‘‘What’’ and ‘‘where’’ in the human auditory system

Claude Alain*†‡, Stephen R. Arnott*†, Stephanie Hevenor*, Simon Graham*§¶, and Cheryl L. Grady*†储
*The Rotman Research Institute, Baycrest Centre for Geriatric Care, Toronto, ON, Canada M6A 2E1; †Department of Psychology and ¶Department of Medical
Biophysics, University of Toronto, Toronto, ON, Canada M8V 2S4; §Imaging兾Bioengineering Research, Sunnybrook and Womens’ College Health Sciences
Centre, Toronto, ON, Canada M4N 3M5; and 储Faculty of Medicine (Psychiatry), University of Toronto, Toronto, ON, Canada M5S 1A8

Edited by P. S. Goldman-Rakic, Yale University School of Medicine, New Haven, CT, and approved August 10, 2001 (received for review April 27, 2001)

The extent to which sound identification and sound localization three-dimensional (3D) auditory environment was created by
depend on specialized auditory pathways was examined by using using synthesized sounds with appropriate free-field acoustic
functional magnetic resonance imaging and event-related brain cues (15, 16). Such a design has proven to be effective for imaging
potentials. Participants performed an S1–S2 match-to-sample task the neural substrates involved in localizing auditory events
in which S1 differed from S2 in its pitch and兾or location. In the pitch (9, 17).
task, participants indicated whether S2 was lower, identical, or
higher in pitch than S1. In the location task, participants were asked Methods
to localize S2 relative to S1 (i.e., leftward, same, or rightward). Fifteen young adults (aged between 21 and 31 years; four males)
Relative to location, pitch processing generated greater activation participated in the study. All were right-handed and reported
in auditory cortex and the inferior frontal gyrus. Conversely, normal hearing. Each participant signed an informed consent
identifying the location of S2 relative to S1 generated greater form approved by the University of Toronto Human Subject
activation in posterior temporal cortex, parietal cortex, and the Review Committee. Data from three participants were excluded
superior frontal sulcus. Differential task-related effects on event- because of head motion greater than 1 mm during the experi-
related brain potentials (ERPs) were seen in anterior and posterior ment. ERPs were recorded in a separate session and were
brain regions beginning at 300 ms poststimulus and lasting for obtained from those 12 participants that were included in the
several hundred milliseconds. The converging evidence from two fMRI analysis.
independent measurements of dissociable brain activity during
identification and localization of identical stimuli provides strong Stimuli and Tasks. Stimuli consisted of five synthesized two-octave
support for specialized auditory streams in the human brain. These band noise bursts starting with a center frequency of 2000 Hz and
findings are analogous to the ‘‘what’’ and ‘‘where’’ segregation of stepping up four times. Stimulus duration was 500 ms including
visual information processing, and suggest that a similar functional 5-ms rise兾fall time. Stimuli were generated digitally with 16-bit
organization exists for processing information from the auditory resolution and a sampling rate of 50 kHz, passed through a
modality. digital-to-analogue converter, and then low pass filtered at 10
kHz by using an anti-aliasing filter (Tucker-Davis Technology,
Gainesville, FL). Stimuli were presented at 85 dB sound pressure
A uditory scene analysis involves identifying the content
(‘‘what’’) and the location (‘‘where’’) of sounds in the
environment. Evidence from anatomical and neurophysiological
level (SPL) by means of circumaural, fMRI-compatible head-
phones (Avotec, Jensen Beach, FL), acoustically padded to
studies in non-human primates (1–5) suggests that identification suppress scanner noise by 25 dB. Stimuli were presented at five
and localization of auditory events may be functionally segre- possible azimuth locations relative to straight ahead (⫺90°, ⫺45°,
gated in specialized auditory streams. Combining anatomical 0°, ⫹45°, ⫹90°). Virtual 3D sources were synthesized by using a

