REVIEWS
The neural basis of visual
body perception
Marius V. Peelen and Paul E. Downing
The human body, like the human face, is a rich source of socially relevant information about
other individuals. Evidence from studies of both humans and non-human primates points
to focal regions of the higher-level visual cortex that are specialized for the visual perception
of the body. These body-selective regions, which can be dissociated from regions involved
in face perception, have been implicated in the perception of the self and the ‘body schema’,
the perception of others’ emotions and the understanding of actions.
Transcranial magnetic
stimulation
(TMS). A technique that
delivers brief, strong electric
pulses through a coil placed on
the scalp. These create a local
magnetic field, which in turn
induces a current in the surface
of the cortex that temporarily
disrupts local neural activity.
Centre for Cognitive
Neuroscience, School of
Psychology, Brigantia
Building, University of
Wales, Bangor, Gwynedd,
LL57 2AS, UK.
Correspondence to P.E.D.
e-mail:
p.downing@bangor.ac.uk
doi:10.1038/nrn2195
Humans, like other primates, are highly social — indeed,
one of our harshest punishments is solitary confinement.
Our lives are intertwined with those of other people,
and so we must be able to efficiently determine their
identities, actions, emotions and intentions. Much of
this information is available from the appearance of the
face, and in the field of cognitive neuroscience there
has been intense interest in understanding the neural
mechanisms that support face perception1–3. By contrast,
the perception of the rest of the body has until recently
received less attention, even though the body shares
many characteristics with the face4: it conveys socially
relevant information; becomes highly familiar with
repeated exposure over the lifespan; is similar in shape
across individuals; and is visually salient, as witnessed by
its ability to capture attention5–7.
The past few years have brought a remarkable
increase in research on the neural basis of visual perception of the human body, which we review here.
Our focus is on visual perception of the body and on
the occipitotemporal brain systems that are thought to
underlie it. We do not review in detail work in related
areas that has recently been covered elsewhere, including
face perception1–3, fronto-parietal action perception or
‘mirror neuron’ networks8,9, biological motion perception10,11, disorders of the body schema12, and emotional
body language13, although our discussion intersects with
all of these topics.
This Review is divided into two main sections. First,
we survey evidence for body-selective neural mechanisms in the visual cortex from single-cell recordings in
primates, and from event-related potential (ERP), functional MRI (fMRI), transcranial magnetic stimulation (TMS)
and neuropsychological studies in humans. Second, we
review evidence on how body-selective brain regions
relate to perception of the self and the ‘body schema’,
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understanding others’ emotions, and action perception
and the ‘mirror’ system. Throughout both sections we
consider how the brain systems that are involved in face
and body perception might be related. Finally, we discuss
some of the many open questions for future research.
Evidence for body-selective neural mechanisms
Non-human primates. Studies on macaque monkeys
have revealed that some neurons in the inferior temporal
cortex (IT) respond selectively to the shapes of human
and monkey bodies and body parts14–18 (FIG. 1). For
example, hand-selective cells in the IT respond strongly
to images of human and monkey hands of different orientation and size, presented at various locations in the
visual field, but not to other complex shapes or to faces15.
Other cells in the IT respond selectively to faces but not
to hands. Various other object categories were used in
this study, but no IT neurons responded selectively to
them, indicating that category selectivity in IT neurons
is restricted to a small number of categories15. A recent
study measured responses in a large number (>600)
of IT neurons to many different images (>1,000) from
natural and artificial object categories 14. An analysis
on the patterns of activity across cells showed that
animate and inanimate images elicited mostly distinct
patterns. In the animate category, faces and bodies elicited distinct patterns, with further distinctions between
different types of face (animal, monkey or human),
and different types of body (human, four-limbed animal
or bird). Analyses of single-cell responses revealed many
cells that responded selectively to human bodies, hands
or faces. Only a few cells selectively responded to a combination of these categories, providing further support
for distinct representations of these categories. Other
studies have reported cells in the anterior part of the
monkey superior temporal sulcus (STS) that respond
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a
Ce
Two recent studies that used fMRI in monkeys provide evidence for two large clusters of body-selective cells
in the STS, with those in the right hemisphere being most
strongly activated24,25 (FIG. 1). Interestingly, in both of these
studies the body-selective area was adjacent to, and partly
overlapped, a face-selective area. Subsequent recordings
from individual neurons in the face-selective area (as
identified with fMRI) revealed that almost all of the cells
were strongly selective for faces26. This result provides a
bridge between single-unit recordings in monkeys and
fMRI findings in humans by showing that dense clusters of highly selective individual neurons can underlie
selectivity measured at a macroscopic level with fMRI.
Ip
L
Ts
Tmp
Tma
Oi
La
b
1
1
1
2
3
3
4
4
5
6
c
5°
d
Faces
Bodies
STS
Figure 1 | Key findings from non-human primates. a | Orange indicates the region
investigated in early single-unit recording studies of macaque inferotemporal cortex15–17.
b | Increasing responses to progressively more hand-like stimuli in a neuron from
macaque inferotemporal cortex. Numbers reflect approximate relative magnitude of
responses to different stimuli. c | Spike histograms and schematic stimuli from an
investigation of a hand-selective neuron15. This neuron seems to be selectively tuned to
the general form of the hand. The response to faces is low, ruling out a general response
to all biological stimuli. The response to hand-like geometric stimuli is also low,
demonstrating the specificity of the response to realistic hand shapes. d | Closely
neighbouring regions of the superior temporal sulcus (STS) in the macaque are
selectively activated in functional MRI by images of faces (left), as compared with bodies,
fruits, artefactual objects and hands. This is also true for images of bodies (right), as
compared with faces, fruits, artefactual objects and hands. Ce, central sulcus;
Ip, intraparietal sulcus; L, lunate sulcus; La, lateral sulcus; Oi, inferior occipital sulcus;
Tma, anterior middle temporal sulcus; Tmp, posterior middle temporal sulcus; Ts, superior
temporal sulcus. Panels a and c modified, with permission, from REF. 15 © (1984)
Society for Neuroscience. Panel b modified, with permission, from REF. 16 © (1972)
The American Psychological Society. Panel d modified, with permission, from Nature
Neurosci. REF. 25 © (2003) Macmillan Publishers Ltd.
to particular body actions and body postures 19–23,
and to specific body postures as a function of the preceding observed action, such that the cell responds to
a particular body posture only when it is preceded by a
particular action20,21.
