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Hemispheric Roles in The Perception of Speech Prosody PDF
Hemispheric Roles in The Perception of Speech Prosody PDF
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NeuroImage 23 (2004) 344 – 357
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Hemispheric roles in the perception of speech prosody
Jackson Gandour, a,* Yunxia Tong, a Donald Wong, b Thomas Talavage, c Mario Dzemidzic, d
Yisheng Xu, a Xiaojian Li, e and Mark Lowe f
a
Department of Audiology and Speech Sciences, Purdue University, West Lafayette, IN 47907-2038, USA
b
Department of Anatomy and Cell Biology, Indiana University School of Medicine, IN 46202-5120, USA
c
School of Electrical and Computer Engineering, Purdue University, IN 47907-2035, USA
d
MDZ Consulting Inc., Greenwood, IN 46143, USA
e
South China Normal University, Guangzhou, PR China
f
Cleveland Clinic Foundation, Cleveland, OH 44195, USA
1053-8119/$ - see front matter D 2004 Elsevier Inc. All rights reserved.
doi:10.1016/j.neuroimage.2004.06.004
J. Gandour et al. / NeuroImage 23 (2004) 344–357 345
features, on a time scale of 200 – 350 ms. Intonation, on the other syllable utterances, 39% of the pairs for the three-syllable utter-
hand, is manifested at the phrase or sentence level, typically on a ances. Stimuli that were identical in both tone and intonation
time scale of seconds. In Chinese, interrogative intonation exhibits a comprised 28% and 22% of the pairs in one-syllable and three-
higher pitch contour than that of its declarative counterpart (Shen, syllable utterances, respectively.
1990) as well as a wider pitch range for sentence-final tones (Yuan
et al., 2002). In English, interrogative sentences do not have overall Recording procedure
higher pitch contours than declarative sentences, nor do they show
any effects of tone and intonation interaction in sentence-final A 52-year-old male native speaker of Mandarin was instructed
position. Chinese interrogative intonation with a final rising tone to read one- and three-syllable utterances at a conversational
has a rising end, which is similar to English, whereas that with a speaking rate in a declarative and interrogative sentence mood. A
final falling tone often has a falling end (Yuan et al., 2002). reading task was chosen to maximize the likelihood of simulating
In a previous fMRI study of Chinese tone and intonation (Gan- normal speaking conditions as much as possible while at the same
dour et al., 2003), both tone and intonation were judged in sentences time controlling the syntactic, prosodic, and segmental character-
presented at a fixed length (three words), and we observed left- istics of the spoken sentences. To enhance the naturalness of
lateralized lexical tone perception in comparison to intonation. producing the three-syllable utterances, he was told to treat them
However, the prosodic unit listeners selectively attended to and as SVO (subject verb object) sentences with non-emphatic stress
the temporal interval of attentional focus were coterminous. In placed on the final syllable. All items in the list were typed in
judgments of lexical tone, the focus of attention was on the final Chinese characters. A sufficient pause was provided between items
word only, whereas judgments of intonation required that the focus to ensure that the speaker maintained a uniform speaking rate. By
be directed to the entire sentence. Whether the principal driving controlling the pace of presentation, we maximized the likelihood
force in hemispheric lateralization of speech prosody is due to the of obtaining consistent, natural-sounding productions. To avoid
temporal interval of attentional focus rather than the hierarchical list-reading effects, extra items were placed at the top and bottom
level of linguistic units is not yet well-established. The aim of the of the list. Recordings were made in a double-walled soundproof
present study is to determine whether the temporal interval in which booth using an AKG C410 headset type microphone and a Sony
prosodic units are presented influences the neural substrates used in TCD-D8 digital audio tape recorder. The subject was seated and
prosodic processing. As such, participants are asked to make per- wore a custom-made headband that maintained the microphone at a
ceptual judgments of tone and intonation in one-syllable and three- distance of 12 cm from the lips.
syllable Chinese utterances. By comparing activation in homolo-
gous regions of both hemispheres, we can assess the extent to which Prescreening identification procedure
hemispheric laterality for speech prosody is driven by the temporal
interval, prosodic unit, or both. Only native Chinese speakers All one- and three-syllable utterances were presented individ-
possess implicit knowledge that relates external auditory cues to ually in random order for identification by five native speakers of
internal representations of tone and intonation. By employing two Chinese who were naive to the purposes of the experiment. They
language groups, one consisting of Chinese speakers, the other of were asked to respond whether they heard a declarative or
English speakers, we are able to determine whether activation of interrogative intonation and to indicate the tone occurring on the
particular brain areas is sensitive to language experience. final syllable. Only those stimuli that achieved a perfect (100%)
recognition score for both intonation and tone were retained for
possible use as stimuli in our training and experimental sessions.
