## Abstract

Sign language activates the auditory cortex of deaf subjects, which is evidence of cross-modal plasticity. Lip-reading (visual phonetics), which involves audio-visual integration, activates the auditory cortex of hearing subjects. To test whether audio-visual cross-modal plasticity occurs within areas involved in cross-modal integration, we used functional MRI to study seven prelingual deaf signers, 10 hearing non-signers and nine hearing signers. The visually presented tasks included mouth-movement matching, random-dot motion matching and sign-related motion matching. The mouth-movement tasks included conditions with or without visual phonetics, and the difference between these was used to measure the lip-reading effects. During the mouth-movement matching tasks, the deaf subjects showed more prominent activation of the left planum temporale (PT) than the hearing subjects. During dot-motion matching, the deaf showed greater activation in the right PT. Sign-related motion, with or without a lexical component, activated the left PT in the deaf signers more than in the hearing signers. These areas showed lip-reading effects in hearing subjects. These findings suggest that cross-modal plasticity is induced by auditory deprivation independent of the lexical processes or visual phonetics, and this plasticity is mediated in part by the neural substrates of audio-visual cross-modal integration.

## Introduction

There is evidence of cross-modal plasticity during the perception of sign language by deaf subjects. Sign language involves the visual perception of hand and face movements. Using functional MRI (fMRI), Neville et al. (1998) observed increased activity in the superior temporal sulcus (STS) during comprehension of American sign language in congenital deaf subjects and hearing signers. They suggested that the STS is involved in the linguistic analysis of sign language. Using PET imaging, Nishimura et al. (1999) found that activity was increased in the auditory association cortex, but not in the primary auditory cortex, of a prelingual deaf individual during comprehension of Japanese sign language. After this patient received a cochlear implant, the primary auditory cortex was activated by the sound of spoken words, but the auditory association cortex was not. Nishimura et al. (1999) suggested that audio-visual cross-modal plasticity is confined to the auditory association cortex, and that cognitive functions (such as sign language) might lead to functional plasticity in the underutilized auditory association cortex. Moreover, Petitto et al. (2000) observed increased activity in the superior temporal gyrus (GTs) in native deaf signers compared with hearing non-signers. These findings suggest that the auditory association cortex is where the changes associated with audio-visual cross-modal plasticity occur.

However, the activation of the auditory cortex by visual stimuli is not specific to deaf subjects. In hearing subjects, lip-reading enhances the perception of spoken language. Combining audible speech with visible articulation movements can improve comprehension to the same extent as altering the acoustic signal-to-noise ratio by 15–20 dB (Sumby and Polack, 1954). Calvert et al. (1997) found that linguistic visual cues during lip-reading activated the auditory cortex and GTs in hearing individuals in the absence of speech sounds, whereas non-linguistic facial movement did not. The findings of Calvert et al. (1997) suggest that this type of audio-visual integration (face/voice matching) occurs in the superior temporal cortices.

Our hypothesis was that audio-visual cross-modal plasticity occurs within areas involved in cross-modal integration. To test this, we recruited deaf and hearing subjects for a fMRI study. Three tasks were included: mouth-movement matching, random dot-movement matching, and sign-related motion matching. The mouth-movement matching task contained open-mouth movements (OPEN), representing visual phonetics, and closed-mouth movements (CLOSED), which are not phonetic. Thus, the difference between the OPEN and CLOSED conditions can be used as a marker for phonetic speech perception (lip-reading effect). Cross-modal plastic changes were evaluated by comparing the task-related activation between deaf and hearing groups during closed-mouth, random-dot and sign-related movements without lexical processing. The bimodal activation of the auditory cortex in hearing subjects suggests that visual and auditory inputs are balanced competitively (Rauschecker, 1995). Hence, with auditory deprivation, the bimodal association auditory cortex is expected to be activated by the visual processing that usually does not activate the auditory cortex. As the CLOSED task does not contain visual phonetics, we expected that it would not activate the association auditory cortex of the hearing, whereas it would activate that of the deaf subjects. The random-dot movement-matching task was included to determine whether the plastic change is specific to the biological mouth motion or related to motion in general. Sign-word/non-word discrimination tasks were also included to determine whether the plastic changes are related to the motion of the hands or linguistic processes. To this end, a direct comparison was made between the deaf signers and the hearing signers.

## Subjects and Methods

### Subjects

Our volunteers were 19 hearing subjects (11 women and eight men) aged 36.0 ± 10.7 years (mean ± SD), and seven prelingual deaf subjects (one woman and six men) aged 31.6 ± 7.1 years. Nine of the hearing subjects and all of the deaf subjects were fluent in Japanese sign language. The deaf subjects also had been trained in lip-reading at deaf school. The characteristics of the subjects are summarized in Table 1. The subjects were all right-handed according to the Edinburgh handedness inventory (Oldfield, 1971). None of the subjects had any history of neurological or psychiatric illness, and none had any neurological deficits other than deafness. The protocol was approved by the ethical committee of Fukui Medical University, Japan, and the National Institute for Physiological Sciences. All subjects gave their written informed consent for the study.

Table 1

Characteristics of the participants

Subject no.

Age

Sex

Age at start of sign language (years)

Age at onset of deafness (years)

Cause of deafness

Residual hearing (dB)

Deaf
34 <3 Prematurity 100
44 Prematurity 100
32 Prematurity >100
26 19 Prematurity 100
33 Fever >100
21 Inherited >100
31 Fever >100
Hearing signers
51 46
35 29
37 31
30 23
25 17
43 35
53 45
59 49
46 20
Hearing non-signers
25
25
27
27
28
29
29
29
39
10

46

M

Subject no.

Age

Sex

Age at start of sign language (years)

Age at onset of deafness (years)

Cause of deafness

Residual hearing (dB)

Deaf
34 <3 Prematurity 100
44 Prematurity 100
32 Prematurity >100
26 19 Prematurity 100
33 Fever >100
21 Inherited >100
31 Fever >100
Hearing signers
51 46
35 29
37 31
30 23
25 17
43 35
53 45
59 49
46 20
Hearing non-signers
25
25
27
27
28
29
29
29
39
10

46

M

### fMRI Experiment

This experiment consisted of three tasks: a lip-movement task, a moving-dot task and signed-word discrimination. Each task was performed in separate sessions. Hearing signers, hearing non-signers and deaf signers underwent the mouth-movement and dot-motion tasks. Hearing signers and deaf signers participated in the signing tasks. The lip-motion and dot-motion sessions were repeated twice.