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Intracranial recordings in humans. Intracranial surfaceelectrode recordings of electrical activity in human brains,
performed on patients being evaluated for surgery to treat
epilepsy, benefit from both high spatial and high temporal
resolution. Studies using this technique have found bodyselective responses in various posterior cortical regions,
including the extrastriate visual cortex (FIG. 2). In one
study, several electrodes showed hand-selective responses
that peaked at about 230 ms after stimulus onset27. These
hand-selective electrodes were located in various regions,
including the right ventral visual cortex, the left STS and
the left inferior parietal cortex. Another study recorded
responses in the right lateral occipitotemporal cortex and
found that one electrode, at the approximate location of
the extrastriate body area28, recorded selective responses
to images of whole bodies (without heads), with a bodyselective response starting at 190 ms and peaking at 260 ms
after stimulus onset29. Interestingly, in these studies, bodyselective electrodes did not record significant responses to
faces, and face-selective electrodes did not record significant responses to bodies, providing further evidence that
the body and its parts activate different neurons to those
that are activated by the face.
Evoked potentials in humans. Recordings of electrical
and magnetic stimulus-evoked potentials on the scalp show
enhanced responses to faces, with peaks at approximately 100 ms (REFS 30–33), 170 ms (REFS 34–37), and
250 ms (REFS 38,39) after stimulus onset. Most attention
has been focused on the negative potential that peaks at
posterior sites at about 170 ms (the ‘N170’), although
the face selectivity of this waveform has been questioned33,40. Recent findings indicate that human bodies,
or body parts such as hands, produce a similar evoked
response41,42. For both bodies and faces, the response
that peaks at around 170 ms is enhanced and delayed by
image inversion43 — an index of configural processing 44–46
— and reduced by image distortion47. Another study
found a strong response to bodies that peaked at 190 ms,
about 20 ms later than the response to faces48 (FIG. 3).
The body selectivity of this ‘N190’ peak also generalized
to stick figures and silhouettes, but not to scrambled
versions of these figures48. In these studies, the spatial
distribution of responses to bodies differed from that
of responses to faces, and source localization identified distinct lateral occipitotemporal sources for the
face-selective N170 and body-selective N190 peaks.
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a
b
P310
P165
+
100ms
N230
100µm
+
N230
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50µm
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Amplitude (µV)
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0
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40
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100
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0
250
500
750
1,000
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Time (ms)
Figure 2 | Key findings from intracranial recordings in humans with epilepsy. a | A summary of hand-selective
negative potentials (indicated by red lines) peaking at approximately 230 ms post-stimulus. Blue lines indicate the
response to images of faces, and green lines the response to images of objects. Red spots indicate body-selective
recording sites across different patients. Red crosses indicate the average position of the inserted electrodes.
b | Pre-surgical intracranial recordings from a patient with epilepsy reveal an electrode at the approximate location of
the extrastriate body area28 (electrode 15; indicated by the yellow box) that is body-selective (red line) from 190 ms poststimulus, relative to faces (blue line), tools (green line) and animals (black line)29. Panel a modified, with permission, from
REF. 27 © (1999) Oxford University Press. Panel b modified, with permission, from REF. 29 © (2007) Elsevier Science.
Stimulus-evoked potential
An electrical or magnetic
potential, resulting from
coordinated neural activity,
that is time-locked to the
onset of a stimulus.
Configural processing
The recognition of an object
by the specific spatial
relationships among (or
configuration of) its parts.
Sometimes referred to as
‘holistic’ processing.
Source localization
A technique used in electroencephalogram (EEG) and
magnetoencephalogram (MEG)
research to estimate the
location of the brain areas that
give rise to the electrical or
magnetic responses
that are measured on the
scalp.
Structure-from-motion
Even a few dots can create the
vivid perception of an object or
structure when they move in a
way that is typical of that
object.
Developmental studies have used event-related potentials (ERPs) to measure the neural response to bodies and
to biological motion in infants. Babies as young as 4–6
months old look longer at intact, as opposed to scrambled,
human biological motion patterns presented in pointlight displays (PLDs), indicating a capacity to perceive the
distinction between these stimuli. These displays, which
consist of only a few dots, give a vivid perception of the
human form through the process of structure-from-motion49.
This looking preference is restricted to patterns that
represent upright bodies50,51. In babies of 8 months of
age, differences are apparent between ERPs recorded
in response to intact versus scrambled52, and to upright
versus inverted53, PLDs. These differences emerge in
roughly the first 200–400 ms after the stimulus onset and
have a right-hemisphere bias. Like adults, infants of just 3
months of age show similar ERPs in response to images of
static faces and static bodies. These responses differ from
those elicited by distorted face and body images, and the
differences in the waveforms emerge at about 450 ms
(REF. 47); these effects are similar for both categories.
This sensitivity to configuration in very young infants
is surprising, given that structural disruption of static
body images does not seem to affect looking preferences
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until 18 months of age54, although this discrepancy may
be a result of stimulus differences between studies55.
Clearly, further developmental experiments are needed
that combine behavioural and neuroimaging measures,
compare moving and static body, face and object images
at a range of ages, and consider the effects of different
types of image disruption.