Materials and methods
Task procedure
Subjects
The experimental paradigm consisted of four active tasks (Table
Ten native speakers of Mandarin (five male; five female) and 1) and a passive listening task. The active tasks required discrim-
ten native speakers of American English (five male; five female) ination judgments of intonation (I) and tone (T) in paired three-
were closely matched in age/years of education (Chinese: M = 29/ syllable (I3, T3) and one-syllable (I1, T1) utterances. Subjects were
19; English: M = 27/19). All subjects were strongly right-handed instructed to focus their attention on either the utterance-level
(Oldfield, 1971) and exhibited normal hearing sensitivity. All intonation or the lexical tone of the final syllable, make discrim-
subjects gave informed consent in compliance with a protocol ination judgments, and respond by pressing a mouse button (left =
approved by the Institutional Review Board of Indiana University same; right = different). The control task involved passive listening
Purdue University Indianapolis and Clarian Health. to the same utterances, either one-syllable utterances (L1) or three-
syllable utterances (L3). Subjects responded by alternately pressing
Stimuli the left and right mouse button after each trial.
A scanning sequence consisted of two tasks presented in a
Stimuli consisted of 36 pairs of three-syllable Chinese utter- blocked format alternating with rest periods (Fig. 2). The one-
ances, and 44 pairs of one-syllable Chinese utterances. Utterances syllable and three-syllable utterance blocks contained 11 and 9
were designed with two intonation patterns (declarative, interrog- trials, respectively. The order of scanning runs and trials within
ative) in combination with the four Chinese tones on the utterance- blocks were randomized for each subject. Instructions were deliv-
final syllable (Fig. 1). Focus was held constant on the utterance- ered to subjects in their native language via headphones during rest
final syllable. No adjacent syllables in the three-syllable utterances periods immediately preceding each task: ‘‘listen’’ for passive
formed bisyllabic words to minimize lexical-semantic processing. listening to speech stimuli, ‘‘intonation’’ for same – different judg-
Tone or intonation each differed in 36% of the pairs for the one- ments on Chinese intonation, and ‘‘tone’’ for same – different
346 J. Gandour et al. / NeuroImage 23 (2004) 344–357
Fig. 1. Acoustic features of sample Chinese speech stimuli. Broad-band spectrograms (SPG: 0 – 8 kHz) and voice fundamental frequency contours (F0: 0 – 400
Hz) are displayed for utterance pairs consisting of same tone/different intonation in three-syllable utterances (top left), same tone/different intonation in one-
syllable utterances (top right), different tone/same intonation in three-syllable utterances (bottom left), and different tone/same intonation in one-syllable
utterances (bottom right).
judgments on Chinese tone. Average trial duration was about 2.9 All speech stimuli were digitally edited to have equal maximum
and 3.5 s, respectively, for the one-syllable and three-syllable energy level in dB SPL. Auditory stimuli were presented binaurally
utterance blocks, including a response interval of 2 s. using a computer playback system (E-Prime) and a pneumatic-
Table 1
Samples of Chinese tone and intonation stimuli for tasks involving one-syllable and three-syllable utterances
Note. I1 (T1) and I3 (T3) represent intonation (tone) tasks in one-syllable and three-syllable utterances, respectively.
J. Gandour et al. / NeuroImage 23 (2004) 344–357 347
Fig. 2. Sequence and timing of conditions in each of the four functional imaging runs. I3 and I1 stand for intonation in three-syllable and one-syllable Chinese
utterances, respectively; T3 and T1 stand for tone in three-syllable and one-syllable Chinese utterances, respectively; R = rest interval; L3 and L1 stand for
passive listening to three-syllable and one-syllable Chinese utterances, respectively.