PSYCHOLOGY
and electrophysiological recording methods in non-human pri- head-related transfer function that replicated the acoustic ef-
mates, Romanski et al. (5) have recently identified two separate fects of the head and ears of an average listener (18).
auditory streams that originate in caudal and rostral auditory Participants performed a delayed match-to-sample task in
cortex, respectively, and project to different regions within the which the first acoustic stimulus (S1) was held in memory (for
frontal lobe. The functional significance of these separate path- 500 ms) for comparison with the second (S2) stimulus. Each trial
ways has not been determined, although they suggest functional began with a binaural warning tone (1000 Hz, 500-ms duration,
dissociations for auditory processes analogous to the ‘‘what’’ and 5-ms rise兾fall time). After a delay of 1000 ms, a noise burst (S1)
‘‘where’’ or ventral and dorsal cortical information streams for was presented at one of three possible locations and one of three
identifying and localizing visual (6, 7) and somatosensory (8) possible pitches that excluded the two extreme locations and the
stimuli. two extreme pitches. A second noise burst (S2) was presented
Auditory neuroimaging studies employing positron emission 500 ms after the offset of S1. S2 was presented at one of three
tomography or functional magnetic resonance imaging (fMRI) possible and equally probable locations that included the same
have revealed enhanced blood flow in parietal areas during location of S1 and the two adjacent locations (immediately
sound localization (9–11). In comparison, tasks requiring indi- rightward and leftward). For example, if S1 was presented at 45°
viduals to make tone discriminations (12) or identify auditory to left of center, S2 could have been presented along the azimuth
stimuli (e.g., words or environmental sounds) show enhanced at 90° left of center, 45° left of center, or 0°. Similarly, the
activation in inferior frontal cortex (13, 14). Although these frequency of S2 was either identical or adjacent (lower or higher)
results suggest that the processing of sound identity and sound than S1. The likelihood of having the second stimulus lower,
location is functionally separable, the segregation in auditory
information processing has yet to be demonstrated within the
same individuals when using the same set of stimuli. This paper was submitted directly (Track II) to the PNAS office.

The present study was designed to directly test, by using fMRI Abbreviations: BOLD, blood oxygenation level-dependent; 3D, three-dimensional; ERP,
and event-related brain potentials (ERPs), the hypothesis that event-related brain potentials; fMRI, functional magnetic resonance imaging.

specialized streams exist in humans for processing sound identity

‡Towhom reprint requests should be addressed at: Rotman Research Institute, Baycrest
Centre for Geriatric Care, 3560 Bathurst Street, Toronto, ON, Canada M6A 2E1. E-mail:
and sound location. Young adults were presented with the same calain@[Link].
stimuli but were required to perform two different tasks: a pitch The publication costs of this article were defrayed in part by page charge payment. This
(what) and a location (where) judgment task. Because free-field article must therefore be hereby marked “advertisement” in accordance with 18 U.S.C.
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auditory stimulation is not possible within the scanner, a virtual §1734 solely to indicate this fact.

[Link]兾cgi兾doi兾10.1073兾pnas.211209098 PNAS 兩 October 9, 2001 兩 vol. 98 兩 no. 21 兩 12301–12306

 Fig. 1. The first two columns (Left) show the group mean activation during the pitch and location tasks vs. rest, respectively. The color scale below is based
on t values ranging from 6 to 15 (P ⬍ 0.00001). The rightmost column shows the difference in brain activation between the location and the pitch tasks. The color
scale for this image is based on t values ranging from 2.9 to 7 (P ⬍ 0.01). Areas with greater activity during the location task are shown in blue, whereas those
more active during the pitch task are illustrated in orange and yellow. The right hemisphere is shown on the left side of the images. Slice locations are indicated
by green lines on the accompanying midline sagittal image.