Functional MRI in humans. Perhaps the clearest evidence for body-selective brain regions in humans comes
from fMRI studies that compare responses to images
of (headless) bodies and body parts with responses
to control images (FIG. 4). This approach has revealed
that a focal region of the lateral occipitotemporal cortex responds strongly and selectively to static images
of human bodies and body parts, but weakly to faces,
objects and object parts. The responses of this region
generalize to non-photographic depictions of bodies,
such as line drawings, stick figures and silhouettes,
indicating that this brain area has a body representation that is abstract across specific visual features28. The
response of this region to non-human animals is significantly lower than to humans, but higher than to objects,
and it is also higher to mammals than to birds and fish,
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REVIEWS
spatially-smoothed or group-averaged data — as a reflection of the level of activity in the EBA (or hMT or LO),
when this interpretation is simply based on the average
peak activation coordinates.
Recent fMRI studies have provided evidence for a
second body-selective area that is anatomically distinct
from the EBA (FIG. 4). This region, located in the fusiform
gyrus and so known as the fusiform body area (FBA),
responds selectively to whole bodies and body parts, as
well as to schematic depictions of the body 60,66,67. The
FBA is adjacent to and partly overlaps the face-selective
fusiform face area (FFA)68. Interestingly, neighbouring
face- and body-selective patches have also been found
in the monkey STS24,25.
The close proximity of the FBA and FFA also raises
the question of whether the apparent body selectivity
found in the fusiform gyrus could instead be due to
an indirect activation of face-selective neurons, for
suggesting that this area is partly activated by objects
with a body plan similar to that of humans56. On the
basis of these and other findings, this region has been
labelled the extrastriate body area (EBA)28.
The EBA is found bilaterally in the posterior inferior
temporal sulcus/middle temporal gyrus57,58. Depending
on the statistical threshold used in imaging studies, bodyselective voxels overlap with the human motion-selective
area MT (hMT) and with the dorsal focus of the objectform selective area LO28,59. This presents methodological
difficulties for the characterization of these regions using
group-average fMRI data or even single-subject functional
region-of-interest designs. Two recent studies59,60 overcame
these difficulties by using multi-voxel pattern analyses
(MVPA) of fMRI data61–65 to disentangle, at a finegrained level, these regions’ patterns of selectivity (BOX 1;
FIG. 5a). These considerations indicate that caution is
required when interpreting activations — particularly in
Voxel
In MRI research, a voxel refers
to the smallest measured
volume unit, analogous to a
three-dimensional pixel. In
fMRI studies, these are
typically of the order of
30 mm3, although much smaller
voxel volumes have been
achieved in more recent work.
Human motion-selective
area MT
An area in the human
extrastriate visual cortex that
responds strongly to visual
displays containing moving
items. It can be functionally
localized with fMRI by
contrasting activation relating
to moving stimuli with that
relating to static stimuli.
b
c
10
8
a
10
Bodies
Faces
Objects and scenes
6
6
Amplitude (µV)
Amplitude (µV)
Silhouettes
Scambled silhouettes
Stick figures
Scrambled stick
figures
8
4
2
0
4
2
0
–2
P8
–2
–4
–4
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0
–50
50 100 150 200 250 300 350 400 450 500
0
50 100 150 200 250 300 350 400 450 500
Time (ms)
Time (ms)
d
–7
–7
Upright
–6
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0
2
4
6
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–2
0
2
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200
Time (ms)
300
400
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–4
Amplitude (µV)
–2
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–6
Inverted
–4
Amplitude (µV)
Amplitude (µV)
–4
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Upright
–2
0
2
4
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200
Time (ms)
300
400
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0
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400
Time (ms)
Figure 3 | Event-related potentials reveal similar, but distinct, responses to faces and bodies. a | A body-selective
event-related potential (ERP) negative (N1) peak is centred approximately at electrode P8, the site from which the data in
the remaining panels were acquired. b | The body-selective response (red line) peaks at 190 ms post-stimulus, about 20 ms
later than the response that is elicited by pictures of faces48 (blue line), and roughly the latency at which body selectivity
has been observed in intracranial recordings of occipitotemporal sites in human patients with epilepsy (FIG. 2). c | This
body-selective ‘N190’ generalizes to abstract depictions of the body, distinguishing between intact and scrambled stick
figures, and between intact and scrambled human silhouettes48. d | For both bodies (left graph) and faces (middle graph),
but not objects (right graph), stimulus inversion delays and amplifies the N1 response43. Panels b and c modified, with
permission, from REF. 48 © (2006) Elsevier Science. Panel d modified, with permission, from REF. 43 © (2004) Lipincott
Williams & Wilkins.
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a
b
X=48
X=51
X=50
X=52
X=54
X=52
Body selective
Face selective
EBA
OFA
c
FFA/FBA
120
100
Response
80
60
40
20
0
Stimulus
Figure 4 | Body- and face-selective regions of the human occipitotemporal
cortex, as revealed by functional MRI. a | The extrastriate body area (EBA; shown
here in the right hemisphere of six subjects) is found in the posterior inferior temporal
sulcus. X values indicate Talairach coordinates for each slice plane. b | Body- and faceselective regions of the human visual cortex, in a ventral view of the right hemisphere of
one individual, rendered on an inflated anatomical scan from the same individual.
Orange indicates body-selective regions (bodies versus tools); green indicates faceselective regions (faces versus tools). Bodies and faces activate similar, but not
identical66,67, regions of the fusiform gyrus (the fusiform body area (FBA), and fusiform
face area (FFA), respectively). Posterior to this region are nearby but distinct bodyselective (EBA) and face-selective (occipital face area (OFA)) regions.
c | Responses of the functionally defined EBA to various stimuli, indicating the bodyselective response of this region. Values are taken from several experiments, each of
which included photographs of varied body parts as one condition. Magnitudes reflect
the mean response to each stimulus type, averaged across several subjects, with all
responses scaled so that the response to body parts is equal to 100. Stimulus conditions,
from left to right: body silhouettes; whole bodies; assorted body parts; hands;
stick figures of bodies; mammals; scrambled silhouettes; scrambled stick figures; faces;
scenes; object parts; and whole objects. Panel a modified, with permission, from
REF. 119 © (2007) Oxford University Press.