based audio system (Avotec). The plastic sound conduction tubes Imaging analysis
were threaded through tightly occlusive foam eartips inside the
earmuffs that attenuated the average sound pressure level of the Image analysis was conducted using the AFNI software pack-
continuous scanner noise by f30 dB. Average intensity of all age (Cox, 1996). All data for a given subject were motion-
experimental stimuli was 92 dB SPL as compared to 80 dB SPL corrected to the fourth acquired volume of the first functional
scanner noise. imaging run. To remove differences in global intensity between
Accuracy, reaction time, and subjective ratings of task difficulty runs, the signal in each voxel was detrended across each functional
were used to measure task performance. Each task was self-rated by scan to remove scanner signal drift, and then normalized to its
listeners on a 1- to 5-point graded scale of difficulty (1 = easy, 3 = mean intensity. Each of the four functional runs was analyzed to
medium, 5 = hard) at the end of the scanning session. Before obtain cross-correlation for each of three reference waveforms with
scanning, subjects were trained to a high level of accuracy using the measured fMRI time series for each voxel. The first reference
stimuli different from those presented during the scanning runs: I3 waveform corresponded to one of the four active conditions (I1, I3,
(Chinese, 93% correct; English, 88%); I1 (Chinese, 92%; English, T1, T3) presented in a single run (Fig. 2). The second and third
77%); T3 (Chinese, 99%; English, 82%); T1 (Chinese, 99%; reference waveforms corresponded to the two control conditions,
English, 85%). L1 and L3, respectively, presented during the two runs with the
same temporal interval for the intonation and tone conditions (L1
Imaging protocol for I1 and T1; L3 for I3 and T3). After the resulting EPI volumes
were transformed to 1-mm isotropic voxels in Talairach coordinate
Scanning was performed on a 1.5T Signa GE LX Horizon space (Talairach and Tournoux, 1988), the correlation coefficients
scanner (Waukesha, WI) equipped with birdcage transmit – re- were converted to z scores for purposes of analyzing multisubject
ceive radiofrequency head coils. Each of four 200-volume echo- fMRI data (Bosch, 2000), and spatially smoothed by a 5.2-mm
planar imaging (EPI) series was begun with a rest interval FWHM Gaussian filter to account for intersubject variation in brain
consisting of 8 baseline volumes (16 s), followed by 184 anatomy and to enhance the signal-to-noise ratio.
volumes during which the two comparison tasks (32 s) alternated Direct comparison of active conditions (I1, I3, T1, T3) across
with intervening 16 s rest intervals, and ended with a rest interval runs was accomplished by computing the average z score for each
of 8 baseline volumes (16 s) (Fig. 2). Functional data were of the four active conditions relative to its corresponding control
acquired using a gradient-echo EPI pulse sequence with the condition. Averaged z scores for the control conditions were then
following parameters: repetition time (TR) 2 s; echo time (TE) subtracted from those obtained for their corresponding intonation
50 ms; matrix 64 64; flip angle (FA) 90j; field of view (FOV) or tone conditions (e.g., DzI1 = zI1 zL1, DzI3 = zI3 zL3).
24 24 cm. Fifteen 7.5-mm-thick, contiguous axial slices were Evaluating each active condition to a control of the same temporal
used to image the entire cerebrum. Before functional imaging interval also makes it possible to compare active conditions across
runs, high-resolution, and anatomic images were acquired in 124 temporal intervals (e.g., DzI1 vs. DzI3).
contiguous axial slices using a 3D Spoiled-Grass (3D SPGR) Within- and between-group random effects maps (I1 vs. L1, T1
sequence (slice thickness 1.2 – 1.3 mm; TR 35 ms; TE 8 ms; 1 vs. L1, I3 vs. L3, T3 vs. L3) were also generated for display purposes
excitation; FA 30j; matrix 256 128; FOV 24 24 cm) for by applying voxel-wise ANOVAs on the z (e.g., Chinese zI1 vs.
purposes of anatomic localization and coregistration to a standard Chinese zL1) and Dz (e.g., Chinese DzI1 vs. English DzI1) values,
stereotactic system (Talairach and Tournoux, 1988). Subjects respectively. The individual voxel threshold for between-group
were scanned with eyes closed and room lights dimmed. The maps was set at P = 0.01. For within-group maps, significantly
effects of head motion were minimized by using a head – neck activated voxels ( P < 0.001) located within a radius of 7.6 mm were
pad and dental bite bar. grouped into clusters, with a minimum cluster size threshold
348 J. Gandour et al. / NeuroImage 23 (2004) 344–357
Fig. 3. Location of fixed spherical ROIs in frontal (open circle), parietal (checkered circle), and temporal (barred circle) regions displayed in left sagittal
sections (top and middle panels), and on the lateral surface of both hemispheres (bottom panels). LH = left hemisphere; RH = right hemisphere. Stereotactic x
coordinates that appear in the top and middle panels are derived from the human brain atlas of Talairach and Tournoux (1988). See also Table 2.
ROIs relative to the Chinese group (Fig. 6; Table 4). In the frontal pSTG (Fig. 4i) was more active bilaterally across tasks (I1, I3, T1,
lobe, all four ROIs (Figs. 4a – d) were more active bilaterally for the T3), whereas greater activity in the aSTG (Fig. 4g) was observed
tone tasks (T1, T3). In the parietal lobe, IPS (Fig. 4e) activity was across tasks in the RH only.
greater in both the LH and RH for T1. In the temporal lobe, the
Within group comparisons
Table 3
Behavioral performance and self-ratings of task difficulty Hemisphere effects for the Chinese group revealed complemen-
a tary leftward and rightward asymmetries, as measured by Dz,
Language group Task Accuracy (%) Reaction time (ms) Difficulty
depending on ROI and task (Table 5). Laterality differences
Chinese I1 91.2 (1.5) 682 (48) 3.3 (0.4) favored the LH in the frontal aMFG (Figs. 4a and 7, upper panel)
T1 97.3 (0.9) 504 (41) 1.4 (0.16)
for intonation tasks only, irrespective of temporal interval (I1, I3).
