Common and distinct brain activation to viewing dynamic sequences of face and hand movements.
ABSTRACT The superior temporal sulcus (STS) and surrounding lateral temporal and inferior parietal cortices are an important part of a network involved in the processing of biological movement. It is unclear whether the STS responds to the movement of different body parts uniformly, or if the response depends on the body part that is moving. Here we examined brain activity to recognizing sequences of face and hand movements as well as radial grating motion, controlling for differences in movement dynamics between stimuli. A region of the right posterior STS (pSTS) showed common activation to both face and hand motion, relative to radial grating motion, with no significant difference between responses to face and hand motion in this region. Distinct responses to face motion relative to hand motion were observed in the right mid-STS, while the right posterior inferior temporal sulcus (pITS) and inferior parietal lobule (IPL) showed greater responses to hand motion relative to face motion. These findings indicate that while there may be distinct processing of different body part motion in lateral temporal and inferior parietal cortices, the response of the pSTS is not body part specific. This region may provide input to other parts of a network involved with processing human actions with a high-level visual description of biological motion.
- SourceAvailable from: Chantal Delon-Martin[show abstract] [hide abstract]
ABSTRACT: Perception of biological motions plays a major adaptive role in identifying, interpreting, and predicting the actions of others. It may therefore be hypothesized that the perception of biological motions is subserved by a specific neural network. Here we used fMRI to verify this hypothesis. In a group of 10 healthy volunteers, we explored the hemodynamic responses to seven types of visual motion displays: drifting random dots, random dot cube, random dot cube with masking elements, upright point-light walker, inverted point-light walker, upright point-light walker display with masking elements, and inverted point-light walker display with masking elements. A gradient in activation was observed in the occipitotemporal junction. The responses to rigid motion were localized posteriorly to those responses elicited by nonrigid motions. Our results demonstrate that in addition to the posterior portion of superior temporal sulcus, the left intraparietal cortex is involved in the perception of nonrigid biological motions.NeuroImage 06/2001; 13(5):775-85. · 6.25 Impact Factor
- [show abstract] [hide abstract]
ABSTRACT: We report a functional neuroimaging study with positron emission tomography (PET) in which six healthy adult volunteers were scanned while watching silent computer-presented animations. The characters in the animations were simple geometrical shapes whose movement patterns selectively evoked mental state attribution or simple action description. Results showed increased activation in association with mental state attribution in four main regions: medial prefrontal cortex, temporoparietal junction (superior temporal sulcus), basal temporal regions (fusiform gyrus and temporal poles adjacent to the amygdala), and extrastriate cortex (occipital gyrus). Previous imaging studies have implicated these regions in self-monitoring, in the perception of biological motion, and in the attribution of mental states using verbal stimuli or visual depictions of the human form. We suggest that these regions form a network for processing information about intentions, and speculate that the ability to make inferences about other people's mental states evolved from the ability to make inferences about other creatures' actions.NeuroImage 10/2000; · 6.25 Impact Factor
- [show abstract] [hide abstract]
ABSTRACT: Functional magnetic resonance imaging (fMRI) was used to localize brain areas that were active during the observation of actions made by another individual. Object- and non-object-related actions made with different effectors (mouth, hand and foot) were presented. Observation of both object- and non-object-related actions determined a somatotopically organized activation of premotor cortex. The somatotopic pattern was similar to that of the classical motor cortex homunculus. During the observation of object-related actions, an activation, also somatotopically organized, was additionally found in the posterior parietal lobe. Thus, when individuals observe an action, an internal replica of that action is automatically generated in their premotor cortex. In the case of object-related actions, a further object-related analysis is performed in the parietal lobe, as if the subjects were indeed using those objects. These results bring the previous concept of an action observation/execution matching system (mirror system) into a broader perspective: this system is not restricted to the ventral premotor cortex, but involves several somatotopically organized motor circuits.European Journal of Neuroscience 02/2001; 13(2):400-4. · 3.75 Impact Factor
Common and distinct brain activation to viewing dynamic
sequences of face and hand movements
James C. Thompson,a,⁎Jillian E. Hardee,b,fAnita Panayiotou,d,e
David Crewther,dand Aina Puceb,c,f
aDepartment of Psychology and Krasnow Institute for Advanced Study, George Mason University, MSN 3F5, Fairfax, VA 22030, USA
bCenter for Advanced Imaging, West Virginia University, Morgantown, WV, USA
cDepartment of Radiology, West Virginia University, USA
dBrain Sciences Institute, Swinburne University, Melbourne, Australia
eSchool of Psychological Science, La Trobe University, Melbourne, Australia
fDepartment of Neurobiology and Anatomy, West Virginia University, USA
Received 28 February 2007; revised 17 April 2007; accepted 23 May 2007
Available online 14 June 2007
The superior temporal sulcus (STS) and surrounding lateral temporal
and inferior parietal cortices are an important part of a network
involved in the processing of biological movement. It is unclear
whether the STS responds to the movement of different body parts
uniformly, or if the response depends on the body part that is moving.