Object-form selective
area LO
A region of the human
extrastriate visual cortex that
responds to object form. It can
be functionally localized by
contrasting fMRI activation
relating to intact objects with
activation relating to scrambled
objects.
instance through mental imagery or the contextual
association of bodies and faces69. However, several
studies argue against this explanation by showing that
the body- and face-selective activations can be anatomically and functionally dissociated. For example,
a high-resolution fMRI study found small neighbouring patches of the fusiform gyrus that were either
selective for bodies but not faces, or for faces but not
bodies67. Furthermore, the application of MVPA has
shown that the local patterns of selectivity for faces
and bodies across fusiform voxels are unrelated60. In
other words, there was no relationship between face
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and body selectivity across voxels — voxels that were
relatively strongly face selective were not necessarily
also strongly body selective, as would be expected if
both categories activated the same underlying neuronal
populations (BOX 1; FIG. 5).
The evidence that the body representation in the
EBA and FBA is independent of low-level image features is further supported by the finding that these
regions are also activated in response to PLDs of human
motion60,70–75. The functionally localized EBA and FBA
are activated significantly more by such PLDs than by
‘scrambled’ control displays60. In standard region-ofinterest analyses, the area hMT (which overlaps the EBA)
and the FFA (which overlaps the FBA) showed similar
responses. However, when MVPA was applied to local
patterns of activity in these areas, only body selectivity,
and not motion or face selectivity, was related to the
voxelwise pattern of selectivity to PLDs60. This indicates
that PLDs specifically activated body-selective neuronal
populations (BOX 1) in both brain areas. Note that the
activation of the EBA and the FBA by PLDs is unlikely
to reflect processing of biological motion patterns per se,
as these regions are strongly activated by static body
stimuli, even when they do not imply motion28. Instead,
the processing of biologically plausible motion has been
linked to other areas that are typically activated by PLDs,
such as the posterior STS (pSTS)76–78.
Finally, a key question is whether the EBA and FBA
can be dissociated on the basis of their functional properties. A recent study of the response to body images
of varying completeness (fingers, hands, arms, torsos
and whole bodies) found more selectivity for body
parts in the EBA than in the FBA, and a relative bias
in selectivity for more complete images of the body in
the FBA79. This finding suggests a possible distinction
between these two areas, with the analysis of bodies
in the EBA being focused on individual parts, and the
function of the FBA being to create a more holistic body
representation.
TMS and lesion studies. Are body-selective brain areas
necessary for the perception of bodies and body parts?
TMS offers the opportunity to test this question in
healthy subjects. Because the EBA is located on the
lateral surface of the brain, close to the scalp, it is relatively easy to create a ‘virtual lesion’80,81 of this region
with TMS, whereas the FBA is out of reach of direct
TMS effects. The first TMS study to investigate whether
the EBA is essential for body processing showed that
disrupting activation in this area 150–250 ms after the
onset of the stimulus selectively impaired performance
on a delayed match-to-sample task involving images of
body parts, but not face or motorcycle parts82 (FIG. 6).
Thus, the EBA is causally involved in creating and/or
maintaining an accurate representation of the shape of
body parts, but not of object parts. Note that the latency
of the TMS pulses in this study, between 150 and
250 ms, generally coincides with the latency of a bodyselective peak in ERPs, as measured with electrodes
on the scalp (190 ms)48 and intracranial recordings
(230–260 ms)27,29.
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Box 1 | Multi-voxel pattern analysis to interpret overlapping activations
Although functional MRI (fMRI) is hailed for its high spatial resolution compared with
other imaging techniques, each measured unit (voxel) reflects the summed activation of
thousands of neurons. Therefore, where neuronal populations with different functional
characteristics are intertwined at a relatively fine scale, fMRI cannot separate these
populations. Recent studies have shown that analysis techniques that take into account
patterns of activation across voxels can partly overcome this problem59–61,65,97,145,156,157. The
application of multi-voxel pattern analysis (MVPA) is best illustrated by an example from
the primary visual cortex (V1), the neural architecture of which is well known from animal
work. Single-unit recordings in the monkey brain have shown that neurons in V1 are
sensitive to the orientation of bars, with some neurons responding optimally to
horizontal bars whereas others prefer bars at other orientations. These differently tuned
neurons are intertwined at a sub-voxel scale, and each fMRI voxel therefore contains a
mixture of these neurons. Importantly, each voxel will show a slight orientation bias due
to differential sampling of orientation-selective neurons. These biases are generally too
weak to pick up when each voxel is analysed independently. However, when analysing all
V1 voxels simultaneously by looking at the pattern of activation across voxels, these
weak (but reliable) biases become informative, because the patterns in response to, for
example, horizontal and vertical bars are significantly different. This information can be
used with linear classifiers to determine which orientation was viewed by a subject
during a block of trials, or even an individual trial65,145,158.
Other studies have used similar logic to dissociate overlapping neuronal populations
in the higher-level visual cortex, including the body-selective visual cortex. For
example, using MVPA, it has been possible to dissociate body-selective from motionselective populations in the lateral occipitotemporal cortex59,60 (FIG. 5a), and
body-selective from face-selective neuronal populations in the fusiform gyrus60
(FIG. 5b). More generally, these results show that, when a group of voxels is activated
by multiple conditions (for example, bodies and faces, or horizontal and vertical bars),
this cannot be assumed to reflect activation of the same underlying neuronal
population. MVPA might therefore be a useful tool for interpreting overlapping
activations anywhere in the brain62–64.