I3 93.9 (1.3) 559 (36) 2.7 (0.3)
In the parietal lobe, significantly more activity was observed in the
T3 96.9 (1.2) 485 (26) 1.8 (0.25)
English I1 76.8 (2.5) 668 (49) 4.0 (0.30) left IPL (Figs. 4f and 5) across tasks, and in the left IPS (Figs. 4e
T1 70.9 (2.9) 642 (53) 3.3 (0.37) and 7, lower panel) for T3 (cf. Gandour et al., 2003). In the
I3 85.3 (2.4) 565 (40) 3.1 (0.28) temporal lobe, activity was greater in the left pSTG (Figs. 4i and 8)
T3 72.2 (3.0) 656 (47) 3.4 (0.27) and aSTG (Fig. 4g) across tasks regardless of temporal interval. In
Note. Values are expressed as mean and standard error (in parentheses). See contrast, laterality differences favored the RH in the frontal mMFG
also note in Table 1. (Fig. 4b) and temporal mSTS (Figs. 4h and 8) across tasks.
a Hemisphere effects for the English group were restricted to
Scalar units are from 1 to 5 (1 = easy; 3 = medium; 5 = hard) for self-
ratings of task difficulty. frontal and temporal ROIs in the RH (Table 5). Rightward asymme-
350 J. Gandour et al. / NeuroImage 23 (2004) 344–357
Fig. 4. Comparison of mean Dz scores between language groups (Chinese, English) per task (I1, T1, I3, T3) and hemisphere (LH, RH) within each ROI. Frontal
lobe, a – d; parietal, e – f; temporal, g – i. I1 is measured by DzI1; T1 by DzT1; I3 by DzI3; T3 by DzT3. Error bars represent F1 SE.
tries were observed in the frontal mMFG (Fig. 4b) and temporal
mSTS (Fig. 4h) across tasks. These functional asymmetries favoring
the RH were identical to those for the Chinese group. No significant
leftward asymmetries were observed for any task across ROIs.
Task effects for the Chinese group revealed laterality differ-
ences, as measured by Dz, related to the prosodic unit. Intonation
(I1, I3), when compared to tone (T1, T3), favored the LH in the
aMFG (Figs. 4a and 7). In the pMFG (Fig. 4c), I3 was greater than
T3 in the RH; I1 was greater than T1 in both hemispheres.
For both groups, a cluster analysis revealed significant (P <
.001) activation in the supplementary motor area across tasks. The
Chinese group showed predominantly right-sided activation in the
lateral cerebellum across tasks. In the caudate and thalamus,
increased activation was observed in the Chinese group for the
intonation tasks only (I1, I3), but across tasks in the English group.
Fig. 5. A random effects fMRI activation map obtained from comparison of
discrimination judgments of intonation in one-syllable utterances (I1)
relative to passive listening to the same stimuli (L1) between the two Discussion
language groups (DzI1 Chinese vs. DzI1 English). Left/right sagittal sections
through stereotaxic space are superimposed onto a representative brain
anatomy. The Chinese group shows increased activation in the left IPL, as Hemispheric roles in speech prosody
compared to the English group, centered in ventral aspects of the
supramarginal gyrus, and extending into the bordering edge of the Sylvian The major findings of this study demonstrate that Chinese tone
fissure. Similar activation foci in the IPL are also observed in I3 vs. L3, T1 and intonation are best thought of as a mosaic of multiple local
vs. L1, and T3 vs. L3 comparisons. See also Fig. 4. asymmetries that allows for the possibility that different regions
J. Gandour et al. / NeuroImage 23 (2004) 344–357 351
Table 4
Group effects per task-and-hemisphere from statistical analyses on mean Dz within each spherical ROI
Group Hemi Task Frontal Parietal Temporal
aMFG mMFG pMFG FO IPS IPL aSTG mSTS pSTG
C>E LH I1 ***
T1 ***
I3 ***
T3 ***
RH I1
T1
I3
T3
E>C LH I1 **
T1 *** * *** ** *** **
I3 **
T3 ** * ** * **
RH I1 * **
T1 *** * *** ** *** * **
I3 * **
T3 ** * ** * * **
Note. C = Chinese group; E = English group; Hemi = hemisphere. LH = left hemisphere; RH = right hemisphere. *F(1, 18), P < 0.05; **F(1, 18), P < 0.01;
***F(1, 18), P < 0.001. See also notes to Tables 1 and 2.
may be differentially weighted in laterality depending on language-, that are lateralized to the LH in response to all tasks or subsets of
modality-, and task-related features (Ide et al., 1999). Earlier tasks are found in the Chinese group only (Fig. 9). Conversely, the
hypotheses that focus on hemispheric function capture only part two regions in the temporal and frontal lobes that are lateralized to
of, but not the whole, phenomenon. Not all aspects of speech the RH are found in both language groups. We infer that LH
prosody are lateralized to the RH. Cross-language differences in laterality reflects higher-order processing of internal representations
laterality of particular brain regions depend on a listener’s implicit of Chinese tone and intonation, whereas RH laterality reflects
knowledge of the relation between external stimulus features lower-order processing of complex auditory stimuli.