Here we examined brain activity to recognizing sequences of face and
hand movements as well as radial grating motion, controlling for
differences in movement dynamics between stimuli. A region of the
right posterior STS (pSTS) showed common activation to both face
and hand motion, relative to radial grating motion, with no significant
difference between responses to face and hand motion in this region.
Distinct responses to face motion relative to hand motion were
observed in the right mid-STS, while the right posterior inferior
temporal sulcus (pITS) and inferior parietal lobule (IPL) showed
greater responses to hand motion relative to face motion. These
findings indicate that while there may be distinct processing of different
body part motion in lateral temporal and inferior parietal cortices, the
response of the pSTS is not body part specific. This region may provide
input to other parts of a network involved with processing human
actions with a high-level visual description of biological motion.
© 2007 Elsevier Inc. All rights reserved.
Keywords: Biological motion; STS; IPL; IFG; fMRI
The ability to accurately interpret another's behavior is critical
for social communication and understanding. Recent neuroimaging
studies have revealed that regions in the inferior frontal, ventral
premotor, lateral temporal and inferior parietal cortex form a
network associated with the perception of human movement
(Beauchamp et al., 2002; Bonda et al., 1996; Buccino et al., 2001,
2004; Grezes et al., 2001; Pelphrey et al., 2005; Puce et al., 1998;
Rizzolatti et al., 2001; Wheaton et al., 2004). A key factor in under-
standing biological movement recognition is to determine if the
movement of different body parts is processed in a uniform manner
within the network, or whether regions in the network respond
differentially to various body parts. Functional magnetic resonance
imaging (fMRI) studies suggest that within the ventral premotor
cortex viewing the movement of different body parts activation is
consistent with the known somatotopy of motor responses in this
region: viewing leg motion activates a region dorsal and posterior to
areas active to viewing face motion, while a region between the two
activates to viewing hand motion (Buccino et al., 2001; Wheaton et
al., 2004). However, it is less clear if there is any organization in
regions making up the biological movement recognition network.
The superior temporal sulcus (STS) makes an important
contribution to the recognition of other people's movements
(Allison et al., 2000; Puce and Perrett, 2003), in particular for
movements of articulated joints (Beauchamp et al., 2002) and for
the body as a whole (Thompson et al., 2005). One role that has
been attributed to the STS is to provide other brain regions in the
network with a visual description of the details of human actions
(Iacoboni et al., 2001). It is not known if this visual description
maintains details of the body part that generated the motion,
implying an intermediate-level representation, or if instead a
higher-level representation is generated which is independent of
moving body part. In a recent study, Pelphrey and colleagues
(2005) described a posterior-to-anterior organization of fMRI
responses to viewing hand, eye, and mouth movement within the
superior temporal sulcus (STS). Other studies have indicated a
different response profile to face and hand motion across the STS
versus parietal cortex (Thompson et al., 2004; Wheaton et al.,
2004). These data suggest that an intermediate-level description is
NeuroImage 37 (2007) 966–973
E-mail address: firstname.lastname@example.org (J.C. Thompson).