Functional region-of-interest
design
A design used in fMRI research
in which one or more brain
areas are defined on the basis
of their functional properties
(typically in each subject
individually), and their
response properties are further
investigated in subsequent
experiments.
Delayed match-to-sample
task
A task in which subjects have
to choose which of multiple
target stimuli matches a
previously presented sample
stimulus that is held in
memory.
Linear classifier
A statistical procedure in which
items are divided into two or
more groups on the basis of a
weighted linear combination of
their features.
Corollary discharge
A copy of the motor signal that
can be used to adjust for
changes in sensory input that
result from the motor action.
A related issue is whether permanent lesions to the
extrastriate cortex as a result of brain injury can cause
body-perception deficits. Focal lesions to the cortex that
encompass the EBA are rare (especially bilaterally), as
evidenced by the scarcity of patients reported with selective damage to the area hMT (which overlaps closely
with the EBA). A case study investigating a patient with
bilateral lesions involving the hMT reported significant
loss of motion perception, whereas other perceptual
functions (including face perception) remained relatively intact83,84. Disordered body perception was not
reported but might not have been tested. A more recent
study examined deficits of body perception in a large,
unselected group of stroke patients with lesions in a
single hemisphere85. The pattern of deficits in various
body-related tasks indicated a three-way dissociation
between representations of the body: postural (for
example, imagining and performing actions), semantic
(such as judging functions of body parts) and structural
(for instance, localizing pictured or touched body parts)
(see also REF. 54). The semantic and structural representations were associated with lesions of the left temporal
lobe, although the anatomical evidence from this study
was not precise enough to determine whether this
specifically involved the EBA or the FBA.
The recent evidence for a body-selective area in
the fusiform gyrus raises the further question of why
body-perception deficits are not reported in conjunction with face-perception deficits (prosopagnosia) that
arise from occipitotemporal lesions1,86. Given their close
NATURE REVIEWS | NEUROSCIENCE
proximity, lesions to the FFA would be expected to significantly affect the FBA as well. However, it is not clear
whether the FFA overlaps with the crucial lesion site for
acquired prosopagnosia — for example, although some
fMRI studies report relatively normal FFA responses in
patients with acquired prosopagnosia87–89, others report
abnormal90,91 face-related activity in this region. It is also
possible that people with prosopagnosia might have subtle defects in body perception that are masked by their
difficulties with faces and that are not routinely tested for
(although this has been ruled out in one comprehensive
test of an individual with congenital prosopagnosia92).
Finally, it is possible that deficits that result from permanent lesions in the EBA or the FBA are quickly compensated for by the remaining intact body-selective area
or by other brain areas that respond to the human form
(for example, the pSTS).
Body perception and other processes
In the following sections, we review the implications of
the above findings, particularly of the existence of two
body-selective regions in the extrastriate cortex, for
several related issues in cognitive neuroscience.
Self perception and the body schema. As we act in the
world, we maintain a continuous sense of our posture
and the positions of our limbs. This sense has been
referred to as the ‘body schema’93. Disruption of this
schema has been proposed to underlie a number
of neurological syndromes, mostly those caused by
damage to the parietal cortex12, although the diversity of these syndromes indicates that there may be a
multitude of body schemas rather than a single one.
Updating of the schema is thought to depend largely
on tactile or somatosensory information (as evidenced,
for example, by intact motor functioning in the
congenitally blind12).
A recent study has indicated that the EBA might
receive signals that update the representation of the body
after movement; this would indicate a novel coupling
between visual and motor representations. Specifically,
the EBA responds to the execution of visually guided
(but unseen) voluntary movements of a subject’s hands
and feet94, (see also REF. 95). This finding was interpreted
to indicate that the execution of a movement might
affect the actor’s body representation through movement-related proprioceptive inputs. Alternatively, or in
addition, the EBA could be activated through corollary
discharge signals from motor areas. These signals might
dynamically update the body representation in the
EBA, and adjust for sensory input resulting from
the movement (a function that is generally attributed
to the inferior parietal cortex). The integration of such
internal action signals with external visual input could
ultimately serve to distinguish between one’s own and
someone else’s body parts96. A subsequent study showed
that, although body movements activate the EBA,
this activation might not be directly related to visual
body perception97. This study replicated the finding
of increased activation in the EBA during execution of
motor actions, but reported relatively low spatial overlap
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a
b
Body selectivity
Object selectivity
Body selectivity
k
‘Biomotion’ selectivity
Motion selectivity
Face selectivity
‘Biomotion’ selectivity
Strong correlation
Weak correlation
Figure 5 | Disentangling the regions of the occipitotemporal cortex. a | Multi-voxel pattern analyses (MVPA) of
functional MRI (fMRI) data disentangle inferior temporal sulcus (ITS) responses to bodies, objects, simple visual motion and
biological motion (see also BOX 1). Bodies, objects, ‘point light’ animations of human movements and simple oscillatory
motion all activate similar regions of the ITS in human fMRI studies (see rendering at top of figure). These responses can be
difficult to disambiguate — for example, even when only the single most object-selective voxel from this region is tested
across individuals, significant motion, biological motion and body selectivity is found59. Pattern analyses consider the local
variations in selectivity across the voxels of the region as a whole. They reveal that body and biological motion selectivity
have significant positive correlation, indicating that they engage similar underlying neural populations (that is, the
extrastriate body area). By contrast, the variations in motion and object selectivity are uncorrelated with each other and
with body selectivity, indicating distinct underlying neural systems59,60. b | Similarly, MVPA of fMRI data disentangle
posterior fusiform gyrus responses to bodies, faces and biological motion (see also BOX 1). Voxelwise pattern analyses
reveal that body and biological motion selectivity show significant positive correlation, indicating that they engage similar
underlying neural populations (that is, the fusiform body area). By contrast, these patterns are uncorrelated with the
pattern that is elicited by faces, indicating a distinct underlying neural system (that is, the fusiform face area)60.