(acoustic/auditory) and internal conceptual representations (linguis- Previous models of speech prosody processing in the brain have
tic/prosodic). All regions in the frontal, temporal, and parietal lobes either focused on linguistics or acoustics as the driving force
underlying hemispheric lateralization. In this study, tone and
intonation are lateralized to the LH for the Chinese group. Despite
their functional differences from a linguistic perspective, they both
recruit shared neural mechanisms in frontal, temporal, and parietal
regions of the LH. The finding that intonation is lateralized to the
LH cannot be accounted for by a model that claims that ‘‘supra-
segmental sentence level information of speech comprehension is
subserved by the RH’’ (Friederici and Alter, 2004, p. 268). Neither
can this finding be explained by a hypothesis based on the size of
the temporal integration window (short ! LH; long ! RH)
(Poeppel, 2003). In spite of the fact that both intonation and tone
meet his criteria for a long temporal integration window, they are
lateralized to the LH instead of the RH.
Instead of viewing hemispheric roles as being derived from
either acoustics or linguistics independently, we propose that both
linguistics and acoustics, in addition to task demands (Plante et
al., 2002), are all necessary ingredients for developing a neuro-
biological model of speech prosody. This model relies on dynamic
interactions between the two hemispheres. Whereas the RH is
engaged in pitch processing of complex auditory signals, includ-
ing speech, we speculate that the LH is recruited to process
categorical information to support phonological processing, or
Fig. 6. Random effects fMRI activation map obtained from comparison of even syntactic and semantic processing (cf. Friederici and Alter,
discrimination judgments of tone in one-syllable utterances (T1) relative to
2004). With respect to task demands, I1 elicits greater activation
passive listening to the same stimuli (L1) between the two language groups
(DzT1 English vs. DzT1 Chinese). An axial section reveals increased
than T1 in the left aMFG and bilaterally in the pMFG. These
activation bilaterally in both frontal and parietal regions, as well as in the differences cannot be explained by ‘‘prosodic frame length’’
supplementary motor area, for the English group relative to the Chinese (Dogil et al., 2002) since both tone and intonation are presented
group. Similar activation foci are also observed in the T3 vs. L3 in an identical temporal context (one-syllable). These findings
comparison. See also Fig. 4. cannot be explained by a model that claims that segmental, lexical
352 J. Gandour et al. / NeuroImage 23 (2004) 344–357
Table 5
Within-group hemisphere effects per task from statistical analyses on mean Dz within each spherical ROI
Group Hemi Task Frontal Parietal Temporal
aMFG mMFG pMFG FO IPS IPL aSTG mSTS pSTG
C LH > RH I1 + ** *
T1 ** *
I3 ++ ** *
T3 ** *
RH > LH I1 * *
T1 * *
I3 * *
T3 * *
E LH > RH I1
T1
I3
T3
RH > LH I1 * *
T1 * *
I3 * *
T3 * *
Note. *F(1, 9), P < 0.05; **F(1, 9), P < 0.01; +tTukey-adjusted(9), P < 0.05. See also notes to Tables 2 and 4. ++
tTukey-adjusted(9), P < 0.01.
(i.e., tone), and syntactic information is processed in the LH, intonation judgment of the second stimulus competes for more
suprasegmental sentence level information (i.e., intonation) in the attentional resources and leads to greater effort in memory retrieval
RH (Friederici and Alter, 2004). Rather, they most likely reflect of intonation from the first stimulus. This process presumably
task demands related to retrieval of internal representations elicits greater activity in the left frontopolar region for intonation
associated with tone and intonation. tasks in Chinese listeners. English listeners, on the other hand,
employ a different processing strategy regardless of linguistic
Functional heterogeneity within a spatially distributed network function. Without prior knowledge of the Chinese language,
retrieving auditory information from working memory and making
Frontal lobe discrimination judgments is presumed to be equally difficult
Activation in the frontopolar cortex (BA 10) was bilateral between tone and intonation, resulting in bilateral activation of
across all tasks for English listeners, but predominantly left-sided frontopolar cortex for all tasks.