Available online on ScienceDirect (www.sciencedirect.com).
1053-8119/$ - see front matter © 2007 Elsevier Inc. All rights reserved.
formed in the STS in which details of the body part generating the
motion are maintained, or that there may be a preference for face
motion in the STS. In contrast, a number of other studies have
demonstrated that the STS also responds to non-biological
geometric objects that move in a manner that implies animacy or
intentionality (Castelli et al., 2000; Blakemore et al., 2003; Schultz
et al., 2005; Martin and Weisberg, 2003). It is possible therefore
that the STS may form a high-level representation of moving
stimuli, rather than maintaining and conveying information about
the body part that generated the motion.
An important consideration when examining brain responses to
motion of different body parts is that patterns of responses might,
in part, be due to differences in the movement kinematics and
dynamics of different body parts (Gentilucci et al., 2001).
Observers are highly sensitive to the dynamic features of biological
movement, and can use abstract temporal cues alone to
discriminate between self- and other-produced actions (Flach et
al., 2004). The importance of dynamic information when viewing
the motion of different body parts is reinforced by findings that
attending to the timing of dynamic sequences of non-biological
visual stimuli can activate the ventral premotor, inferior parietal,
and superior temporal cortex (Coull, 2004; Schubotz and von
Cramon, 2004). Previous studies examining brain responses in the
STS to viewing the movement of different body parts have used
passive viewing (Pelphrey et al., 2005; Puce et al., 1998; Wheaton
et al., 2004), or detection tasks that may have allowed different
strategies for processing mouth and hand movements (Thompson
et al., 2004). Here, we used computer animated stimuli to represent
face, hand and non-biological (control) movement. We asked
subjects to focus on the dynamics of the stimuli and detect target
motion sequences that were identical across stimulus types. If the
STS forms an intermediate representation of biological movement
where details of body part are maintained, we would expect to see
non-overlapping responses to face and hand motion, once move-
ment dynamics and meaning or task-relevance of the stimuli are
In contrast, if there are overlapping responses to face and hand
motion in STS this may suggest a high-level representation of
biological movement that is not body-part dependent.
Materials and methods
Eleven healthy, neurologically normal right-handed subjects
(5 males) aged 22–40 years participated in a study approved by the
Ethics Committee of Swinburne University of Technology,
Melbourne, Australia. All subjects had normal or corrected-to-
Subjects lay supine and viewed movies of face, hand, or radial
grating motion presented through the MRI scanner's control room
window on a screen place at the edge of the patient gurney of the
MRI scanner via a mirror mounted on the quadrature head coil.
Stimuli were presented on a PC running Presentation (Neurobe-
havioral Systems, Albany CA).
Each stimulus category was presented in blocks of mouth,
finger, or radial grating motion each lasting 40 s alternating with
blocks of fixation lasting 20 s. During fixation, subjects fixated on
a red cross presented in the center of a grey screen. Pseudorandom
sequences of short and long movements within each block were
presented in each block (Fig. 1). Both the left and right hands were
used as stimuli. A total of 4 runs were presented, with 2 blocks of
each stimulus per run. Motion consisted of short (212 ms) or long
(696 ms) movements of a face with opening mouth, an extending
index finger on a hand, or inwardly moving rings of a radial motion
stimulus (Fig. 1, top panel). Between each movement a static
version of a face with closed mouth, a closed hand, or a radial
pattern were presented with a random ISI of 1–3 s. Subjects
responded with a button press when a target motion sequence of
short–short–long movements occurred, and with another button
press when they observed a second target sequence of long–long–
short movements (Fig. 1, bottom panel). Successive target
sequences were separated by at least one non-target motion
stimulus triplet. A total of 28 target sequences were presented for
each stimulus type (mouth, hand, radial pattern).