in individual subjects (14–19%) between areas of the
brain that were activated by movements and those that
were activated by body perception, especially when body
movements were contrasted with a stringent control condition97. Furthermore, even within this region of overlap,
MVPA showed no relationship between perceptual body
selectivity and movement-related modulation. That is,
variations in body selectivity were uncorrelated, across
voxels, with variations in movement-related modulation:
voxels that showed strong body selectivity did not necessarily show strong movement-related modulation, and
vice versa97 (BOX 1). This finding indicates that there may
be overlapping but functionally independent mechanisms, similar to the independence of selectivity for the
perception of simple visual motion and bodies in this
region59. Indeed, several studies have linked extrastriate activity in response to body movements to motion
perception in the nearby middle temporal area (MT)
or medial superior temporal area (MST) rather than in
the EBA98–100. Follow-up studies that localize areas that
are involved in body perception, motion perception and
action responses are needed.
Although schema updating is thought to depend
largely on somatosensory input, visual perception of
body parts also contributes significantly to our subjective
642 | AUGUST 2007 | VOLUME 8
sense of the position and ‘ownership’ of our own limbs.
This can be demonstrated by the visual capture of the
sense of limb position that is elicited by the ‘rubber hand
illusion’101. A rubber arm is placed in the subject’s view,
above and parallel to his or her own arm, which is hidden
below a screen. When a brush is stroked over both arms
simultaneously, the tactile sensation is often ‘captured’ by
the sight of the brush on the rubber arm, and the sense
of limb ownership is associated with the rubber arm.
Neuroimaging studies have linked cerebellar, parietal,
premotor and frontal opercular areas with this illusion,
and the premotor and frontal areas in particular with the
sense of ownership102–104. These studies did not reveal
illusion-related activity in posterior occipitotemporal
areas that might coincide with the EBA or the FBA,
although no study has specifically tested for this with
functionally defined regions of interest. Importantly,
any such study would potentially be confounded by differential visual attention or patterns of eye movements
to the limb when the illusion occurs.
Finally, vision is also relevant to self perception in
that some views of body parts are possible only from
an allocentric perspective. Two studies of EBA activity compared views of bodies105 or hands and feet106
that would be consistent with egocentric or allocentric
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of another familiar individual105. Together, these findings indicate that the body representation in the EBA is
relatively blind to higher-level factors such as identity
and sense of ownership, although this issue might be
revisited with stimuli that include full-motion, live views
of subjects’ own body parts.
a
EBA
b
vPMc
V1
500 ms
150 ms
500 ms
Until
key-press
rTMS;
delay, 150 ms
c
d
*
*
900
1,000
Response times (ms)
Response times (ms)
1,100
900
800
700
*
*
*
800
700
600
500
400
300
200
100
600
Body parts
Face parts
Motorcycle
parts
0
Form
Action
Figure 6 | Transcranial magnetic stimulation studies of the involvement of the
extrastriate body area in body-perception tasks. a | The cortical sites stimulated in two
studies82,134 — the extrastriate body area (EBA), the ventral premotor cortex (vPMC) and the
primary visual cortex (V1). b | Stimulus timing parameters. Two transcranial magnetic
stimulation (TMS) pulses were delivered in the interval of a short-term visual memory task
in which subjects were asked to compare the initial sample item with two test items, and in
which the comparison was based either on identity (form) or on the action implied by the
image. c | Repetitive TMS (rTMS) over the EBA (red bars) slows response times to judgments
on body parts, but not face or object parts, whereas rTMS over the V1 (green bars) and
sham stimulation (grey bars) have no effect82. d | rTMS over the EBA (red bars) and the vPMC
(white bars) has differential effects on judgments of body part identity (form) or implied
action: form tasks are disrupted by EBA stimulation, and action-judgment tasks are
disproportionately slowed by vPMC stimulation134. Panels a and d modified, with
permission, from Nature Neurosci. REF. 134 © (2007) Macmillan Publishers Ltd. Panels b and
c modified, with permission, from REF. 82 © (2004) Elsevier Science.
views (the FBA was not examined). Both studies found
higher activity in the left parietal cortex elicited by egocentric compared with allocentric views, in contrast to
a small but reliable enhancement of activity in response
to allocentric views in the right (but not the left) EBA.
Furthermore, the EBA showed no sensitivity to whether
the views were of the subject’s own body or the body
NATURE REVIEWS | NEUROSCIENCE
Body perception and emotion. We perceive others’ emotional states from multi-modal cues, including postures
and movements of the body and face. Although emotional face perception has been studied extensively107,108,
emotional body perception has, until recently, been
relatively neglected13. Recent behavioural and imaging
results have shown that emotional body postures can
bias attention and modulate activity in the visual cortex,
indicating that many previous findings on emotional face
processing might extend to emotional body processing.
For example, images of emotional body postures presented in the contralesional visual space can reduce the
attentional bias towards the ipsilesional space in patients
with hemi-spatial neglect7, as was shown previously for
emotional faces109. Furthermore, emotional body postures can be perceived implicitly in the absence of the
primary visual cortex110, which indicates that emotional
signals from the body might be partly encoded by a
subcortical visual pathway that involves the pulvinar
and amygdala and is known to have a role in processing
emotional faces111–114.
Recent fMRI studies have shown that, like emotional
faces, images of emotionally expressive bodies and
body parts increase activation in the visual cortex when
compared with neutral controls110,115–119. In one study,
short movie clips of emotional bodies, compared with
motion-matched movie clips of neutral bodies, selectively modulated the EBA and the FBA119. Furthermore,
this modulation was related, across subjects, to concurrent amygdala activation, indicating that the amygdala
is a likely source of this modulation. Although other
studies of emotional body perception did not functionally localize the EBA or the FBA120–122, the coordinates
of the observed modulations in these studies similarly
indicate that emotions expressed by bodies modulate
both of these body-selective regions, in the same way
that emotional faces modulate the activity of faceselective regions. For example, an fMRI comparison of
responses to static fearful body postures and emotionally neutral but semantically meaningful body postures
found enhanced activation in the fusiform gyrus115.