in the intonation tasks (I1, I3) for Chinese listeners (Table 5). The Dorsolateral prefrontal cortex, including BA 46 and BA 9, is
frontopolar region has extensive interconnections with auditory involved in controlling attentional demands of tasks and maintain-
regions of the superior temporal gyrus (Petrides and Pandya, ing information in working memory (Corbetta and Shulman, 2002;
1984). Thus, when presented with a competing articulatory sup- Knight et al., 1999; MacDonald et al., 2000; Mesulam, 1981). The
pression task, bilateral activation of frontopolar cortex has been rightward asymmetry in the mMFG (BA 46) that is observed in all
reported in a verbal working memory paradigm (Gruber, 2001). Its tasks (I1, I3, T1, T3) in both language groups (Table 5) points to a
functional role is inferred to be that of integrating working memory stage of processing that involves auditory attention and working
with the allocation of attentional resources (Koechlin et al., 1999), memory. Functional neuroimaging data reveal that auditory selec-
or applying greater effort in memory retrieval (Buckner et al., tive attention tasks elicit increased activity in right dorsolateral
1996; Schacter et al., 1996). prefrontal cortex (Zatorre et al., 1999). In the music domain,
These cross-language differences in frontopolar activation are perceptual analysis and short-term maintenance of pitch informa-
likely to result from the linguistic function of suprasegmental tion underlying melodies recruits neural systems within the right
information in Chinese and English. As measured by RT and prefrontal and temporal cortex (Zatorre et al., 1994). In this study,
accuracy, Chinese listeners take longer and are less proficient in activation of the prefrontal mMFG and temporal mSTS is similarly
judging intonation than tone. The relatively greater difficulty in lateralized to the RH across tasks in both language groups. These
intonation judgments presumably reflects the fact that in Chinese, data are consistent with the idea that the right dorsolateral
all syllables carry tonal contours obligatorily. Tones are likely to be prefrontal area (BA 46/9) plays a role in auditory attention that
processed first, as compared to intonation, due to this syllable-by- modulates pitch perception in sensory representations beyond the
syllable processing. By comparison, intonation contours play a lateral belt of the auditory cortex, and actively retains pitch
comparatively minor role in signaling differences in sentence information in auditory working memory (cf. Plante et al., 2002).
mood. In this study, the unmarked (i.e., minus a sentence-final Albeit in the speech domain, this frontotemporal network in the RH
particle) yes – no interrogatives are known to carry a light func- serves to maintain pitch information regardless of its linguistic
tional load (Shen, 1990). relevance. A frontotemporal network for auditory short-term mem-
In the present study, subjects were required to keep tone or ory is further supported by epileptic patients who show significant
intonation information of the first stimulus in a pair in their deficits in retention of tonal information after unilateral excisions
working memory while concurrently accessing tone or intonation of the right frontal or temporal regions (Zatorre and Samson,
identification of the second stimulus. Due to the functional 1991). In nonhuman primates, a processing stream for sound-
difference between tone and intonation for Chinese listeners, object identification has been proposed that projects anteriorly
J. Gandour et al. / NeuroImage 23 (2004) 344–357 353
Fig. 7. Random effects fMRI activation maps obtained from comparison of discrimination judgments of intonation (I3; upper panel) and tone (T3; bottom
panel) in three-syllable utterances relative to passive listening to the same stimuli (L3) for the Chinese group (zI3 vs. zL3; zT3 vs. zL3). In I3 vs. L3 and I1 vs. L1
(not shown), increased activity in frontopolar cortex (aMFG) shows a leftward asymmetry (upper panel; x = 35), whereas activation of the middle (mMFG)
region of dorsolateral prefrontal cortex shows the opposite laterality effect (upper panel; x = + 35, + 40, + 45). In T3 vs. L3, IPS activity is predominant in the
LH (bottom panel; x = 35, 40, 45). In I3 (upper panel; x = + 35, + 40, + 45) vs. T3 (lower panel; x = + 35, + 40, + 45), activation of the right pMFG is
greater in the I3 than the T3 task. See also Fig. 4.
