presented pitch interval differed by approxi- dataset showed head motion below 2.5 mm of maximal translation (in
mately two or five semitones. For musical incongruent trials, the au- any direction) and 1.0° of maximal rotation throughout the course of
ditorily presented pitch interval and the visually presented pitch scanning, and none outliers happened in all functional dataset.
The individual functional data analysis was performed in SPM12.

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Fig. 5. Contrast results for the sound-congruence-judgment task. The threshold was set at FDR-corrected p < 0.05 with a minimum cluster size of 30 voxels. (A) The
linguistic sound-congruence-judgment condition was associated with significant greater activation of the insula, striatum, superior/middle temporal gyrus, inferior
frontal gyrus, dorsal premotor cortex, supplementary motor area, caudate, and inferior parietal lobule relative to the baseline condition. (B) Significantly increased
activation in the superior/inferior parietal lobules, bilateral dorsal premotor cortex (rostral subregion), and right precuneus was observed for the contrast of the
musical minus linguistic sound-congruence-judgment conditions.

Statistical inference was based on a random effect approach at two the contrast of the linguistic sound-congruence-judgement condition
levels. At the single-subject level, the data of each participant were minus baseline and the contrast of the musical minus linguistic sound-
analyzed using the general linear model by fitting the time series data congruence-judgement conditions. When comparing the linguistic
with the canonical hemodynamic response function (HRF) modeled at sound-congruence-judgement condition to baseline condition, greater
events (Josephs and Henson, 1999). The events were modeled for the activation was observed within the right superior/middle temporal
response period in each of the five conditions. In the group-level ana- gyri, superior/inferior parietal lobules, dlPFC, IFG, SMA, dPMC, ante-
lysis, paired t-tests were used to calculate the contrasts of the linguistic rior insula, anterior cingulate cortex, precuneus, striatum, cerebellum,
auditory-imagery and linguistic sound-congruence-judgement condi- and thalamus/midbrain. Moreover, the musical sound-congruence-
tions minus the baseline condition, as well as the contrasts of the mu- judgement condition was associated with greater activation in the
sical minus linguistic auditory-imagery conditions and the musical dPMC, superior/inferior parietal lobules, and dorsal precuneus com-
minus linguistic sound-congruence-judgement conditions. The statis- pared to the linguistic sound-congruence-judgement condition.
tical significant threshold was set at p < 0.05 (FDR-corrected) and a
minimum cluster size of 30 voxels in a whole brain analysis.
4. Discussion

3. Results Perception of lexical tones in a tonal language and perception of


musical melodies rely on parsing the relationship between pitches,
Fig. 3 shows the results of the correct response rates for the two namely, RP processing. In the current study, we used fMRI and working
sound-congruence-judgement conditions. Participants had significantly memory paradigms to compare the neural substrates underlying the
lower correct rates in the musical sound-congruence-judgement con- explicit RP processing of musical and linguistic pitch intervals. The
dition compared to the linguistic sound-congruence-judgement condi- linguistic stimuli were Taiwanese bi-character words with level tones.
tion (p = 0.0317, one-tailed t-test). This result suggests that the musical Their steady pitches allow for a direct comparison to the scale tones in
sound-congruence-judgement task was more difficult than the linguistic Western music. Pitch intervals of Taiwanese words are less fine-grained
sound-congruence-judgement task. compared to musical pitch intervals. Analysis of the behavioral data
Table 2 and Fig. 4 show the significantly activated brain regions for confirmed that the musical sound-congruence-judgement task was more
the contrast of the linguistic auditory-imagery condition minus baseline difficult than the linguistic sound-congruence-judgement task. As ex-
and the contrast of the musical minus linguistic auditory-imagery pected, we found that several brain regions implicated in higher order
conditions. When comparing the linguistic auditory-imagery condition cognitive operations such as working memory and attention control
to baseline condition, greater activation was observed within the su- showed enhanced activity for the musical versus linguistic conditions.
perior temporal gyrus, inferior/superior parietal lobules, precuneus, All the four experimental conditions were associated with sig-
inferior frontal gyrus (IFG), anterior insula, dPMC, dlPFC, supplemen- nificantly greater activity within the dPMC and PPC relative to the
tary motor cortex (SMA), lingual gyrus, cuneus, parahippocampal baseline. This finding is concordant with the results from prior fMRI
gyrus, striatum, thalamus, and cerebellum. Moreover, the musical au- studies showing the involvement of the dPMC-PPC network in both
ditory-imagery condition was associated with greater activation in the linguistic and musical RP processing (Klein et al., 2001; Gandour et al.,
right dPMC, right superior/inferior parietal lobules, and precuneus 2004; Bermudez et al., 2009; Schulze et al., 2009; Foster and Zatorre,
compared to the linguistic auditory-imagery condition. 2010; Li et al., 2010; Angenstein et al., 2012; Brown et al., 2013). The
Table 3 and Fig. 5 show the significantly activated brain regions for dPMC-PPC network has been termed the dorsal attention network