Stimuli were created using Poser 4.0 (Curious Labs, Santa
Cruz, CA). The face stimulus covered a total area approximately
3°×4.5° of horizontal and by vertical visual angle, respectively.
The hand and the radial gratings covered approximately 3°×3° of
horizontal and vertical visual angle. Movies of face and hand
motion were created by first initializing the face with mouth closed
or the index finger curled in a closed hand (Fig. 1). A keyframe of
the face with mouth fully open or finger fully extended was then
created as the middle frame of 7 (short) or 23 (long) keyframes
using a rate of 33 frames/s. Smooth motion from the initial
keyframe to the middle keyframe and back was then performed
with spline interpolation. For the radial grating stimuli, each ring
was approximately 0.16° wide, and only the inner six of the nine
rings moved. The color of the darkest and lightest rings matched
the minimum and maximum RGB value of the face and hand
stimuli. Gratings moved outwards for either 7 or 23 frames at
MRI data acquisition and analysis
A total of 160 volumes/condition were collected in four
imaging runs in each subject. Gradient echo EPI BOLD signal was
acquired from whole-brain at 3T (GE Medical Systems Horizon
LX MRI scanner) from 17 oblique slices (+30° from axial; 5 mm
thick, 1 mm gap, in plane resolution=1.875 mm, TR/TE=2000/
40 ms). This slice prescription allowed near whole-brain coverage
except for the ventral-most surface of the posterior occipital lobes.
Anatomical images consisting of low-resolution T2 and high-
resolution SPGR (0.5 mm×0.5 mm×1.5 mm) images were also
Data analysis was performed using SPM2 (FIL, UCL London,
UK), and Matlab 6.1 (Mathworks, Natick, MA). The low-
resolution T2 volume was first coregistered to the high resolution
SPGR. GE-EPI images were then co-registered to the T2 volume
and corrected for motion and slice timing differences. The SPGR
image was normalized to the Montreal Neurological Institute
template, and the parameters determined for this normalization
were applied to functional images. Voxels from the SPGR and
functional images were resampled to 2 mm3isovoxel resolution.
967 J.C. Thompson et al. / NeuroImage 37 (2007) 966–973
Finally, the functional images were smoothed with a Gaussian
kernel with a full width half maximum (FWHM) of 8 mm.
Regressors for each of the experimental conditions were created
by modeling the BOLD response to each stimulus condition as a
box car with an ‘ON’ cycle equal to the length of each block,
convolved with a hemodynamic response function. Included in the
model were six motion covariates (three translation and three
rotation parameters) determined from the motion correction, and a
constant term to account for drift. A high-pass temporal filter of
1/128 s was also applied to the data to remove low frequency drifts
in the MR signal.
Parameter estimates were created for each subject, and contrasts
comparing all experimental conditions to fixation, in addition to
contrasts comparing face or hand movement to the radial grating
were created. The significance of each of these contrasts was
examined by using one-sample t-tests at the group level in a
random effects analysis. As the focus of this study was on posterior
temporal and parietal cortex, we created a volume of interest (VOI)
that contained the temporal, parietal and lateral occipital cortex
using the WFU_Pickatlas (www.rad.wfubmc.edu/fmri) (Fig. 2).
Results of the contrast for all experimental conditions relative to
fixation were thresholded at pb0.05 (corrected for multiple
comparisons at the cluster level). Within the voxels that survived
this threshold, contrasts between face or hand movement relative to
radial gratings were thresholded at pb0.005 (uncorrected) and
clusters of N30 voxels. Overlap in activation to face and hand
movement relative to radial gratings was determined by finding the
intersection of each of these contrasts thresholded as above
(Nichols et al., 2005). For the comparison between face and hand
movement, we extracted the beta weights from the voxel of peak
significance from the comparison of face or hand movement
relative to radial gratings and compared them using a t-test.