### Experimental Setup

Visual stimuli were displayed on a half-transparent screen hung 285 cm from the subject's eyes, and were shown via an LCD projector (Epson ELP-7200L, Tokyo, Japan) connected to a digital video camera (SONY, Tokyo, Japan). Subjects viewed the screen through a mirror. The screen covered a visual angle of ∼10°. The subject's right hand was placed on a button box connected to a microcomputer that recorded the subject's responses.

### MRI

A time-course series of volumes was acquired using T2*-weighted gradient echo-planar imaging (EPI) sequences with a 3.0 T magnetic resonance imager (VP, General Electric, Milwaukee, WI). Each volume consisted of 28 slices, with a slice thickness of 4.0 mm and a 1.0 mm gap, and included the entire cerebral and cerebellar cortex. The time interval between two successive acquisitions of the same image was 2500 ms, the echo time was 30 ms and the flip angle was 81°. The field of view (FOV) was 19 cm. The in-plane matrix size was 64 × 64 pixels with a pixel dimension of 3.0 × 3.0 mm. Tight, but comfortable, foam padding was placed around the subject's head to minimize head movement.

Immediately after five fMRI sessions, T2-weighted fast-spin echo images were obtained from each subject using location variables that were identical to those of the EPIs used for anatomical reference. In addition, high-resolution whole-brain MRIs were obtained with a conventional T2-weighted fast-spin echo sequence. A total of 112 transaxial images were obtained. The in-plane matrix size was 256 × 256 mm, slice thickness was 1.5 mm and pixel size was 0.742 × 0.742 mm.

The event-related design consisted of three event conditions: OPEN, CLOSED and a still face. Throughout the session, the subjects fixated a small cross-hair in the center of the screen. During the OPEN condition, four consecutive frames (frame duration = 500 ms) of a face pronouncing a vowel (/a/, /i/, /u/, /e/, or /o/) were presented (Fig. 1). No auditory input was provided. Following the stimulus, we presented a question mark (‘?’) for 400 ms. This prompted the subject to press a button with their right index finger if the movements of the lips in the first and last frames were the same, and with the middle finger if the movements were not the same. The CLOSED condition was similar to the OPEN condition, except that the mouth was always closed during the movement, which consisted of five patterns of twitches of the lips (Fig. 1). During the still-face event condition (STILL), a face without any movement was presented for 2 s followed by the 400 ms presentation of an arrow pointing to the right or left. The subject was instructed to press the button with their right index finger if the leftward arrow was shown, and with the middle finger if the rightward arrow was shown. The inter-trial interval (ITI) was fixed at 3 s. Each condition was repeated 30 times, so there were 90 events in total.

Figure 1.

Task design. (Top) The event-related design of the face-motion discrimination task consisted of three conditions: OPEN, CLOSED and a still face. During the OPEN condition, four consecutive frames (each frame = 500 ms duration) of a face pronouncing a vowel (/a/, /i/, /u/, /e/, or /o/) were presented. A 400 ms presentation of a question mark ‘?’ followed, which prompted the subject to press the button with their right index finger if the movement of the lips in the first and last frames was the same, and otherwise to press the button with their right middle finger. The CLOSED condition was similar to the OPEN condition, except that the mouth was closed during the movement. During the still-face condition (STILL), a face without any movement was presented for 2 s followed by the 400 ms presentation of an arrow that pointed either right or left. The subject was instructed to press the button with their right index finger if the leftward arrow was displayed, and with the middle finger if the rightward arrow appeared. Each condition was repeated 30 times. (Middle) Block design of the dot-motion discrimination task. (Bottom) The event-related design of the sign-word discrimination task consisted of a sign-word/pseudo-word discrimination condition (D), a non-discrimination (ND) condition using flipped images, and control conditions [Still (S) and Flipped Still (FS)], during which corresponding still images were presented. During the D condition, a single frame of an animated character moving her hands and arms was presented for 2 s, followed by the presentation of a question mark ‘?’ for 400 ms, which cued the subject to press the button with their right index finger if the movements constituted a real word, and otherwise to perform the button press with their right middle finger. ND was similar to D, except that the reversed upside-down images were presented, followed by the presentation of arrows indicating the button press for 400 ms. S was similar to ND, except that the still images of the same character, both flipped and non-flipped, were presented. The ITI was fixed at 3 s. Each condition was repeated 30 times.

Figure 1.

Task design. (Top) The event-related design of the face-motion discrimination task consisted of three conditions: OPEN, CLOSED and a still face. During the OPEN condition, four consecutive frames (each frame = 500 ms duration) of a face pronouncing a vowel (/a/, /i/, /u/, /e/, or /o/) were presented. A 400 ms presentation of a question mark ‘?’ followed, which prompted the subject to press the button with their right index finger if the movement of the lips in the first and last frames was the same, and otherwise to press the button with their right middle finger. The CLOSED condition was similar to the OPEN condition, except that the mouth was closed during the movement. During the still-face condition (STILL), a face without any movement was presented for 2 s followed by the 400 ms presentation of an arrow that pointed either right or left. The subject was instructed to press the button with their right index finger if the leftward arrow was displayed, and with the middle finger if the rightward arrow appeared. Each condition was repeated 30 times. (Middle) Block design of the dot-motion discrimination task. (Bottom) The event-related design of the sign-word discrimination task consisted of a sign-word/pseudo-word discrimination condition (D), a non-discrimination (ND) condition using flipped images, and control conditions [Still (S) and Flipped Still (FS)], during which corresponding still images were presented. During the D condition, a single frame of an animated character moving her hands and arms was presented for 2 s, followed by the presentation of a question mark ‘?’ for 400 ms, which cued the subject to press the button with their right index finger if the movements constituted a real word, and otherwise to perform the button press with their right middle finger. ND was similar to D, except that the reversed upside-down images were presented, followed by the presentation of arrows indicating the button press for 400 ms. S was similar to ND, except that the still images of the same character, both flipped and non-flipped, were presented. The ITI was fixed at 3 s. Each condition was repeated 30 times.

To maximize the efficiency of detecting the differences between the CLOSED and STILL, OPEN and STILL, and OPEN and CLOSED conditions, the distribution of stimulus onset asynchronies (SOAs) of each condition were determined as follows (Friston et al., 1999b): the order of the 90 events (30 for each condition) was permutated randomly to generate a set of three vectors (1 × 90 matrix) indicating the presence (1) or absence (0) of a particular event, and hence representing the distribution of the SOAs of each condition (SOA vectors). A design matrix (X) incorporating the three conditions (CLOSED, OPEN and STILL) was created by convolving a set of SOA vectors with a hemodynamic-response function:

$X{=}[{\vec{c}},{\vec{o}},{\vec{s}}]{\otimes}{\vec{h}}$
where c⃗ is for CLOSED, o⃗ is for OPEN and s⃗ is for STILL.