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because it mediates the top-down guided voluntary allocation of at- processing salient sounds (Watkins et al., 2007), effortful auditory
tention to stimuli features, and links stimuli to appropriate responses imagery (Huijbers et al., 2011), and shifts of auditory attention
(Corbetta et al., 2008). Recently, Genon et al. (2016) employed con- (Shomstein and Yantis, 2006). In an fMRI study of music score reading
nectivity-based parcellation to divide the right dPMC into the central, and playing, Stewart et al. (2003) showed increased dorsal precuneus
ventral, dorsal, rostral, and caudal subregions. The central subregion of activation when participants used the right hand to play melodies from
the right dPMC tends to work in tandem with the PPC to support musical notation after training, suggesting the role of this area in
working memory maintenance (Genon et al., 2016). In the present translating musical notation into sensorimotor codes. Moreover,
study, the right PPC and the right central dPMC showed increased ac- Meister et al. (2005) found that pianists’ dorsal precuneus exhibited
tivation for the musical versus linguistic auditory-imagery conditions. increased activation during silently playing musical scores with the
This result is consistent with Schon et al.’s (2010) finding of increased right hand, relative to passively reading the score of repetition of a
activity in the right central dPMC for a melody discrimination task than single note. In the present study, the musical conditions entailed the
a speech discrimination task. We suggest that the right dPMC-PPC transformation from musical notations into laryngeal sensorimotor
network may be responsible for the voluntary allocation of attention to codes for subvocal rehearsal. During the auditory-imagery and sound-
pitch features of the stimuli and the attention-based maintenance of congruence-judgement tasks, increased activation in the right dorsal
working memory (Berryhill et al., 2011) for the musical auditory-ima- precuneus for the musical versus linguistic conditions may reflect its
gery, musical sound-congruence-judgement, and linguistic sound-con- role in attention control and the spatial/sensorimotor processing of the
gruence-judgement conditions. The activation clusters in the dPMC and more fine-grained pitch structures of music compared to lexical tones.
PPC for the linguistic auditory-imagery condition were more limited. While the present fMRI study recruited individuals with good RP,
This may be due to the relatively lower attention to pitch features of the our findings also echo several studies on congenital amusia, a disorder
imagined Taiwanese words. affecting the processing of musical pitch. Past studies have demon-
Experimental and theoretical studies have suggested that the right strated that the impaired fine-grained processing of pitch in congenital
PPC contains the neural substrate of a general magnitude system for amusics seems to be characterized by reduced activity in the right IFG
space, time, and numbers (Walsh, 2003; Cohen Kadosh et al., 2007; in response to musical melodies (Hyde et al., 2011), reduced white
Bueti and Walsh, 2009). This general magnitude system may support matter concentration in the right IFG (Hyde et al., 2006), as well as the
accurate representation and processing of pitch relations for the task of reduced connectivity between the right IFG and auditory cortex (Loui
judging the melodic congruence between an imagined musical pitch et al., 2009; Peretz, 2016). In the present study, both the contrast of the
interval and a heard interval. Notably, there are twelve pitch categories musical minus linguistic auditory-imagery conditions and the contrast
within an octave, whereas there are only three level lexical tones in of the musical minus linguistic sound-congruence-judgement conditions
Taiwanese. The significantly enhanced activity in the right PPC for the showed subthreshold (uncorrected p < 0.003 with cluster size > 50
musical versus linguistic auditory-imagery conditions may reflect the voxels for overall AlphaSim corrected p < 0.05) activation in the right
increased cognitive load for the more fine-grained pitch processing. IFG (extending into the ventral premotor cortex). Based on this and
Moreover, it was proposed that the right dPMC-PPC network may previous studies, we suggest that the right vPMC/IFG may contribute to
contribute to systematically transforming and comparing pitch in- sensorimotor and auditory encoding of musical pitches.
formation in a manner similar to visuospatial information (Zarate and Besides deficits in sensorimotor and auditory encoding, there is
Zatorre, 2008; Foster et al., 2013). Converging with this view, our emerging evidence linking amusia with higher order cognitive control