leftward, identical, higher, or rightward was equally probable. location or of the same pitch. Participants kept their eyes closed
The intertrial interval was controlled by the participant, whose during the scanning.
response initiated the presentation of the next trial 1000 ms later. The tasks carried out during the ERP measurements were
In the pitch task, participants indicated by pressing one of three similar to those used during the fMRI procedure. For each
buttons whether S2 was lower, identical, or higher in pitch than condition, participants were given five blocks of 60 trials. The
S1, regardless of its location. Similarly, in the spatial discrimi- stimuli were the same as those used for the fMRI experiment and
nation task, participants indicated by pressing one of three the trials themselves were identical (i.e., warning tone, then S1,
buttons whether S2 was at a leftward, an identical, or a rightward and then S2 500 ms later). The order of conditions was coun-
position relative to S1, regardless of changes in pitch. The stimuli terbalanced across participants.
were identical in both conditions; only the task instructions
differed. Participants performed each designated task (e.g., fMRI Procedure. Participants’ regional cerebral activity was as-
location comparisons) for 30 s followed by a 30-s rest period in sessed by using a 1.5-T Signa MR scanner with a standard head
which no stimuli were presented. This on兾off sequence was coil (CV兾i hardware, LX8.3 software; General Electric Medical
repeated five times for a total duration of 5 min. Three to four Systems, Waukesha, WI). Each scan sequence consisted of five
pitch and location sequences were performed on each partici- 30-s task blocks alternating with 30-s blocks in which no sound
was presented. Functional imaging was performed to measure
pant, which alternated over the course of the experiment. The
brain activation by means of the blood oxygenation level-
order of conditions (pitch or location) was counterbalanced
dependent (BOLD) effect (19) with optimal signal contrast.
across participants.
Eighteen axial slices 7 mm thick were obtained. Functional scans
To ensure that changes in brain activation were not due to were obtained by using a single shot T2*-weighted pulse se-
differences in task difficulty, the pitch separation was adjusted quence with spiral readout, offline gridding, and reconstruction
for each individual such that they performed equally well in both (ref. 20; TR ⫽ 2000 ms, TE ⫽ 40 ms, flip angle 80°, 90 ⫻ 90
tasks. This separation varied between 3 and 10% (mean 6.6 ⫾ effective acquisition matrix). For each participant, standard
2.4%). Responses and latencies were obtained by using two volumetric anatomical MRI was performed before functional
fMRI-compatible response pads (Lightwave Technologies, Sur- scanning by using a standard 3D T1-weighted pulse sequence
rey, BC, Canada), each containing two buttons side-by-side. On (TR ⫽ 12.4 ms, TE ⫽ 5.4 ms, flip angle 35°, 22 ⫻ 16.5 field of
each trial, participants indicated whether the sound was leftward view, 256 ⫻ 192 acquisition matrix, 124 axial slices 1.4 mm thick).
or lower in pitch by pressing the extreme left button of the pad Data processing and analyses were performed by using Anal-
in their left hand with their left middle finger (button 1). They ysis of Functional NeuroImages (AFNI) software (21, 22). Time
pressed the rightmost button (button 4) on the pad in their right series data were spatially coregistered to correct for head motion
hand with their right middle finger for sounds that were right- by using a 3D Fourier transform interpolation, and detrended to
ward or higher in pitch. Lastly, they simultaneously pressed the a constant reference scan by using a fifth-order polynomial.
remaining buttons (2 and 3) on each response pad with their left Percent changes in signal intensity with respect to rest were
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and right index fingers when the sounds were at the same analyzed by using voxel-wise correlations of the location and

12302 兩 [Link]兾cgi兾doi兾10.1073兾pnas.211209098 Alain et al.

 Table 1. Brain regions where there were significant differences between pitch and location
Pitch vs.
Task vs. Rest Location

Region x y z L-R P-R (P-R)-(L-R)

Location ⬎ Pitch
Frontal
Right superior frontal gyrus (BA 6兾8) 25 5 59 6.72** 0.88 ⫺4.13*
Left middle frontal gyrus (BA 8) ⫺26 3 48 6.10** ⫺1.15 ⫺5.13**
Parietal
Right superior parietal (BA 7) 15 ⫺76 49 10.39** 3.74* ⫺4.70*
Left precuneus (BA 7) ⫺12 ⫺72 49 10.54** 0.82 ⫺6.87**
Right inferior parietal (BA 40) 25 ⫺45 39 10.29** 2.86 ⫺5.25**
Left inferior parietal (BA 40) ⫺27 ⫺48 41 17.23** 11.34** ⫺4.16*
Temporal
Right posterior middle temporal gyrus (BA 21) 58 ⫺53 1 7.90** 1.30 ⫺4.66**
Left posterior middle temporal gyrus (BA 37) ⫺56 ⫺59 ⫺1 3.69* ⫺2.51 ⫺4.38*
Occipital
Left superior occipital gyrus (BA 19) ⫺32 ⫺74 37 ⫺1.09 ⫺6.96** ⫺4.15*
Pitch ⬎ Location
Frontal
Right inferior frontal gyrus (BA 45) 45 25 10 5.76** 11.44** 4.02*
Right inferior frontal gyrus (BA 44) 51 18 12 4.71** 9.68** 3.52*
Temporal
Right superior temporal gyrus (BA 22) 60 ⫺21 2 4.45** 9.25** 3.39*
Left superior temporal gyrus (BA 22) ⫺53 ⫺20 6 20.13** 26.04** 4.18*
Left superior medial temporal gyrus (BA 22) ⫺37 ⫺25 5 3.79* 9.47** 4.02*
Right primary auditory cortex (BA 41) 42 ⫺21 11 5.09** 8.47** 4.18*
Occipital
Right cuneus (BA 19) 17 ⫺87 23 ⫺13.14** ⫺5.20** 5.62**
Left cuneus (BA 18) ⫺11 ⫺97 3 ⫺15.94** ⫺6.03** 7.01**