The efficiency of the estimation of the CLOSED–STILL, OPEN–STILL and OPEN–CLOSED comparisons was evaluated by taking the inverse of the covariance of the contrast between parameter estimates (Friston et al., 1999b),

$\mathrm{var}{\{}c^{T}\mathrm{{\hat{{\beta}}}}{\}}{=}{\sigma}^{2}c^{T}(X^{T}X)^{{-}1}c$

$Efficiency{=}trace{\{}c^{T}(X^{T}X)^{{-}1}c{\}}^{{-}1}$
where c = (1, 0, −1) for CLOSED–STILL, (0, 1, −1) for OPEN–STILL and (−1, 1, 0) for OPEN–CLOSED. From the 1000 randomly generated sets of SOA vectors, we selected the most efficient based on the maximum of the sum of the square of the efficiency for three contrasts. The session took 4.5 min, and tried twice per subject.

The fMRI session with random-moving dot stimuli consisted of three periods of stationary dots and three periods of moving dots; each period was 30 s long, alternating stationary and moving stimuli (Fig. 1). Throughout the session, the subjects were asked to fixate a small cross-hair in the center of the screen. During the 30 s stationary period, every 3 s stationary random dots were presented for 2 s, followed by the 400 ms appearance of an arrow pointing either right or left. The frame size of the presented dots was 5°. The subject was instructed to press a button with their right index finger if the leftward arrow appeared, and with the middle finger if the rightward arrow appeared. During the 30 s moving-dot period, every 3 s four consecutive frames of coherently moving random dots were presented. In each frame, the random dots moved for 500 ms in a single horizontal direction (right or left). The dots moved at a speed of 2.5°/s. This was followed by a question mark, which was presented for 400 ms. This cued the subject to press the button with the right index finger if the direction of the movement in the first and last frames was the same, and otherwise to respond by pressing the middle finger button. Thus, the subject was required to match the movement direction of the random dots in the first and last frames. The session took 3 min, and tried twice per subject.

We selected 30 sign words that can be expressed by moving the hands and arms, and without facial expression. The animated sign words were generated with commercially available software (Mimehand II, Hitachi, Tokyo, Japan) implemented on a personal computer (Endevor Pro-600L, Dell, Tokyo, Japan). This software can modify all components of the movement separately. As the sign words are composed of several component movements of the hands or arms, pseudo-words can be generated by modifying a single component of each word. Thirty pseudo-words were generated from the 30 words. Non-words were generated by flipping the 30 words and 30 pseudo-words upside down. Finally, a still image was generated.

The event-related design consisted of three event conditions: word discrimination (D), non-discrimination (ND) and still (S). The order of presentation was determined to maximize the efficiency of the predefined contrasts, as in the face-movement tasks. During the D condition, a single frame (2000 ms duration) of an animated character moving her hands and arms appeared, followed by a question mark (400 ms), which cued the subject to press the button with their right index finger if the movements constituted a real word, and otherwise to press the button with the middle finger. The ND condition was similar to D, except that the reversed upside-down images were presented, and the arrows were presented for 400 ms. The subject was instructed to press the button with their right index finger if the leftward arrow appeared, and with the middle finger if the rightward arrow appeared. During the S condition, the same characters were presented but without any movement, and both flipped and non-flipped images were presented. The stimuli lasted 2 s and were followed by an arrow pointing either right or left for 400 ms. The subject was instructed to press the button with their right index finger if the leftward arrow appeared, and with the middle finger if the rightward arrow appeared. The ITI was fixed at 3 s. Each condition was repeated 60 times and there were 180 events in total. The session took 9 min, and tried once per subject.

### Data Analysis

The data were analyzed using statistical parametric mapping (SPM99, Wellcome Department of Cognitive Neurology, London, UK) implemented in Matlab (Mathworks, Sherborn, MA; Friston et al., 1994, 1995a,b). As our scanning protocol used sequential axial-slice acquisition in ascending order, signals in different slices were measured at different time points. To correct for this, we interpolated and resampled the data (Büchel and Friston, 1997). Following realignment, all images were co-registered to the high-resolution three-dimensional (3-D) T2-weighted MRI, using the anatomical MRI with T2-weighted spin-echo sequences from locations identical to those of the fMRI images. The parameters for affine and nonlinear transformation into a template of T2-weighted images already fitted to standard stereotaxic space (MNI template; Evans et al., 1994) were estimated using the high-resolution 3-D T2-weighted MRI employing least-square means (Friston et al., 1995b). The parameters were applied to the co-registered fMRI data. The anatomically normalized fMRI data were filtered using a Gaussian kernel of 8 mm (full width at half maximum) in the x, y and z axes.

### Statistics

Statistical analysis was conducted at two levels. First, individual task-related activation was evaluated. Secondly, individual data were summarized and incorporated into a random-effect model so that inferences could be made at a population level (Friston et al., 1999a).

#### Individual Analysis

The signal was scaled proportionally by setting the whole-brain mean value to 100 arbitrary units. Proportional scaling was performed to adjust for the fluctuation of the sensitivity of the MRI scanner which affects both global and local signals. Here we assume that the global signal (which is an indirect measure of global cerebral blood flow) is unchanged and that the local change will not affect the global signal. The signal time-course of each subject was modeled with two box-car functions convolved with a hemodynamic-response function, high-pass filtering and session effect. To test hypotheses about regionally specific condition effects, the estimates for each condition were compared using linear contrasts of the matching task versus the rest period. The resulting set of voxel values for each comparison constituted a statistical parametric map (SPM) of the t statistic (SPM{t}). The SPM{t} was transformed to normal-distribution units (SPM{Z}). The threshold for the SPM{Z} of individual analyses was set at P < 0.05, correcting for multiple comparisons at the voxel level for the entire brain (Friston et al., 1996).

#### Group Analysis with the Random-effect Model

The weighted sum of the parameter estimates in the individual analysis constituted ‘contrast’ images that were used for the group analysis (Friston et al., 1999a). The contrast images obtained by individual analysis represent the normalized task-related increment of the MR signal of each subject, comparing the matching task versus rest period. To examine the effect of the tasks by group (deaf and hearing) and the task × group interaction, the contrast images of the prelingual deaf and hearing groups were entered into a random-effect model. For the mouth-movement and dot-motion tasks, hearing signers and hearing non-signers were pooled, as there was no significant group difference. The comparison was then made between the deaf (n = 7) and hearing (n = 19) groups. For the sign tasks, the deaf subjects (n = 7) and hearing signers (n = 9) were compared. Significant signal changes for each contrast were assessed using t-statistics on a voxel-by-voxel basis (Friston et al., 1995a). The resulting set of voxel values for each contrast constituted an SPM of the t-statistic (SPM{t}). The SPM{t} was transformed into normal-distribution units (SPM{Z}). The threshold for the SPM{Z} was set at Z > 3.09 and P < 0.05, with a correction for multiple comparisons at the cluster level for the entire brain (Friston et al., 1996).