findings indicated that the visuospatial working memory function of the deficits. Zhang et al. (2017) recruited Cantonese-speaking amusics and
right dPMC-PPC network plays a more important role in discrimination musically intact controls to investigate the processing of pitch intervals,
of musical pitch intervals compared to linguistic pitch intervals. finding abnormally strong activation in amusics’ precuneus during the
One crucial question arising from the present study is the functional repetition of pitch stimuli. This result resonates with our finding of
specialization of the dPMC during RP processing. We observed that the increased activation in the precuneus for the music minus linguistic
musical sound-congruence-judgement condition was associated with conditions. In addition to the precuneus, the dorsal prefrontal cortex
significantly greater activity within the rostral (y > 3) dPMC than the might also plays a role in cognitive control functions for the fine-
linguistic sound-congruence-judgement condition. Compared with the grained RP processing of music. Albouy et al. (2013) reported that the
right central dPMC, the right rostral dPMC typically supports higher impairment in maintaining musical melodic information in amusics
cognitive functions, including error-monitoring (Bengtsson et al., might be linked to decreased gamma oscillations within the right dlPFC
2009), evaluating hand motor capability (Hirose et al., 2010), response during the retention of musical melodic information. A follow-up study
selection/inhibition (Rowe et al., 2008; Batterink et al., 2010; Padmala by Schaal et al. (2015) showed that modulating the right dlPFC with
and Pessoa, 2010; Cai et al., 2014), attention control (Vohn et al., 2007; 35 Hz transcranial alternating current stimulation in amusics selectively
Natale et al., 2009; Wang et al., 2010; Braga et al., 2013), mental cal- improved pitch memory performance. The critical role of this region in
culation (Fehr et al., 2008), and working memory (Veltman et al., 2003; the processing of musical pitch intervals is supported by our finding
Manoach et al., 2004; Mitchell, 2007; Takahama et al., 2010; Harding that the rostral dPMC, a transitional zone between the dPMC and dlPFC,
et al., 2016). It is also recognized that the left rostral dPMC may showed greater activity for the musical versus linguistic sound-con-
mediate error-monitoring (Chen et al., 2006; Provost et al., 2010) and gruence-judgement conditions.
top-down attention to memory retrieval (Ciaramelli et al., 2010). We Several limitations of the study need to be pointed out. First, the
suggest that the bilateral rostral dPMC may support attention control, final sample of fMRI data consisted of 11 females and 3 males. Future
working memory, evaluation of motor activities, and monitoring me- research should recruit a larger sample size with a good gender balance.
chanisms during judging the congruence between the imagined and Second, the major screening criterion to be able to participate in the
heard musical pitch intervals. fMRI experiment was the score of a RP pre-test, which was a shortened
In the present study, the contrast of musical minus linguistic sound- version of the congruence-sound-judgement task in the fMRI experi-
congruence-judgement conditions yielded co-activation of the bilateral ment. While this pre-test selected participants with good RP, our study
rostral dPMC and dorsal precuneus. This result accords well with Genon did not take into account the effect of music training. Moreover, future
et al.’s (2016) finding that the right rostral dPMC connects with the research could explore whether RP-related musical training is beneficial
precuneus. In visual modality, the dorsal precuneus has been implicated for sensorimotor/spatial processing, working memory, attention con-
in higher order cognitive operations such as spatial working memory trol, or error-monitoring.
and attention control (Lepsien et al., 2005; Natale et al., 2006). In au- To summarize, the present study has examined the neural substrates
ditory modality, the dorsal precuneus plays a role in detecting/ underlying the explicit RP processing of musical and linguistic pitch

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ment of the dPMC-PPC network in RP-related tasks (Klein et al., 2001; Foster, N.E., Zatorre, R.J., 2010. A role for the intraparietal sulcus in transforming mu-
sical pitch information. Cereb. Cortex 20, 1350–1359.
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and Zatorre, 2010; Li et al., 2010; Schon et al., 2010), linking RP with 2004. Hemispheric roles in the perception of speech prosody. NeuroImage 23,
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