We also performed a whole-brain, voxel-wise analysis by
identifying regions that responded to all experimental conditions
relative to baseline, thresholded with a cluster level threshold of
pb0.01 (corrected for multiple comparisons). Voxel-wise contrasts
of face or hand movement relative to radial gratings were then
thresholded at pb0.005 (uncorrected) with clusters N30 voxels.
Overlap in activation was determined by finding the intersection of
activation as outlined above.
To examine the possible overlap in response to face and hand
movement relative to radial gratings in more detail, we also
conducted a single-subject analysis. Procedures used to transform
individual brains into stereotaxic coordinates such as the MNI or
Talairach coordinate systems align on the basis of anatomy rather
than function (Brett et al., 2002; Beauchamp et al., 2004). In
particular, volume-based warping procedures such as those used by
SPM2 do not match individual brains to the template on the basis of
Fig. 1. Activation task stimuli and sequence. A grid showing still images of Face (top), Hand (middle) and Radial motion pattern (bottom) stimuli shown before
andafterthe movement(leftandright),andduringmotionendpoint(center).Thereweretwopossiblemotiondurations(designatedas ‘short’or ‘long’).Atypical
ongoing experimental sequence is shown at the bottom of the figure.
968J.C. Thompson et al. / NeuroImage 37 (2007) 966–973
sulcal anatomy (Brett et al., 2002). If the STS varies considerably in
anatomy across individuals, the location of coordinates correspond-
ing to the STS in the template brain may not necessarily correspond
to the STS in individuals. It is also possible that normalization
procedures may lead to a distortion in the underlying functional
distribution of responses to different stimuli. To examine single-
subject responses, we conducted a whole-brain analysis on each
to 2 mm3and smoothed with an 8 mm FWHM Gaussian kernel. We
then identified regions that responded to all experimental stimuli
relative to baseline with a cluster threshold of pb0.05 (corrected for
multiple comparisons). Voxel-wise contrasts of face or hand
Fig. 2. Group activation to face and hand movement relative to radial grating motion. (A) Group activation data within the temporal and parietal VOI are overlaid
on the right hemisphere of a representative subject showing the response to viewing face vs radial (red), hand vs radial (blue), overlap of face vs radial and hand
vs motion (purple), and all motion including radial vs fixation (green). (B) Histograms of mean beta weights and standard errors for the three stimulus conditions
vs fixation are shown for the posterior inferior temporal sulcus (pITS), middle superior temporal sulcus (midSTS), posterior STS (pSTS) and the inferior parietal
lobule (IPL). (C) Group activation data from the whole-brain analysis are overlaid on the right hemisphere of a representative subject showing the response to
viewing face vs radial (red), hand vs radial (blue), overlap of face vs radial and hand vs motion (purple), and all motion including radial vs fixation (green). (D)
Histograms of mean beta weights and standard errors for the three stimulus conditions vs fixation are shown for right medial frontal gyrus (MFG), left inferior
frontal gyrus (IFG), left ventral premotor cortex (vPM), and left putamen/basal ganglia (Put). L=left.
969J.C. Thompson et al. / NeuroImage 37 (2007) 966–973
movement relative to radial gratings were then thresholded at
pb0.005. Overlap in activation was determined by finding the
intersection in activation as outlined for the group analysis. In order
to reference the location of the group activation to the single-subject
anatomy and activation patterns, we obtained the deformations used
these deformations, using the Deformations Toolbox in SPM2.
Then, for each subject, we applied the inverted deformations to the
MNI coordinates of the peak activations in temporal and parietal
cortex of the group analysis and compared the location of group
activation to single-subject anatomy.