Other studies have reported similar emotional or motivational modulation at the approximate location of the
FBA in the fusiform gyrus116,118, and at the approximate
location of the EBA in the lateral occipitotemporal cortex110,116,117,123–125. The function of this emotional modulation could be to increase the speed and accuracy of
detection and recognition of emotionally salient bodily
actions.
The similarity of the effects of emotion on body and
face perception raises the question of how analyses of
emotions in these stimuli might interact — after all, bodies and faces are normally seen together. An ERP study
on emotion recognition found behavioural and neural
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evidence that facial and bodily emotions have congruent
effects: subjects recognized emotions from the face more
quickly and accurately when the emotion displayed by
the body was congruent126. Remarkably, ERPs showed
a congruency-related modulation as early as 115 ms
after stimulus onset. This early congruency effect shows
that emotional signals from both the face and body are
processed and integrated rapidly.
Action perception and ‘mirror’ systems. Viewing human
body actions activates the pSTS73,76–78,127, the parietal
cortex 78,128 and the ventrolateral premotor cortex
(vPMC)128–131, as well as the EBA and the FBA. Whereas
the pSTS responds specifically to the observation of biologically plausible motion patterns, the parietal cortex
and the vPMC are both involved in the perception and
the execution of actions8. After the discovery in the
macaque of neurons that respond to both the perception
and the performance of specific actions132 — known as
‘mirror’ neurons — these areas have been suggested to
constitute the human ‘mirror’ system8,133. Can the EBA
and the FBA be functionally dissociated from the ‘mirror’ system? If so, what role do these areas have in action
observation?
Two recent studies have reported functional dissociations between the EBA and the FBA and the frontoparietal mirror areas. TMS was used to investigate what
aspect of body processing is impaired after the disruption of activity in the EBA or the vPMC134. Subjects
had to discriminate either the identity or the (implied)
action of a pictured body part. Disruption of the EBA
with TMS impaired identity discrimination, whereas
disruption of the vPMC impaired action discrimination134. A similar dissociation was found in a recent fMRI
study135. Subjects viewed short movies of whole-body
actions, parsed into static frames that were presented
slowly, in either the correct or an incorrect sequence.
As expected, frontoparietal mirror areas and the pSTS
showed increased activity in response to coherent meaningful actions. By contrast, the EBA and the FBA showed
a greater response to the incoherent action sequences,
which contained more frame-to-frame differences in
body posture135. Together, these studies indicate that the
EBA and the FBA are not involved in the mirroring of
actions, but that they instead respond to the form of the
body that is present in displays of bodily actions.
One study that brings this proposal into question
found that group-average fMRI activity in a region that
is similar to the EBA (along with other parts of the visual
cortex) was greater when participants observed actions
in order to imitate them, rather than simply observe
them — a finding that would be consistent with EBA
involvement in a mirror system136. However, uncertainties in the localization of the EBA in this study, and
possible confounding influences of attention (due to
increased task demands and stimulus relevance in the
imitate condition), leave these results open to alternative
interpretations.
Instead, it seems that the EBA is not a core component of the ‘mirror neuron’ network. A more likely alternative, in our view, is that the EBA creates static, visual
644 | AUGUST 2007 | VOLUME 8
representations of the body — ‘snapshots’ that, along
with form and motion information from other areas,
contribute to a full visual representation of human
actions135. This might, in turn, support the perceptual
input to a mirror system, but in this hypothesis the EBA
is not a core part of this system, because it does not
exhibit mirror-like properties. These suggestions are in
line with the results of a detailed computational model of
biological motion perception, a key feature of which is a
collection of static body-form representations137.
Conclusions and future directions
There is now substantial evidence that the perception of
bodies, like that of faces, evokes a consistent and selective pattern of neural activity in the extrastriate visual
cortex. These findings raise many questions about the
functional role of these mechanisms and their interactions with other brain systems. Below we discuss some
of the numerous topics that we feel it is important for
future research to address.
Body and face perception in the visual cortex. Both
bodies and faces provide cues to the identity, emotions, intentions, age and gender of other people. To
a large extent, the similarities between bodies and
faces are reflected in similarities between the spatial
and temporal organization of the neural responses
they produce. Bodies and faces elicit globally similar
patterns of evoked potentials, similar effects on neural
activity when inverted or displaying emotion, adjacent
or overlapping selective activity in the monkey STS,
and closely overlapping fMRI responses in the posterior fusiform gyrus in humans. Notwithstanding these
global similarities, evidence from all of the methods
reviewed here points to functional and anatomical
distinctions between the neural systems involved in
face and body processing. These are sometimes subtle,
as in the closely overlapping body- and face-selective
fusiform responses. There are also broader differences;
perhaps the most striking is the apparent absence of
a body-related homologue of prosopagnosia after
damage to the extrastriate cortex.
The evidence for similar neural responses to bodies
and faces raises the possibility, as yet largely untested, that
body and face perception are closely functionally integrated18,69,126,138. A system that is built for fast and accurate
recognition of the information present in bodies and faces
would benefit from close and rapid interactions between
the systems that encode them. The posterior fusiform
gyrus, in which body and face representations are in close
proximity66,67, would be an ideal place for local interactions between face and body processing. For example,
previous fMRI studies have indicated that the FFA might
be important for processing face identity139–141. The FBA
could similarly be involved in processing body identity.