## Results

### Face Motion

In the hearing subjects, during the CLOSED condition the matching accuracy was 93.0 ± 5.9%, and accuracy during the OPEN condition was 90.8 ± 5.5%, with no significant difference between the conditions. In the deaf subjects, during the CLOSED condition the matching accuracy was 92.7 ± 5.7%, and accuracy during the OPEN condition was 86.5 ± 9.6%, with no significant difference.

During the CLOSED condition compared with the STILL condition, activation was found in the bilateral inferior (GFi) and middle (GFm) frontal gyrus, temporo-occipital junction, pre- and post-central gyri, superior (LPs) and inferior (LPi) parietal lobules, dorsal premotor cortex (PMd), supplementary motor cortex (SMA), posterior lobe of the cerebellar hemisphere, and left basal ganglia and thalamus. The left GTs, corresponding to the planum temporale (PT), was more active during the CLOSED condition in deaf subjects compared with hearing subjects (Fig. 2). During the OPEN condition compared with the STILL condition, activation was found in almost identical regions to those found in the CLOSED–STILL comparison. However, in the hearing subjects, when the OPEN condition was compared with the CLOSED, activation was found in the left middle and inferior frontal gyrus, the bilateral GTs along the superior temporal sulci and the SMA (Table 2, Fig. 3). Post hoc tests confirmed that the responses in the left PT of the hearing signers and non-signers were similar (P > 0.1, two-sample t-test), and that both hearing and deaf subjects showed a significant lip-reading effect (OPEN–CLOSED) (Fig. 2).

Figure 2.

Group analysis of CLOSED. (Upper half) SPMs of the average neural activity within each group, deaf on the left and hearing on the right, during CLOSED compared with the rest period. The maps are shown in standard anatomical space. The 3-D information was collapsed into 2-D sagittal and transverse images (that is, maximum-intensity projections viewed from the right and top of the brain). The statistical threshold was P < 0.05 with a correction for multiple comparisons at the cluster level. (Lower half) There was more prominent activation in the PT on the left in deaf subjects than in hearing subjects during the CLOSED condition. The focus of activation on a pseudocolor fMRI superimposed on a high-resolution anatomical MRI in the sagittal (upper left), coronal (upper right) and transaxial (lower left) planes, as indicated by the blue lines that cross at (−56, −18, 6), corresponding to the left PT (Brodmann area 42). The significance level of activity is as indicated by the color bar, increasing as red proceeds to white. The statistical threshold was P < 0.05, corrected for multiple comparisons at the cluster level. Lower right, percent signal change in the left PT of the deaf (open square) and hearing (closed square) during CLOSED, OPEN and OPEN–CLOSED. The right PT counterpart showed the same tendency with less prominent increments. * indicates P < 0.001 (two-sample t-test), ** indicates P = 0.041 (one-sample t-test), and +, P = 0.001 (one-sample t-test). Post hoc tests in the region of interest in the left PT are specified by the group comparisons of the CLOSED condition.

Figure 2.

Group analysis of CLOSED. (Upper half) SPMs of the average neural activity within each group, deaf on the left and hearing on the right, during CLOSED compared with the rest period. The maps are shown in standard anatomical space. The 3-D information was collapsed into 2-D sagittal and transverse images (that is, maximum-intensity projections viewed from the right and top of the brain). The statistical threshold was P < 0.05 with a correction for multiple comparisons at the cluster level. (Lower half) There was more prominent activation in the PT on the left in deaf subjects than in hearing subjects during the CLOSED condition. The focus of activation on a pseudocolor fMRI superimposed on a high-resolution anatomical MRI in the sagittal (upper left), coronal (upper right) and transaxial (lower left) planes, as indicated by the blue lines that cross at (−56, −18, 6), corresponding to the left PT (Brodmann area 42). The significance level of activity is as indicated by the color bar, increasing as red proceeds to white. The statistical threshold was P < 0.05, corrected for multiple comparisons at the cluster level. Lower right, percent signal change in the left PT of the deaf (open square) and hearing (closed square) during CLOSED, OPEN and OPEN–CLOSED. The right PT counterpart showed the same tendency with less prominent increments. * indicates P < 0.001 (two-sample t-test), ** indicates P = 0.041 (one-sample t-test), and +, P = 0.001 (one-sample t-test). Post hoc tests in the region of interest in the left PT are specified by the group comparisons of the CLOSED condition.

Figure 3.

Group analysis of OPEN–CLOSED. The task-related increments of the MR signal in the regions that showed lip-reading effects. SPMs of the OPEN–CLOSED (O–C) comparison in the hearing subjects are shown in standard anatomical space (left column). The 3-D information was collapsed into 2-D sagittal, coronal and transverse images. (Right column) Focus of activation on a pseudocolor fMRI superimposed on a high-resolution anatomical MRI in the sagittal (upper right), transaxial (middle right) and coronal (lower right) planes.

Figure 3.

Group analysis of OPEN–CLOSED. The task-related increments of the MR signal in the regions that showed lip-reading effects. SPMs of the OPEN–CLOSED (O–C) comparison in the hearing subjects are shown in standard anatomical space (left column). The 3-D information was collapsed into 2-D sagittal, coronal and transverse images. (Right column) Focus of activation on a pseudocolor fMRI superimposed on a high-resolution anatomical MRI in the sagittal (upper right), transaxial (middle right) and coronal (lower right) planes.

Table 2

Fields significantly activated by lip-motion discrimination (mouth open versus closed)

Cluster P Cluster equiv. k Voxel Z Talairach's coordinates

Side Location BA

x

y

z

Deaf
N.S.
Hearing (signers + non-signers)
0.017 174 4.07 −46 14 −4 GFi 47
512 4.16 −48 12 34 GFi 44
3.71 −44 48 GFm
5.58 −48 −2 42 GprC
485 5.27 −2 64 SMA
4.6 −4 16 64 GFs
391 4.68 −62 −16 GTs 42
4.25 −56 −4 −6 GTm 21
3.88 −60 −32 GTs 22
0.023 162 4.54 62 −24 GTs 22
Deaf > hearing
N.S.