No significant differences in mean accuracy for detecting the
target sequences were observed between the three movement
conditions Mean [±S.D.] % Accuracy: Face=70.8±19.9; Hand=
70.6±18.0; Radial Gratings=71.7±39.0). Similarly, there were no
significant differencesinmean reaction time (RT)to target sequence
detection between the three conditions (Mean [±S.D.] RT in
ms: Face=755.7±134.4; Hand=798.7±232.7; Radial Gratings=
Significant activation to the experimental conditions relative to
fixation was observed throughout the temporal and parietal VOI
(Figs. 2A and B, green). Within the areas that responded to the
experimental conditions relative to fixation, significantly greater
activation to the mouth movement than to radial gratings was
observed in the right midSTS (pb0.001) and right pSTS
(pb0.001) (Fig. 2A, red; Table 1). Significantly greater activation
to the hand movement than to the radial gratings was observed in
the right posterior inferior temporal sulcus (pITS) (pb0.0001),
right pSTS (pb0.001) and right IPL (pb0.001) (Fig. 2A, blue;
Table 1). Importantly, as the main focus of our experiment we
observed overlapping activation across face and hand movements,
when compared to radial gratings, in the right pSTS (x=56, y=
−48, z=12; Fig. 2A, purple). There was no significant difference
between face movements relative to radial gratings when compared
to hand movement relative to radial gratings (pN0.2) in this region.
Comparing responsivity in the midSTS revealed significantly
greater activation to face movements compared to radial gratings
relative to activation to hand movements compared to radial
gratings (pb0.05) (Fig. 2B). In contrast, there was significantly
greater activation to hand movements compared to radial gratings
than to face movements compared to radial gratings in the right
pITS (pb0.005) and right IPL (pb0.05).
Whole-brain analysis revealed activation all three experimental
conditions relative to fixation in the left ventral premotor cortex
(vPM), left inferior frontal gyrus (IFG), left putamen, and right
medial frontal gyrus (MFG) (Fig. 2C). Significantly greater
activation to face motion relative to radial gratings, and hand
motion relative to radial gratings, was observed in the left IFG, left
putamen, and right MFG (Table 1). All three areas active to both
face motion relative to radial gratings, and hand motion relative to
radial gratings (Fig. 2D IFG: x=−49, y=8, z=2; MFG: 2, 29, 52;
Put: −18, −2, 14;). In the left vPM there was significant activation
to all three conditions compared to fixation, with no significant
difference between conditions (Fig. 2D −46, −8, 46).
The single-subject analysis revealed a distribution of responses
to face and hand movement relative to radial gratings that was
consistent with the group activation. In particular, the location of
coordinates corresponding to the peak activations in the pITS,
midSTS, pSTS, and IPL in the group corresponded to these
anatomical locations in single-subjects. Overall, the finding of
overlapping activation to face movement relative to gratings and
hand movement relative to gratings in the pSTS was observed in 7
out of 11 subjects using a threshold of pb0.005. Lowering the
threshold to pb0.05 revealed that 9 out of 11 subjects showed
overlapping activation in pSTS. The overlap of activation to face
movement and hand movement relative to radial gratings from the
11 subjects is presented in Fig. 3. These results confirm that the
results of the group analysis were not due to distortion of
underlying response patterns by spatial normalization and instead
reflect common processing of face and hand movement in this
region. There was more variation across subjects in the response
patterns of the pITS, midSTS and IPL in which there was distinct
activation in the group analyses. For example, we observed
overlapping activation to both face and hand movement relative to
radial gratings in midSTS in several subjects (e.g. subjects AP and
TA in Fig. 3).
The present study examined fMRI responses as subjects
monitored and detected dynamic target sequences of face, hand,
and radial grating motion. Cortex in and around the STS has been
shown to consistently respond to the movements of other
biological forms (Allison et al., 2000; Puce and Perrett, 2003;
MNI Talaraich coordinates of activation foci to face and hand movements
versus radial grating motion
Brain regionCoordinates Volume
Superior temporal sulcus—mid
Superior temporal sulcus—posterior
Inferior temporal sulcus
Superior temporal sulcus—posterior
Inferior parietal lobule
Inferior frontal—dorsal pars opercularis
Inferior frontal—ventral pars opercularis
Medial frontal gyrus
Medial frontal gyrus
Inferior frontal—dorsal pars opercularis
Medial frontal gyrus
970J.C. Thompson et al. / NeuroImage 37 (2007) 966–973
Bonda et al., 1996; Puce et al., 1998; Beauchamp et al., 2002).