The Talairach coordinates are based on the peak voxel in t value. BA, Brodmann’s Area according to the atlas of Talairach and
Tournoux (24); L-R, t value for Location vs. Rest; P-R, t value for Pitch vs. Rest; (P-R)-(L-R), t value for Location by Pitch interaction (negative
values represent greater activation during the Location vs. the Pitch condition, positive values represent greater activation during the
Pitch vs. the Location condition). *, P ⬍ 0.01; **, P ⬍ 0.001.

pitch time series with square-wave reference vectors (23) shifted The analysis epoch included 200 ms of prestimulus activity and
to account for the delay in hemodynamic response. The statis- 3000 ms of poststimulus activity. Trials contaminated by eye

PSYCHOLOGY
tical cut-off for activation was set at P ⬍ 0.001 or lower, blink or excessive peak-to-peak deflection (⫾150 ␮V) at the
uncorrected. The minimum cluster size was 10 mm3 with a radius electrodes not adjacent to the eyes were automatically rejected
of 2 mm. This produced two activation images per participant, before averaging. The ERPs were then averaged separately for
one for location vs. rest and one for pitch vs. rest. These each site, stimulus type, and listening condition. ERPs were
activation images were then transformed into Talairach coordi- digitally lowpass filtered to attenuate frequencies above 12 Hz.
nates (21, 22, 24) and smoothed with a Gaussian filter of 6 mm For each individual average, the ocular artifacts (e.g., blinks and
full width at half maximum (FWHM) to increase the signal-to- lateral movements) were removed by means of ocular source
noise ratio. The latter step was performed to facilitate the components, using BR A IN ELECTR ICA L SOURCE ANA LYSIS
subsequent group analysis, which consisted of a random effect, (BESA) software (25). ERP waveforms were quantified by
voxel-wise two-factor ANOVA with tasks (location and pitch) as computing mean values in selected latency regions, relative to
within-subject factors. Because the ANOVA was performed on the mean amplitude of the 200-ms prestimulus activity. All
the task vs. rest contrast images, the degrees of freedom were measurements were subjected to repeated measures ANOVA
based on the number of subjects rather than on the number of with task (pitch and location) and electrodes (CP1, CP2, FT9,
scans. For the comparison between pitch and location condi-
FT10) as within-subject factors. Scalp topographies using the 61
tions, the statistical cut-off was set at P ⬍ 0.01 and the minimum
electrodes (omitting the periocular electrodes) were statistically
cluster size was 10 mm3 with a radius of 2 mm.
analyzed after scaling the amplitudes to eliminate amplitude
Recording and Analysis of ERPs. The electroencephalogram (EEG)
differences between conditions (26). The original degrees of
was recorded from an array of 64 electrodes including those from freedom for all analyses are reported throughout the paper. Type
the standard 10–20 placement. Vertical and horizontal eye I errors associated with inhomogeneity of variance were con-
movements were recorded with electrodes at the outer canthi trolled by decreasing the degrees of freedom using the Green-
and at the superior and inferior orbit. Electrophysiological house-Geisser epsilon (␧), and the probability estimates are
signals were digitized continuously (bandpass 0.05–50 Hz; 250 based on these reduced degrees of freedom.
Hz sampling rate) by means of NeuroScan SynAmps and stored
for offline analysis. During the recording, all electrodes were Results
referenced to the midline central electrode (Cz); for data fMRI Experiment. There were no differences in accuracy between
analysis, they were re-referenced to an average reference and the the two tasks: participants correctly judged the pitch or the
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electrode Cz was reinstated. location of S2 in 72% and 70% of the trials, respectively.