Cluster P Cluster equiv. k Voxel Z Talairach's coordinates

Side Location BA

x

y

z

Deaf
N.S.
Hearing (signers + non-signers)
0.017 174 4.07 −46 14 −4 GFi 47
512 4.16 −48 12 34 GFi 44
3.71 −44 48 GFm
5.58 −48 −2 42 GprC
485 5.27 −2 64 SMA
4.6 −4 16 64 GFs
391 4.68 −62 −16 GTs 42
4.25 −56 −4 −6 GTm 21
3.88 −60 −32 GTs 22
0.023 162 4.54 62 −24 GTs 22
Deaf > hearing
N.S.

Abbreviations: GFi, inferior frontal gyrus; GFm, middle frontal gyrus; GprC, precentral gyrus; GTm, middle temporal gyrus; GTs, superior temporal gyrus; SMA, supplementary motor area. BA, Brodmann's areas. All P values are corrected for multiple comparisons. Height threshold, Z = 3.09, P = 0.001. L, left; R, right.

### Dot Motion

Matching accuracy in the deaf group was 87.4 ± 9.1%, and in the hearing group accuracy was 85.1 ± 12.5%, with no significant group difference (P > 0.1, two-sample t-test).

In deaf subjects, the bilateral LPi, GTs and posterior lobe of the cerebellum, the left medial prefrontal cortex and PMd, and the right GFi and GFm were activated by the dot-motion discrimination task. In hearing subjects, the bilateral LPi, LPs, GFm, PMd, inferior and middle occipital gyrus, fusiform gyrus and posterior lobe of the cerebellum, the left medial prefrontal cortex and right GTs were activated. The right GTs (corresponding to the PT) was more active during the dot-motion discrimination task in deaf subjects compared with hearing subjects (Fig. 4). This area of hearing subjects showed a significant lip-reading effect during face motion tasks (contrasting OPEN–CLOSED). There were no areas in which the hearing subjects showed significantly more activation than deaf subjects did.

Figure 4.

Group analysis of dot motion discrimination. (Upper half) SPMs of the average neural activity within each group, deaf on the left and hearing on the right, during the dot-motion discrimination task compared with the rest period. The same format and statistical thresholds are used as in Figure 2. (Lower half) There was more prominent activation in the PT on the right in deaf subjects than in hearing subjects during the dot-motion discrimination task. The focus of activation on a pseudocolor fMRI is superimposed on a high-resolution anatomical MRI in the sagittal (upper left), coronal (upper right), and transaxial (lower left) planes, as indicated by the blue lines that cross at (56, −18, 6) corresponding to the right PT (Brodmann area 42). The significance level of activity is as indicated by the color bar; this increases as red proceeds to white. The statistical threshold was P < 0.05, corrected for multiple comparisons at the cluster level. Lower right, percent signal change in the right PT of the deaf (open square) and hearing (closed square) subjects during DOT, CLOSED, OPEN and O–C. * indicates P < 0.001 (two-sample t test), + P = 0.009 (two-sample t test), ++ P = 0.001 (two-sample t test), and × P = 0.004 (one-sample t test).

Figure 4.

Group analysis of dot motion discrimination. (Upper half) SPMs of the average neural activity within each group, deaf on the left and hearing on the right, during the dot-motion discrimination task compared with the rest period. The same format and statistical thresholds are used as in Figure 2. (Lower half) There was more prominent activation in the PT on the right in deaf subjects than in hearing subjects during the dot-motion discrimination task. The focus of activation on a pseudocolor fMRI is superimposed on a high-resolution anatomical MRI in the sagittal (upper left), coronal (upper right), and transaxial (lower left) planes, as indicated by the blue lines that cross at (56, −18, 6) corresponding to the right PT (Brodmann area 42). The significance level of activity is as indicated by the color bar; this increases as red proceeds to white. The statistical threshold was P < 0.05, corrected for multiple comparisons at the cluster level. Lower right, percent signal change in the right PT of the deaf (open square) and hearing (closed square) subjects during DOT, CLOSED, OPEN and O–C. * indicates P < 0.001 (two-sample t test), + P = 0.009 (two-sample t test), ++ P = 0.001 (two-sample t test), and × P = 0.004 (one-sample t test).

### Sign Discrimination

The accuracy of sign discrimination was 59.8 ± 2.9% in the hearing signers and 53.6 ± 5.4% in the deaf subjects, which was an insignificant difference. The deaf subjects showed an accuracy of 84.0 ± 35.7% in response to real words, and only 35.7 ± 11.3% accuracy in response to pseudo-words. The hearing signers showed 86.7 ± 12.1% accuracy in response to true words, and 40.6 ± 16.9% in response to pseudo-words.

In the hearing signers, the sign-discrimination and non-discrimination conditions activated the SMA, bilateral GFi, bilateral PMd, LPs, LPi, GOm and GTm. Deaf subjects showed a similar activation pattern, except more prominent activation was seen in the left GTs adjacent to the transverse temporal gyrus (Fig. 5, Table 3). The left GTs was activated more prominently in the deaf than in the hearing subjects during closed-lip movement, sign discrimination and non-discrimination. This area of hearing subjects showed a significant lip-reading effect during face motion tasks (contrasting OPEN–CLOSED). There was a marked overlap of the foci activated by lip-reading in hearing subjects and the cross-modal plasticity in the deaf, which was revealed by the sign-word discrimination/non-discrimination and lip-movement tasks (Fig. 6). Figure 6 showed that the common area for cross-modal integration (blue, as lip-reading in the hearing) and cross-modal plasticity (yellow for sign and red for CLSOED lip movement) is in the PT adjacent to the anterior border, the sulcus behind Heschl's gyrus (Westbury et al., 1999). On the other hand, the area for the lip-reading in the hearing (blue) extended to the posterior STS area, more prominent on the left. Sign discrimination compared with non-discrimination activated the bilateral PMd, GFm and GFi of both hearing and deaf signers but not the temporal cortices, whereas no activation was observed in the PT (Fig. 7).

Figure 5.

Group analysis of sign. (Upper half) SPMs of average neural activity within each group, deaf on the left and hearing on the right, during the sign-word discrimination and non-discrimination tasks compared with the rest period. The same format and statistical thresholds are used as in Figure 2. (Lower half) There was more prominent activation in the left PT in deaf subjects than in hearing subjects during the sign-word discrimination/non-discrimination tasks. The focus of activation on a pseudocolor fMRI is superimposed on a high-resolution anatomical MRI in the sagittal (upper left), coronal (upper right) and transaxial (lower left) planes, as indicated by the blue lines that cross at (−62, −20, 4) corresponding to the left PT (Brodmann area 42). The significance level of the activity is indicated by the color bar; this increases as red proceeds to white. The statistical threshold was P < 0.05, corrected for multiple comparisons at the cluster level. Lower right, percent signal change in the left PT (− 62, −20, 4) of the deaf (open square) and hearing (closed square) during word-discrimination and non-discrimination tasks, CLOSED, OPEN and O–C. *, ** and + indicate P < 0.001 (two-sample t-test), ++ P = 0.002 (two-sample t-test), × P = 0.004 and ×× P < 0.001 (one-sample t-test).