along the lateral temporal cortex (Rauschecker and Tian, 2000), possible explanation has to do with the temporal interval. One
leading to the lateral prefrontal cortex (Hackett et al., 1999; might argue that the difference between I3 and T3 is due to the
Romanski et al., 1999a,b). A similar anterior processing stream time interval of focused attention for the prosodic unit: I3 = three
destined for the lateral prefrontal cortex in humans presumably syllables; T3 = last syllable only. On this view, shorter prosodic
underlies a frontotemporal network, at least in the RH, for low- frames are processed in the LH, longer frames in the RH. This
level auditory processing of complex pitch information. alternative account of pMFG activity is also ruled out because I1
Intonation elicited greater activity relative to tone in the pMFG elicits similar hemispheric laterality effects as I3. Instead, differ-
(BA 9), bilaterally in the one-syllable condition, right sided only in ential pMFG activity related to direct comparisons between into-
the three-syllable condition (Fig. 4c). The fact that I3 elicited nation and tone are most likely related to task demands (cf. Plante
greater activity than T3 in the posterior MFG of the RH replicates et al., 2002). As measured by RT and self-ratings of task difficulty,
Gandour et al. (2003). One possible explanation focuses on the intonation tasks are more difficult than tone for Chinese listeners
prosodic units themselves. Tones are processed in the LH, intona- (Table 3). Equally significant is the fact that the English group
tion predominantly in the RH. However, this account is untenable shows greater activation for tonal processing (T1, T3) than the
because I1 elicits greater activation bilaterally as compared to T1. Chinese group in the pMFG bilaterally (Table 4). These findings
Moreover, intonation (I1, I3) and tone (T1, T3) tasks separately together are consistent with the idea that the pMFG coordinates
elicit no hemispheric laterality effects in the pMFG. Another attentional resources required by the task.
354 J. Gandour et al. / NeuroImage 23 (2004) 344–357
Parietal lobe
There appear to be at least two distinct regions of activation in
the parietal cortex, one located more superiorly (IPS) in the
intraparietal sulcus and adjacent aspects of the superior parietal
lobule, another more inferiorly (IPL) in the anterior supramarginal
gyrus (SMG) near the parietotemporal boundary (cf. Becker et al.,
1999). Our findings show greater activation in the IPS bilaterally in
T1 for the English group compared to Chinese (Table 4). It has
been proposed that this area supports voluntary focusing and
shifting of attentional scanning across activated memory represen-
tations (Chein et al., 2003; Corbetta and Shulman, 2002; Corbetta
et al., 2000; Cowan, 1995; Mazoyer et al., 2002). The efficacy of
selective attention depends on how external stimuli are encoded
into internal phonological representations. English listeners expe-
rienced more difficulty in focusing and shifting of attention in T1
because lexically relevant pitch variations do not occur in English Fig. 9. Laterality effects for ROIs in the Chinese group only, and in both
monosyllables. Chinese and English groups, rendered on a three-dimensional LH template
for common reference. In the Chinese group (top panel), IPL, aSTG, and
In contrast, the Chinese group shows left-sided activity in the
pSTG are left-lateralized (LH > RH) across tasks; aMFG (I1, I3) and IPS
IPS for T3 (Table 5). This finding replicates our previous study of (T3) are left-lateralized for specific tasks. In both language groups (bottom
Chinese tone and intonation (Gandour et al., 2003), reinforcing the panel), mMFG and mSTS are right-lateralized (RH > LH) across tasks
view that a left frontoparietal network is recruited for the process- (bottom right panel). Other ROIs do not show laterality effects. No ROI
ing of lexical tones (Li et al., 2003). In T1, listeners extract tone elicited either a rightward asymmetry for the Chinese group only, or a
from isolated monosyllables. In T3, they extract tone from a fixed leftward asymmetry for both Chinese and English groups. See also Table 5.
J. Gandour et al. / NeuroImage 23 (2004) 344–357 355
activation of this area is observed in the English group. Its LH These findings collectively support functional segregation of
activity co-occurs with a leftward asymmetry in the pSTG across temporal lobe regions, and their functional integration as part of a
tasks. Co-activation of the IPL reinforces the view that it is part of temporofrontal network (Davis and Johnsrude, 2003; Scott, 2003;
an auditory – articulatory processing stream that connects posterior Scott and Johnsrude, 2003; Specht and Reul, 2003). LH networks
temporal and inferior prefrontal regions. An alternative conceptu- in the temporal lobe that are sensitive to phonologically relevant
alization is that the phonological storage component of verbal parameters from the auditory signal are in anterior and posterior, as
working memory resides in the IPL (Awh et al., 1996; Paulesu et opposed to central, regions of the STG/STS (Giraud and Price,
al., 1993). This notion predicts that both passive listening and 2001). The anterior region appears to be part of an auditory-
verbal working memory tasks should elicit activation in this region, semantic processing stream, the posterior region part of an audi-
since auditory verbal information has obligatory access to the store tory-motor processing stream. Both processing streams, in turn,
(Chein et al., 2003). I1, I3, T1, and T3 were all derived by project to convergence areas in the frontal lobe.
subtracting their corresponding passive listening control condition.
Contrary to fact, this notion would wrongly predict no increased Effects of task performance on hemispheric asymmetry
activation in the IPL.