Alain et al. PNAS 兩 October 9, 2001 兩 vol. 98 兩 no. 21 兩 12303

 extending anteriorly to auditory association cortices on the
supratemporal plane. We also found concurrent increases in
BOLD signal in the right inferior frontal gyrus. These differ-
ences in brain activation were seen primarily in the right
hemisphere (see Fig. 2 and Table 1). Conversely, selectively
processing sound locations was associated with greater bilateral
activation in posterior temporal areas, and in inferior and
superior parietal cortices compared with the pitch judgment
(Fig. 2, Table 1). Importantly, there was a parallel increase in
BOLD signal in the right superior frontal sulcus, an area very
similar to that observed during a visual spatial location task (27).
Another way of examining task-specific changes is to see
whether areas with greater increases of activity in a given task
also show larger correlations among their activity measures
during that task. To explore these functional correlations, we
compared the time course of changes in BOLD signal in the peak
voxels of four brain areas; primary auditory cortex, superior
parietal cortex, and inferior and superior prefrontal gyrus. These
regions of interest were chosen because (i) they were proposed
to be part of a dual pathway model (5, 27) and (ii) they were
differentially active during the pitch and location tasks. Across
participants, the time series for each condition were averaged
into a 60-s sequence containing 30 data points (15 ON and 15
Fig. 2. Three-dimensional pattern of cortical activation to highlight differ- OFF). Pair-wise correlation coefficients were then computed
ences between pitch and location discrimination tasks. The right hemisphere among these group mean time courses of BOLD signal changes
is foremost in the figure and part of the temporal lobe has been removed to for the four regions.
show activation in temporal and inferior frontal cortices. The enhanced signal
in posterior visual areas during the location task was caused by greater signal
Table 2 shows the correlation matrices of interregional cor-
reduction during the pitch task rather than increased activity during the relations for the location and the pitch task. During the location
location task. See Table 1 for atlas coordinates and statistical measures for task, the enhanced BOLD signal in auditory cortex was corre-
these areas. The color scale for this image is based on t values ranging from 2.9 lated with that of parietal cortex. Furthermore, there was a
to 7 (P ⬍ 0.01). significant correlation between the BOLD signal in parietal
cortex and that of the superior frontal gyrus. Importantly, there
was no significant correlation between BOLD signal in auditory
Similarly, response latency did not significantly differ between cortices and inferior frontal cortex during the location task. In
pitch (1121 ⫾ 308 ms) and location (1069 ⫾ 204 ms) judgments. contrast, we found a significant correlation between the ob-
For the group as a whole, we calculated the mean percent served changes in BOLD signal in auditory cortices and that of
increase in BOLD signal intensity from rest for the pitch and the inferior prefrontal gyrus during the pitch task. There was no
location conditions separately (Fig. 1). In both location and pitch significant correlation between activity in the parietal cortex and
tasks, there was bilateral activity enhancement in primary and superior frontal gyrus during the pitch task. This pattern of
secondary auditory cortices, inferior and superior parietal cor- functional correlations therefore supports distinct ventral and
tices, and the superior and inferior frontal gyri (Table 1). Both dorsal networks active during nonspatial and spatial auditory
auditory discrimination tasks also were associated with a de- tasks, respectively.
crease in BOLD signal in visual cortex.
Specificity of brain activity for localizing and identifying ERP Experiment. As in the fMRI experiment, there were no
auditory events was determined by directly comparing the differences in accuracy between the two tasks. Accuracy in both
changes in hemodynamic response obtained during the pitch task the pitch and the location tasks was 82%. However, participants
vs. rest with changes in activity measured during the location task were slower in the pitch (1026 ⫾ 162 ms) than in the location task
vs. rest. Relative to the location task, pitch judgment was [954 ⫾ 134 ms; F(1,11) ⫽ 5.52, P ⬍ 0.05].
associated with greater activation in primary auditory cortices, Fig. 3 shows event-related brain potentials elicited during the

Table 2. Correlations between BOLD signal changes in regions of interest

Right superior Right superior Right inferior Right superior
frontal gyrus (BA 6兾8) parietal cortex (BA 7) frontal gyrus (BA 45) temporal gyrus (BA 22)

Location task
Right superior frontal gyrus (BA 6兾8) 1.000
Right superior parietal cortex (BA 7) 0.670** 1.000
Right inferior frontal gyrus (BA 45) 0.014 0.034 1.000
Right superior temporal gyrus (BA 22) 0.291 0.469** ⫺0.174 1.000
Pitch task
Right superior frontal gyrus (BA 6兾8) 1.000
Right superior parietal cortex (BA 7) 0.064 1.000
Right inferior frontal gyrus (BA 45) ⫺0.165 0.573** 1.000
Right superior temporal gyrus (BA 22) ⫺0.473** 0.532** 0.558** 1.000
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*, P ⬍ 0.05; **, P ⬍ 0.01.