Figure 5.

Group analysis of sign. (Upper half) SPMs of average neural activity within each group, deaf on the left and hearing on the right, during the sign-word discrimination and non-discrimination tasks compared with the rest period. The same format and statistical thresholds are used as in Figure 2. (Lower half) There was more prominent activation in the left PT in deaf subjects than in hearing subjects during the sign-word discrimination/non-discrimination tasks. The focus of activation on a pseudocolor fMRI is superimposed on a high-resolution anatomical MRI in the sagittal (upper left), coronal (upper right) and transaxial (lower left) planes, as indicated by the blue lines that cross at (−62, −20, 4) corresponding to the left PT (Brodmann area 42). The significance level of the activity is indicated by the color bar; this increases as red proceeds to white. The statistical threshold was P < 0.05, corrected for multiple comparisons at the cluster level. Lower right, percent signal change in the left PT (− 62, −20, 4) of the deaf (open square) and hearing (closed square) during word-discrimination and non-discrimination tasks, CLOSED, OPEN and O–C. *, ** and + indicate P < 0.001 (two-sample t-test), ++ P = 0.002 (two-sample t-test), × P = 0.004 and ×× P < 0.001 (one-sample t-test).

Figure 6.

Summary of the between-group differences. The areas showing more prominent activation in the deaf than the hearing subjects during dot-motion discrimination (DOT, green), the CLOSED condition (red), sign-word discrimination and non-discrimination (SIGN, yellow) were superimposed with the areas activated by OPEN–CLOSED in the hearing subjects (blue outline). Marked overlap among the latter three conditions was seen in the anterior part of the left PT, adjacent to the Heschl's gyrus. On the other hand, the neural substrates of lip-reading in the hearing subjects extended to the ascending portion of the superior temporal sulcus (white arrowhead), passing the posterior border of the PT (black arrow head, descending ramus of the Sylvian fissure) (Westbury et al., 1999).

Figure 6.

Summary of the between-group differences. The areas showing more prominent activation in the deaf than the hearing subjects during dot-motion discrimination (DOT, green), the CLOSED condition (red), sign-word discrimination and non-discrimination (SIGN, yellow) were superimposed with the areas activated by OPEN–CLOSED in the hearing subjects (blue outline). Marked overlap among the latter three conditions was seen in the anterior part of the left PT, adjacent to the Heschl's gyrus. On the other hand, the neural substrates of lip-reading in the hearing subjects extended to the ascending portion of the superior temporal sulcus (white arrowhead), passing the posterior border of the PT (black arrow head, descending ramus of the Sylvian fissure) (Westbury et al., 1999).

Figure 7.

Areas more prominently activated during the sign-word discrimination task compared with the non-discrimination task in deaf subjects (top) and hearing subjects (bottom). The areas are superimposed on surface-rendered high-resolution MRIs viewed from the right, front and left. Both hearing and deaf signers exhibited activation of the bilateral frontal cortex without activation in the temporal cortex. The statistical threshold is P < 0.05, corrected for multiple comparisons at the cluster level.

Figure 7.

Areas more prominently activated during the sign-word discrimination task compared with the non-discrimination task in deaf subjects (top) and hearing subjects (bottom). The areas are superimposed on surface-rendered high-resolution MRIs viewed from the right, front and left. Both hearing and deaf signers exhibited activation of the bilateral frontal cortex without activation in the temporal cortex. The statistical threshold is P < 0.05, corrected for multiple comparisons at the cluster level.

Table 3

Fields significantly activated by sign-word discrimination and non-discrimination

Cluster P Cluster size Voxel Z Talairach's coordinates

Side Location BA

x

y

z

Deaf
1079 5.63 66  SMA
4.75 24 46 GFd
1531 4.85 −48 14 20 GFi 44
4.64 −36 50 PMd
4.45 −30 28 −2 GFi 47
1153 4.16 48 32 GFi 45
46 46 16 GFm 46
320 3.96 36 −56 48 LPs
3.93 40 −46 56 LPi 40
939 4.69 −26 −62 54 LPs
4.56 −26 −48 48 PCu
4.4 −30 −54 52 LPs
0.033 137 4.51 −20 −76 40 PCu
0.002 265 4.61 54 −58 GTm 21
0.041 129 4.97 −60 −18 GTs 22
0.003 240 3.8 62 −10 GTs 22
889 4.92 −48 −70 GOm 37
4.88 −50 −56 GTm 21
4.5 −34 −64 −30 Cerebellum
Hearing signer
837 4.91 42 10 32 GFi 44
4.07 42 56 PMd
602 4.77 −2 66  SMA
4.13 −4 20 52 GFs
1281 4.58 −40 12 32 GFi 44
4.26 −28 62 PMd
4.04 −36 30 16 GFi 45
0.001 311 3.99 30 −62 48 LPs
3.63 38 −48 56 LPi 40
1580 4.88 −30 −56 52 LPs
4.87 −46 −34 40 LPi 40
753 5.23 46 −60 −6 GOm 19
4.06 30 −74 24 GTm 39
1012 4.92 −44 −66 −8 GOm 19
4.88 −54 −62 GTm 37
Deaf > hearing signer
0.001 277 5.08 −62 −20 GTs 22
4.68 −58 −10 GTs 22

3.4

−54

−24

10

L

GTT

41

Cluster P Cluster size Voxel Z Talairach's coordinates

Side Location BA

x

y

z

Deaf
1079 5.63 66  SMA
4.75 24 46 GFd
1531 4.85 −48 14 20 GFi 44
4.64 −36 50 PMd
4.45 −30 28 −2 GFi 47
1153 4.16 48 32 GFi 45
46 46 16 GFm 46
320 3.96 36 −56 48 LPs
3.93 40 −46 56 LPi 40
939 4.69 −26 −62 54 LPs
4.56 −26 −48 48 PCu
4.4 −30 −54 52 LPs
0.033 137 4.51 −20 −76 40 PCu
0.002 265 4.61 54 −58 GTm 21
0.041 129 4.97 −60 −18 GTs 22
0.003 240 3.8 62 −10 GTs 22
889 4.92 −48 −70 GOm 37
4.88 −50 −56 GTm 21
4.5 −34 −64 −30 Cerebellum
Hearing signer
837 4.91 42 10 32 GFi 44
4.07 42 56 PMd
602 4.77 −2 66  SMA
4.13 −4 20 52 GFs
1281 4.58 −40 12 32 GFi 44
4.26 −28 62 PMd
4.04 −36 30 16 GFi 45
0.001 311 3.99 30 −62 48 LPs
3.63 38 −48 56 LPi 40
1580 4.88 −30 −56 52 LPs
4.87 −46 −34 40 LPi 40
753 5.23 46 −60 −6 GOm 19
4.06 30 −74 24 GTm 39
1012 4.92 −44 −66 −8 GOm 19
4.88 −54 −62 GTm 37
Deaf > hearing signer
0.001 277 5.08 −62 −20 GTs 22
4.68 −58 −10 GTs 22