In this study, the BOLD signal magnitude depends on the
Temporal lobe participant’s proficiency in a particular phonological task (Chee et
The anterior superior temporal gyrus (aSTG) displays an LH al., 2001). The two groups differ maximally in relative language
advantage in the Chinese group across tasks (Table 5). A reduced proficiency: Chinese group, 100%; English group, 0%. As reflected
RH, rather than increased LH aSTG activation, appears to in behavioral measures of task performance (Table 3), perceptual
underlie this hemispheric asymmetry across all tasks. Since judgments of Chinese tones require more cognitive effort by
intelligible speech is used in all tasks, phonological input alone English monolinguals due to their unfamiliarity with lexical tones.
may be sufficient to explain the leftward asymmetry in the Their unfamiliarity with the Chinese language results in greater
Chinese group (Scott and Johnsrude, 2003; Scott et al., 2000). BOLD activation for T1 and T3, either bilateral or RH only (cf.
It is also consistent with the notion that this region maps Chee et al., 2001). The effect of minimal language proficiency
acoustic – phonetic cues onto linguistic representations as part of applies only to lexical tone. Intonation, on the other hand, elicits
a larger auditory-semantic integration circuit in speech perception bilateral activation for both groups in the posterior MFG, frontal
(Giraud and Price, 2001; Scott and Johnsrude, 2003; Scott et al., operculum, and intraparietal sulcus (Table 4; Fig. 4). This common
2003). In contrast, English listeners do not have knowledge of frontoparietal activity implies that processing of intonation requires
these prosodic representations. Consequently, they employ a similar cognitive effort for Chinese and English participants.
nonlinguistic pitch processing strategy across tasks and fail to
show any hemispheric asymmetry.
A language group effect is not found in hemispheric laterality Conclusions
of the mSTS (BA 22/21). Both groups show greater RH activity in
the mSTS across tasks (Table 5). This suggests that this area is Cross-language comparisons provide unique insights into the
sensitive to different acoustic features of the speech signal irre- functional roles of different areas of this cortical network that are
spective of language experience. The rightward asymmetry may recruited for processing different aspects of speech prosody (e.g.,
reflect shared mechanisms underlying early attentional modulation auditory, phonological). By using tone and intonation tasks, we are
in processing of complex pitch patterns. In this study, subjects were able to distinguish hemispheric roles of areas sensitive to linguistic
required to direct their attention to slow modulation of pitch levels of processing (LH) from those sensitive to lower-level
patterns (i.e., c300 – 1000 ms) underlying either Chinese tone or acoustical processing (RH). Rather than attribute processing of
intonation. This interpretation is consistent with hemispheric roles speech prosody to RH mechanisms exclusively, our findings
hypothesized for auditory processing of complex sounds in the suggest that lateralization is influenced by language experience
temporal lobe: RH for spectral processing, LH for temporal that shapes the internal prosodic representation of an external
processing (Poeppel, 2003; Zatorre and Belin, 2001; Zatorre et auditory signal. This emerging model assumes a close interaction
al., 2002). Moreover, it is consistent with the view that right between the two hemispheres via the corpus callosum. In sum, we
auditory cortex is most important in the processing of dynamic propose a more comprehensive model of speech prosody percep-
pitch variation (Johnsrude et al., 2000). Both groups show greater tion that is mediated primarily by RH regions for complex-sound
activation in the right mSTS. We therefore infer that this activity analysis, but is lateralized to task-dependent regions in the LH
reflects a complex aspect of pitch processing that is independent of when language processing is required.
language experience.
A left asymmetric activation of the posterior part of the superior
temporal gyrus (pSTG; BA 22) across tasks is observed in the Acknowledgments
Chinese group only (Table 5). It has been suggested that the left
pSTG, as part of a posterior processing stream, is involved in Funding was provided by a research grant from the National
prelexical processing of phonetic cues and features (Scott, 2003; Institutes of Health R01 DC04584-04 (JG) and an NIH
Scott and Johnsrude, 2003; Scott and Wise, 2003). English listen- postdoctoral traineeship (XL). We are grateful to J. Lowe, T.
ers, however, show no leftward asymmetry in the pSTG (Table 5). Osborn, and J. Zimmerman for their technical assistance in the
Moreover, they show greater activation bilaterally relative to the MRI laboratory. Portions of this research were presented at the 11th
Chinese group (Table 4). Therefore, auditory phonetic cues that are annual meeting of the Cognitive Neuroscience Society, San
of phonological significance in one’s native language may be Francisco, April 2004. Correspondence should be addressed to
primarily responsible for this leftward asymmetry. Jack Gandour, Department of Audiology and Speech Sciences,
356 J. Gandour et al. / NeuroImage 23 (2004) 344–357
Purdue University, West Lafayette, IN 47907-2038, or via email: Gandour, J., Wong, D., Hsieh, L., Weinzapfel, B., Van Lancker, D., Hutch-
gandour@purdue.edu. ins, G.D., 2000. A crosslinguistic PET study of tone perception.
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Tong, Y., Li, X., 2002. A cross-linguistic FMRI study of spectral and
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