Consistent with these findings, we found greater responses to face
and hand movements than to radial grating motion in the right mid-
STS and pSTS, as well as the right pITS and IPL. Importantly, the
radial pattern motion also conveyed the same dynamic, task-
relevant information as that presented by the face and hand. This
suggests that these responses are not simply due to greater task-
relevance of the biologically relevant stimuli. Of the regions in
temporal and parietal cortex that responded significantly to face or
hand motion relative to the radial grating, a significantly greater
response to face motion relative to hand motion was observed in
the right mid-STS, whereas a significantly greater response to hand
motion relative to face motion was observed in the right pITS and
right IPL. However, analysis of single-subject data revealed that
there was some variability in the extent to which these regions
showed distinct activation. Importantly, a region of the right pSTS
showed overlapping responses to both face and hand motion,
relative to radial grating motion, and there was no significant
difference between the response to face or hand motion in this
region. The overlapping activation in pSTS was consistent across
individual subjects, indicating that it was not simply a result of the
group averaging process and reflects common processing of face
and hand movement in this region.
It has been suggested that the STS provides a visual description
of motion details to other regions involved in the processing of
other people's actions, such as the IPL, PMv, and IFG (Iacoboni et
al., 1999). However, the nature of this description has been unclear.
A study by Pelphrey and colleagues (2005) indicated a somatotopic
Fig. 3. Single-subject activation to face and hand movement relative to radial grating motion. Activation data from the pSTS are overlaid on the right hemisphere
of each subjects' T1 anatomical images showing the response to viewing face vs radial (light red pb0.005; dark red pb0.05), hand vs radial (light blue pb0.005;
dark blue pb0.05), and overlap of face vs radial and hand vs motion (light purple pb0.005; dark purple pb0.05), Yellow squares show the location of the peak
activation in the pSTS from the group analysis following transformation to each subject's native space.
971J.C. Thompson et al. / NeuroImage 37 (2007) 966–973
arrangement of responses to viewing hand, eye and mouth
movements was present in the STS and surrounding cortex. A
number of other findings have suggested that there may even be a
different pattern of responses to face motion compared to hand
motion in the STS and parietal cortex (Thompson et al., 2004;
Wheaton et al., 2004). These data suggest that the STS may form
an intermediate-level, body-part specific representation of biologi-
cal motion or that this region may prefer face motion to hand
motion. The findings of the present study suggest instead that a
region of the pSTS may form a high-level representation of
biological movement that is not dependent on the particular body
part generating the motion. An alternative hypothesis is that the
distribution of responses in STS and surrounding cortex might in
part be shaped by particular task requirements. In circumstances
when one is focused on higher-level stimulus attributes, such as the
dynamics of the motion in the present study, the pSTS might form
a high-level representation that is not bound to a particular body
part. In circumstances in which the motion of one particular body
part might be the primary source of information, effector-specific
representations may be formed and passed on to other parts of the
network involved in action recognition. Such a flexible mechanism
might explain the distribution of responses to different body part
motion reported by Pelphrey and colleagues (2005), as during
passive viewing subjects might have focused to a greater extent on
the specific effector that was moving. Our findings do suggest,
however, that it is not just the dynamics of the action that are
processed by the pSTS, as the overlapping responses to face and
hand motion were significantly greater than the response to radial
grating motion even though the task-relevant information did not
differ between face, hand and radial gratings.