3.4

−54

−24

10

L

GTT

41

Abbreviations: GF, fusiform gyrus; GFd, medial frontal gyrus; GFi, inferior frontal gyrus; GFm, middle frontal gyrus; GOi, inferior occipital gyrus; GOm, middle occipital gyrus; GpoC, postcentral gyrus; GprC, precentral gyrus; Gsm, supramarginal gyrus; GTi, inferior temporal gyrus; GTs, superior temporal gyrus; LPi, inferior parietal lobule; LPs, superior parietal lobule; PCu, precuneus; PMd, dorsal premotor cortex; SMA, supplementary motor area. BA, Brodmann's areas. All P values are corrected for multiple comparisons. Height threshold, Z = 3.09, P = 0.001. L, left; R, right.

## Discussion

The present study showed that the neural substrates of audio-visual integration during lip-reading in hearing subjects overlap with the substrates underlying cross-modal plasticity due to auditory deprivation: During the CLOSED matching task, the deaf showed more prominent activation than the hearing subjects in the left planum temporale (PT). During dot-motion matching, the deaf subjects showed greater activation in the right PT. Sign-related motion, with or without a lexical component, activated the left PT of the deaf signers more than in the hearing signers. These areas showed a lip-reading effect in hearing subjects. Hence, the PT is the common substrate of audio-visual integration during lip-reading in hearing subjects and of audio-visual cross-modal plasticity in deaf subjects.

In this study, we used the difference between the OPEN and CLOSED conditions as a marker of phonetic speech perception (lip-reading effect). We assumed that the OPEN condition carried phonetic information even if the subjects did not understand which vowel was presented, whereas the CLOSED condition did not. This is based on the fact that visual phonetics shown by lip movements can influence speech perception without any understanding of what is presented in each modality (the McGurk effect) (McGurk and MacDonald, 1976; Driver, 1996). We used phonetic discrimination by visual speech (OPEN) to exclude the influence of the lexicon (Bernstein et al., 2002). To control for the perception of lip movements and working memory processes, a non-phonetic lip-movement (CLOSED) condition was included. Thus, the OPEN–CLOSED comparison reveals the phonetic perception of visual speech that is essential for lip-reading. In the hearing group, during the OPEN condition compared with the CLOSED condition, more prominent activation was seen in the GTs, along the STS bilaterally and in the left lateralized frontal cortex including Broca's area, which is consistent with previous results (Burt and Perrett, 1997; Campbell et al., 2001; Olson et al., 2002; Calvert and Campbell, 2003).

### Broca's Area

Functional neuroimaging studies revealed activity in Broca's area during prepositional speech tasks, including reading words (Price et al., 1994), word generation (McCarthy et al., 1993), decoding syntactically complex sentences (Just et al., 1996) and phoneme discrimination (Zatorre et al., 1992, 1996). Auditory phonemic-discrimination activated secondary auditory cortices bilaterally as well as Broca's area (Zatorre et al., 1992, 1996). Zatorre et al. (1996) proposed that Broca's area is recruited for fine-grained phonetic analysis by means of articulatory decoding. The phonetic analysis of speech depends not only on auditory information but also on access to information about the articulatory gestures associated with a given speech sound (Liberman and Whalen, 2000). This access might be accomplished through a ‘mirror’ system, including Broca's area, which forms a link between the observer and the actor (Rizzolatti and Arbib, 1998; Iacoboni et al., 1999). The mirror system might not be confined to Broca's area (Bucchino et al., 2001). Lip-reading enhances the motor excitability of the left primary motor cortex, presumably through input from premotor areas (Watkins et al., 2003). This could account for the activation of other left-lateralized prefrontal cortical structures, such as BA 44/6 and SMA, as parts of the mirror system.

### Planum Temporale

As expected, the OPEN–CLOSED condition activated the part of the GTs posterior to the transverse temporal gyrus (Heschl's gyrus), corresponding to the PT. Anatomically, the anterior border of the PT is the sulcus behind Heschl's gyrus and the medial border is the point where the PT fades into the insula. The posterior border is the ascending and descending rami of the Sylvian fissures (Westbury et al., 1999). Functionally, the left PT is involved in word detection and generation, due to its ability to process rapid frequency changes (Schwartz and Tallal, 1980; Belin et al., 1998b). The right homologue is specialized for the discrimination of melody, pitch and sound intensity (Zattore et al., 1994; Belin et al., 1998a). Activation of the PT by the OPEN–CLOSED comparison is consistent with previous work (Calvert et al., 1997), and confirms that phonetic information encoded in lip movements (during the OPEN condition) activates the PT. Furthermore, the temporal cortical activation was limited during the CLOSED condition in this study. This supports a previous study, which found that non-phonetic lower-face movement did not activate the temporal cortex (Calvert et al., 1997). Hence, in hearing subjects, the activation of the PT is related to the phonetic information encoded by lip movements, and thus the perception of physical visual speech stimuli (visual phonetics) (Bernstein et al., 2002). Figure 6 shows that the area for the lip-reading in the hearing (blue) extended to the posterior STS area, more prominent on the left. This is consistent with the previous report by Puce et al. (1998) showing that the mouth movement activates the posterior portion of the STS. The more prominent posterior extension on the left than the right is consistent with the lesion study (Campbell et al., 1986): a patient with a left occipitotemporal lesion cannot lip-read whereas another patient with the occipitotemporal lesion on the right did not show deficit in lip-reading. Hence the posterior STS on the left may be functionally related to the nearby PT thought to be involved in lip-reading.

In the deaf group, a similar, but less significant, activation pattern in the PT was found in response to visual phonetics, and hence there is a lip-reading effect (Figs 2, 4 and 5). This might reflect the fact that, despite being profoundly deaf, the deaf subjects in the present study have undergone training in lip-reading at school.