12304 兩 [Link]兾cgi兾doi兾10.1073兾pnas.211209098 Alain et al.

 emission tomography in humans by using a virtual auditory
display (9, 10).
The novel finding of this study is the dissociation of the effects
of task instruction on both fMRI and ERP measures. Processing
pitch information recruited brain areas that were primarily
distributed in the ventral part of the brain, whereas sound
localization recruited areas that were primarily distributed in
dorsal regions. The dissociation in fMRI signals was paralleled
by task-related changes in ERPs. Differences in both BOLD
signal change and in ERP amplitude during the pitch and
location tasks suggest that auditory information processing may
be specialized into nonspatial and spatial domains. The present
study supports and extends previous neuroimaging studies by
providing the first direct evidence that the neural systems
involved in identifying and localizing auditory objects are func-
tionally and neuroanatomically segregated based on task de-
mands even when stimuli are identical across tasks. Our findings
are consistent with animal models of auditory processing (4, 5,
Fig. 3. Group mean event-related waveforms elicited during the pitch (solid 34–37) in which object identification recruits activation in an-
line) and the location (dashed line) judgment tasks. FT9兾FT10, left and right terior temporal and inferior frontal areas, whereas object loca-
frontal-temporal electrodes; CP1兾CP2, left and right central-parietal electrodes.
tion is mediated by posterior temporal areas, parietal cortex, and
dorsal frontal regions. The observed pattern of BOLD signal
correlations between auditory and inferior prefrontal cortices
two tasks. There were two ERP modulations that distinguished
during the pitch task and between parietal and dorsal prefrontal
neural activity associated with processing pitch and location. The
cortices during the location task is also consistent with the
first occurred between 300 and 500 ms after the presentation of
proposal that sound identification and localization are function-
S1, and consisted of greater positivity of the waveform over ally distinct along ventral and dorsal pathways, respectively.
inferior frontotemporal regions during the pitch task and greater Here, the functional segregation is due to the degree to which
positivity over centroparietal regions during the location task the areas are active in the two tasks (Figs. 1 and 2, Table 1),
[F(1,11) ⫽ 6.86, P ⬍ 0.05]. This task effect on the ERPs may similar to the differential activation of the ventral and dorsal
reflect the online processing and maintenance of S1 in working visual streams by objects and locations (27, 38). There was also
memory for an eventual comparison with S2. The second a significant correlation between temporal and parietal cortex
modulation occurred 300 – 400 ms after S2 was presented during both pitch and location judgment tasks. This finding may
[F(1,11) ⫽ 26.05, P ⬍ 0.001], and showed a similar difference in reflect ‘‘cross talk’’ between ventral and dorsal streams as
the waveforms as was seen for S1 (Fig. 3). This modulation previously suggested for visual stimuli (39) and emphasizes that,
preceded the P3b wave at parietal sites and may reflect the although functional segregation may be an important property of
comparison between the representations of S1 and S2. the brain, higher perceptual and cognitive functions involve
interaction among many brain areas.
Discussion Lastly, there is a remarkable similarity between the prefrontal
The present study was designed to examine the extent to which areas recruited during sound identity and sound location in the

PSYCHOLOGY
processing sound identity and sound location depends on spe- current experiment and those observed in previous anatomical
cialized auditory pathways within the same individuals and using studies (4, 5) and reported during a comparable visual task (27).
similar stimuli. Relative to location processing, pitch processing This result suggests that these ventral and dorsal prefrontal areas
was associated with greater activity in auditory cortices and are involved in representing two distinct types of information
inferior prefrontal gyrus. The inferior prefrontal gyrus has about the environment, ‘‘what’’ and ‘‘where,’’ regardless of which
consistently been shown to be active during the processing of stimulus modality provides that information. Together, these
pitch changes (28), auditory word and tone working memory findings suggest that the segregation of information processing
(29), semantic processing of auditory materials (13), and pho- into ‘‘what’’ and ‘‘where’’ pathways may be a fundamental
neme discrimination (14). Conversely, spatial judgment was property of cortical organization.
associated with greater bilateral activation in posterior temporal
areas, and in inferior and superior parietal cortices compared We thank Rhonda Walcarius and Cathy Nangini for technical support,
with the pitch judgment. This finding provides further evidence and Drs. Helen Mayberg, Randy McIntosh, Terry Picton, and Don Stuss
for helpful comments on an earlier version of the manuscript. We also
that the parietal cortex plays an important role in processing the thank Dr. Gary Glover for the use of his spiral imaging pulse sequence.
spatial relation between consecutive auditory events, consistent This research was supported by grants from the Natural Sciences and
with findings from electrophysiological studies in non-human Engineering Research Council of Canada, the Canadian Institutes for
primates (30), lesion studies in humans (31–33), and positron Health Research, and the Canadian Foundation for Innovation.

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