If so, this opens up the possibility that this general region
underlies our ability to identify other people on the basis
of cues from both the body and the face (BOX 2). The close
proximity and interaction of the FFA and FBA would be
especially useful when cues from either the face or the
body alone are not sufficient for recognition. For example,
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Box 2 | Body and face selectivity in the fusiform gyrus, and the expertise debate
A continuing debate in cognitive neuroscience concerns the interpretation of face selectivity in the human fusiform
gyrus. According to a domain-general, process-specific account, activation in this region is related to discriminating
among structurally similar exemplars of categories for which one has substantial expertise159. On this account, faces
activate this region more than other categories because they are automatically processed at the subordinate level, and
because of the greater expertise we have with faces than with most other object categories. By contrast, according to a
domain-specific account, face-selective responses in the fusiform gyrus reflect the activation of a ‘module’ for faces that
is not significantly involved in processing other object categories3. What bearing do the recent findings of closely
overlapping face- and body-selective responses in the fusiform gyrus60,66,67 have on this debate?
Faces and bodies are visually different, yet both are easily processed at the subordinate level, and most people have
developed substantial expertise for both. Thus, the finding of overlapping face- and body-selective responses is
consistent with the process-specific view, but only at a macroscopic level. At a finer-grained level, the existence of
spatially distinct face- and body-selective responses67 supports the domain-specific view, in that different categories
activate separate regions.
when seeing someone from a distance, facial features
alone might not be enough for identification69. In this
case, cues from the body, processed in the FBA, could
inform and perhaps support face-selective processing in
the FFA, and vice versa.
Functional role of body-selective regions. We have
reviewed evidence for two areas in the human visual
cortex that selectively respond to the form of the
human body. What is computed by the neurons in these
regions? One way to address this question is to test
for organizing principles that underlie the activity in
a region. For example, do the body representations
in the EBA and the FBA correspond to a psychophysical space centred around a ‘mean’ body shape, as
has been argued for faces in the FFA140? Furthermore,
these regions should be tested to determine whether
they exhibit retinotopy or somatotopy — that is, are
the representations of different body parts organized
systematically, or are some body parts overrepresented
relative to others, as has been found in the motor and
somatosensory cortices? We can also ask whether the
activity in these regions is sensitive to, or invariant
across, such stimulus variables as retinal size, viewpoint
or identity142. Given their distinct locations in the visual
system, there might also be dissociations between the
EBA and the FBA on one or more of these aspects79,143.
Finally, most studies on the EBA and the FBA have
focused on stimulus manipulations. New evidence
about their functional properties could be gained by
measuring their activity under different task conditions, or by correlating neural activity with behavioural
performance on a trial-by-trial basis144.
Recent advances in MVPA also offer a tool for understanding the neural representations in body-selective
brain regions. We have described how these analyses
are useful for interpreting overlapping fMRI activations64 (BOX 1). MVPA could have further applications
for understanding the neural encoding of bodies. Local
patterns of activity in the EBA and the FBA might carry
information about bodies that is lost when activity is
averaged across the voxels of these regions. Methods
that have been developed for extracting orientation
information from primary visual cortex activity65,145, or
motion direction from hMT activity146, could be adapted
NATURE REVIEWS | NEUROSCIENCE
to determine whether and how the EBA and the FBA
encode information about features such as gender, identity, posture or action. Also, whole-brain applications of
MVPA, which apply an analysis ‘searchlight’ across the
cortex to investigate the information content of different
brain regions147,148, could be used to determine whether
these features are encoded in brain areas that do not
respond significantly more to bodies than to other visual
stimuli, a possibility that has not been tested by previous
fMRI studies.
Interactions among body- and action-perception regions.
What is the interrelationship among the brain regions
that respond to static and dynamic bodies? How do these
areas coordinate in order to provide a coherent representation of other individuals? We have proposed that the
EBA (and possibly the FBA) provides static snapshots of
body postures to action perception systems, for example,
in the pSTS135. More generally, these regions might interact with the frontoparietal mirror areas involved in both
the perception and production of actions. Recent studies
using TMS have demonstrated the effects of stimulation
over some of these regions on performance in various
body- and action-perception tasks82,149,150. In light of
this, a further useful approach might be to measure the
modulation of fMRI activity in the EBA, FBA, pSTS, and
frontal and parietal areas that is caused by stimulation
of any one of these regions with TMS151. Changes in the
functional properties of regions that are distant from
the stimulation site could be used as an index of information flow through this network, and of the functional
contribution of each node within it.
Timecourse of body-selective neural activity. The visual
perception of bodies elicits a selective ERP response (the
N190)48 early in the timecourse of processing. Although
source analyses indicate that this component is generated in the extrastriate cortex, it is not clear whether it
is attributable to activity in the EBA, the FBA or some
other brain region. Constraining ERP source analysis
with fMRI would help to accurately localize the N190
(for an example, see REF. 152). This ERP–fMRI mapping
would also benefit from the identification of functional
distinctions between the EBA and the FBA; that is, to
the extent that these regions respond differentially to a
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stimulus or task manipulation, the N190 can be tested
with similar manipulations to determine whether its
response pattern matches that of one or the other region.
This effort may also be useful for developmental studies, for which ERP recordings are the primary source
of information on neural substrates. If the N190 can be
linked to a specific brain region, it might then be possible to make indirect inferences about the development
of specific cortical areas in infants.
If a link between specific brain regions and bodyspecific potentials can be identified, further ERP studies
might help to shed light on how activity in these regions
evolves over time, an aspect that fMRI studies cannot
measure. For example, the initial sweep of neural activity in the body-selective visual cortex might be largely
stimulus-driven, but activity in later stages might reflect
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Acknowledgements
The authors thank G. Thierry for helpful comments, and the
Biotechnology and Biological Sciences Research Council and
the Wales Institute of Cognitive Neuroscience for funding
support.
Competing interests statement
The authors declare no competing financial interests.
FURTHER INFORMATION
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