### The Effects of Auditory Deprivation

The results also indicate that the neural substrates of the cross-modal plasticity resulting from auditory deprivation lie within the PT. The PT in hearing subjects did not respond to non-phonetic lip movements (CLOSED), dot movement and sign-related hand movement without lexical meaning, whereas the PT of the deaf subjects responded to all these conditions. However, the response of the PT to visual phonetics (OPEN–CLOSED) was seen in the hearing subjects. As these results were found for both hearing signers and hearing non-signers, the ability to sign cannot explain this difference. Similarly, visual phonetics cannot explain these differences because the group difference was seen during the CLOSED condition. Instead, these findings suggest the PT is the common substrate of audio-visual integration during lip-reading (OPEN–CLOSED) in hearing subjects and audio-visual cross-modal plasticity during non-phonetic lip (CLOSED) or hand movements in deaf subjects.

#### Lip Movement and Dot Movement

This study confirmed the results of Finney et al. (2001), who showed that visual stimulation using moving dot patterns activated the right PT of deaf subjects. This finding suggests that the auditory cortex in deaf subjects is recruited for the processing of purely visual stimuli with no lexical component. The right PT is activated by sound motion (Baumgart et al., 1999), probably reflecting the dominance of the right hemisphere for the processing of spatial information. However, in deaf subjects, the left PT was active in response to face movement, which was not seen in hearing subjects. The left-lateralized activity might be related to the notable anatomical and functional asymmetry seen in auditory processing.

Anatomically, the PT is significantly larger in the left hemisphere, with increased cell size and density of neurons. Furthermore, there are more functionally distinct columnar systems per surface unit in the left than the right PT (Galuske et al., 2000). Functionally, the left auditory cortical areas that are optimal for speech discrimination, which is highly dependent on rapidly changing broadband sounds, have a higher degree of temporal sensitivity. By contrast, the right cortical counterpart has greater spectral sensitivity, and thus is optimal for processing the tonal patterns of music, in which small and precise changes in frequency are important (Zatorre et al., 2002). Thus, due to auditory deprivation, in deaf subjects the processing of the rapid temporal alternations of mouth movements might be directed towards the left PT, whereas processing of the global motion of moving dots might be performed in the right PT. This functional asymmetry might also explain why the area of the right PT that was more active in deaf than hearing subjects during the dot-discrimination task showed less overlap with the area activated by OPEN versus CLOSED conditions.

#### Sign

The sign-word discrimination and non-discrimination conditions revealed more prominent activation in the left PT of the deaf signers than hearing signers. This is consistent with Petitto et al. (2000), who found that meaningless but linguistically organized phonetic-syllabic units in sign, as well as real signs, activated the bilateral GTs of deaf subjects. They concluded that semantic content is not an important determinant of the activity of the GTs. This study revealed that the PT was activated by not only signs and pseudo-signs, but also their upside-down reversed moving images. Thus, the linguistic component of the signs might not be causing the PT activation in the deaf subjects. The sign word-discrimination and non-discrimination tasks yielded a similar activation pattern in the left PT: both tasks activated the left PT of the deaf signers, but deactivated that of the hearing signers (Fig. 5). Furthermore, when the sign-word discrimination was compared with the non-discrimination condition, the bilateral frontal cortex was activated, but the temporal cortical areas were not (Fig. 7).

We saw a marked overlap of the enhanced activation in the left PT during sign presentation and face discrimination in the deaf subjects compared with the hearing signers. This finding suggests that the activation of the left PT in the deaf during sign presentation relates neither to sign-language comprehension nor linguistic processing. Instead, the left PT activation might be related to the analysis of the temporal segmental aspects of hand or lip movements. Lip-reading accuracy correlated negatively with sign-language usage in the deaf (Geers and Moog, 1989; Moores and Sweet, 1991; Bernstein et al., 1998), suggesting competition between the processes. Hence, amodal processing of rapid temporal alternation might occur in the left PT, the input to which might be visual or auditory. Although cross-modal plastic changes in the PT are not specific to linguistic processing itself, as shown in the present study, it might promote the sublexical processing of signs.

#### Cross-modal Plasticity in the Deaf and Lip-reading in the Hearing Share the PT

This study revealed that the primary auditory cortex (A1) is not activated by visual phonetics (lip-reading) in the hearing, which was consistent with a previous study (Bernstein et al., 2002). We also found that the plastic changes due to auditory deprivation activated the PT, as did lip-reading in the hearing subjects; the A1 was also spared. In deaf subjects, higher-order auditory areas become activated by visually presented sign language (Nishimura et al., 1999; Petitto et al., 2000). When the auditory nerve is stimulated by cochlear implants, the contralateral A1 becomes active (Lee et al., 2001). The higher-order cortical areas become progressively activated only after increased hearing experience with cochlear implants (Giraud et al., 2001). Therefore, cross-modal reorganization is likely to occur only in the higher-order auditory areas, consistent with the present findings. A previous electrophysiological study revealed no evidence of cross-modal plasticity in the A1 in congenitally deaf cats (Kral et al., 2003). They attributed this to the different inputs that the A1 and the auditory-association cortices receive. The primary auditory cortex receives its main input from the lemniscal auditory pathway, whereas the higher-order auditory cortices receive mainly cortical and extralemniscal thalamic inputs. The extralemniscal thalamic nuclei also receive projections from other modalities (Aitkin et al., 1978, 1981; Robards, 1979; Shore et al., 2000; Schroeder et al., 2001). The sensory representation is determined by the competitive balance of inputs from different modalities (Rauschecker, 1995); therefore, these findings suggest that cross-modal plasticity occurs in the PT.

In conclusion, the cross-modal plastic changes as a result of auditory deprivation might be mediated, at least in part, by the neural substrates of cross-modal integration in the hearing.

This study was supported in part by a Grant-in Aid for Scientific Research B#14380380 (to N.S.) from the Japan Society for the Promotion of Science, and Special Coordination Funds for Promoting Science and Technology from the Ministry of Education, Culture, Sports, Science and Technology, of the Japanese Government.

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## Author notes

1National Institute for Physiological Sciences, Okazaki, Japan, 2Biomedical Imaging Research Center, Fukui University, Fukui, Japan, 3JST (Japan Science and Technology Corporation)/RISTEX (Research Institute of Science and Technology for Society), Kawaguchi, Japan, 4Department of Image-based Medicine, Institute of Biomedical Research and Innovation, Kobe, Japan, 5Department of Education, Fukui University, Fukui, Japan, 6 Department of Education, Kanazawa University, Kanazawa, Japan and 7Graduate School of Human and Environmental Studies, Kyoto University, Kyoto, Japan