It should be noted that, similar to the findings of Pelphrey and
colleagues (2005), we observed greater responses to face motion
than hand motion in the right mid-STS, and greater responses to
lateral temporal cortex (pITS) and the right IPL. There was
variability across subjects in the extent to which each of these
regions showed distinct responses to either face movement or hand
movement relative to radial gratings, and lowering the threshold for
significance indicated that several subjects showed overlapping
responses in these regions. In general these results suggest that in
addition to the common processing of different body part motion in
the pSTS, there is some evidence of distinct responses to different
body parts in the neighboring lateral temporal and inferior parietal
regions (Pelphrey et al., 2005; Thompson et al., 2004; Wheaton et
al., 2004). More anterior regions of the STS, particularly in the right
hemisphere, respond to hearing speech and nonspeech human
vocalizations (Belin et al., 2000). The mid-STS location of the
strongest response to face motion may reflect the recruitment of
with vocalizations. The pITS response in the present study, with
greater activation to hand versus face motion, is close to a region
responding to static images of headless bodies, hands and hand
actions that has been labeled the extrastriate body area, or EBA
during the performance of hand actions, even when the hand is not
visible (Astafiev et al., 2004). The IPL responds to viewing hand
actions (Buccino et al., 2004), as well as the execution of hand
actions (Buccino et al., 2004; Ehrsson et al., 2001). Both the EBA
and IPL may be involved in visually-guided hand actions, with the
right IPL in particular playing an important role in guiding hand
actions in space (Mattingley et al., 1998; Husain et al., 2000).
In addition to activation in the temporal and parietal cortex, we
also observed greater activation to face and hand movements,
relative to radial grating motion, in the right MFG, left IFG and left
putamen. Activation in the MFG and IFG to biological movement
has been reported previously (Aziz-Zadeh et al., 2006; Grafton et
al., 1996; Iacoboni et al., 2005, 1999; Johnson-Frey et al., 2003;
Pelphrey et al., 2005; Rizzolatti et al., 2001; Schubotz and von
Cramon, 2004). Pelphrey and colleagues (2005) found that the
MFG responded to eye and mouth movements, but not hand
movements (Pelphrey et al., 2005). Our data show that the MFG
responded to hand movements and face movements, with a region
of overlap between the two. One could argue that activation in
these regions might also be expected as a function of subjects
using working memory to keep track of presented stimuli in order
to correctly identify the required target sequences. All three
stimulus conditions, including the inwardly moving radial pattern,
required subjects to keep track of the current sequences of previous
three items in working memory. Interestingly, greater activation
occurred to face or hand motion sequences than to the radial
pattern, yet the working memory load and subjects' performance
did not differ across the three stimulus types, indicating that the
ventral prefrontal cortex activation, often associated with manip-
ulation of information in working memory (Wager and Smith,
2003), may not be driven by these processes. In contrast to
previous studies, we did not observe distinct responses to face or
hand motion in another region of frontal cortex, the so-called
ventral premotor (PMv) region (Buccino et al., 2001; Wheaton et
al., 2004). In the present study, all three movement conditions
activated a large section of the left PMv. A recent study by
Schubotz and von Cramon (2004) also reported overlapping
activation to detecting sequences of biological and non-biological
stimuli in the left PMv. Together with the present results, these
findings raise the possibility that this region may encode sequential
information that predicts action outcomes regardless of whether the
stimuli are biological or not. This would be an interesting avenue
of exploration in future studies.
Understanding how different brain regions respond to the
motion of different body parts can help provide insight into the role
of these regions in action recognition. The common response to
face and hand motion in the pSTS suggests that this region may
form a high-level representation of biological movement that is
independent of body part. This response is, however, greater than
the response to a non-biological stimulus matched for motion
dynamics and task-relevance. This high-level visual description of
biological movement may be passed on to other brain regions
involved in making sense of the actions of others.
The authors thank Ms. Mary Pettit for editorial assistance. The
collection and initial analysis of data from this study by James
Thompson was conducted as part of a postdoctoral fellowship in
the Center for Advanced Imaging and Department of Radiology at
West Virginia University. This study was funded by a Project Grant
awarded to Aina Puce by the National Health and Medical
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