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Cerebral Cortex
doi:10.1093/cercor/bhm212
Paying Attention to Social Meaning: An
fMRI Study
Paula Tavares
1
, Andrew D. Lawrence
2,3
and Philip J. Barnard
2
1
Instituto de Biofı
´sica e Engenharia Biome
´dica, Faculdade de
Cie
ˆncias da Universidade de Lisboa, Campo Grande, 1749-016
Lisboa, Portugal,
2
MRC Cognition and Brain Sciences Unit, 15
Chaucer Road, CB2 7EF Cambridge, UK and
3
Wales Institute of
Cognitive Neuroscience, School of Psychology, Cardiff
University, CF10 3AT Cardiff, UK
Animations of simple geometric shapes are readily interpreted as
animate agents engaged in meaningful social interactions. Such
animations have been shown to activate brain regions implicated in
the detection of animate motion, in understanding the intentions of
others as well as areas commonly linked to the processing of social
and emotional information. However, attribution of animacy does
not occur under all circumstances and the precise conditions under
which specific regions are activated remains unclear. In a func-
tional magnetic resonance imaging study we manipulated viewers’
perspective to assess the part played by selective attention.
Participants were cued to attend either to spatial properties of the
movements or to the kind of social behavior it could represent.
Activations that occurred to the initial cue, while observing the
animations themselves and while responding to a postpresentation
probe, were analyzed separately. Results showed that activity in
the social brain network was strongly influenced by selective
attention, and that remarkably similar activations were seen during
film viewing and in response to probe questions. Our use of stimuli
supporting rich and diverse social narratives likely enhanced the
influence of top-down processes on neural activity in the social
brain.
Keywords: amygdala, animacy, autism, dorsomedial prefrontal cortex,
social cognition
Introduction
Animated movements of abstract geometric shapes can readily
be interpreted as depicting social events in which animate
agents are engaged in intentional activity such as pursuit, play,
or fighting (Heider and Simmel 1944; Scholl and Tremoulet
2000; Barrett et al. 2005). That these simple graphical
renderings give rise to intricate conceptual meanings has
attracted considerable interest from social neuroscientists.
Such animations provide an ideal platform for investigating
brain areas involved in the perception of animate motion versus
its mechanical counterparts as well as those areas involved in
the understanding of causality, of ‘‘Theory of Mind’’ and of
social interactions that are associated with particular move-
ment dynamics.
Several studies have shown that brain areas linked to
biological motion processing, social perception, the processing
of the intentions of others, emotional information, and social
knowledge can be activated when viewing Heider--Simmel
animations. They include the lateral fusiform gyrus (FG), the
temporo-parietal junction (TPJ), the amygdala, the posterior
cingulate cortex (PCC), the temporal poles (TPs) and the
medial prefrontal cortex (Castelli et al. 2000, 2002; Martin and
Weisberg 2003; Schultz et al. 2003; Ohnishi et al. 2004),
consistent with the idea of a distributed social brain network
(Brothers 1990). The nature of the activations is nevertheless
subject to considerable variation. Regional activation varies as
a function of the precise kinetic properties of the animations
(e.g., Blakemore et al. 2003), and with the nature of the viewing
task itself (e.g., Schultz et al. 2003). The typical patterns of
activation to animations usually interpreted as having social
content can also be markedly reduced in some individuals—
most notably those with autistic spectrum disorders (Castelli
et al. 2002). There is also evidence that Heider--Simmel
animations, being ambiguous, do not of necessity invoke social
interpretations. For example, Martin and Tversky (2003) have
shown that healthy individuals, on initial exposure interpret
Heider--Simmel animations as involving mechanical rather than
intentional movements (see Epley et al. 2007 for review).
Here, we focus on the role of selective attention in the
interpretation of animate motion and in the preparatory and
response phases that precede and follow the viewing itself. Our
aim is to clarify the part played by top-down influences on
regional activation. There is mounting evidence of the
importance of top-down influences in the interpretation of
ambiguous material within areas linked to the processing of
social stimuli. Kingstone et al. (2004) presented healthy
participants with ambiguous pictures and obtained different
patterns of activation when they were biased to see the images
as an inanimate entity (a car) or as an animate one (eyes and
a hat). The right superior temporal sulcus (STS) was only
observed to activate in the latter condition. In the context of
economic games, Gallagher et al. (2002) and Rilling et al.
(2004) found different patterns of neural activity when
participants thought they were playing with/against another
person or a computer. In McCabe et al. (2001) and Gallagher
et al. (2002) this difference was restricted to the dorsomedial
prefrontal cortex (DMPFC), whereas in Rilling et al. (2004)
increases were seen in DMPFC, PCC, TPJ, and STS. Kumaran and
Maguire (2005) found increased activity in FG, STS, TPJ, PCC,
TP, and DMPFC when people thought about the social, rather
than spatial, relations between the same individuals (see also
Mitchell et al. 2004).
Research specifically using Heider--Simmel style animations
has also involved a range of task conditions. Several studies
have relied on the most unconstrained form of task in which
observers passively view animations (Castelli, et al. 2000, 2002;
Martin and Weisberg 2003; Ohnishi et al. 2004). In 1 previous
study, Castelli et al. (2000) did provide prior knowledge of the
type of animation about to be encountered (‘‘Theory of Mind,’’
‘‘Goal Directed,’’ or ‘‘Random Motion’’) on 50% of occasions.
They found no significant differences in activation as a function
of that prior knowledge.
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Other studies suggest a role for top-down influences on
patterns of activation. Schultz et al. (2003) presented ‘‘Theory
of Mind’’ animations and asked their participants either to
decide whether the animated shapes were friends or not, or to
pretend that the shapes were cars and decide whether they
shared the physical property of being equally heavy. Whereas
a contrast between the physical and social decisions revealed
activation only in the dorsal bank of the intraparietal sulcus, the
opposite contrast between attention to social and physical
properties showed more widespread activation in the FG,
amygdala, TP, STS, and DMPFC. Schultz et al. (2004) created
animations depicting a pursuit scenario between 2 objects, in
which the chasing object used different strategies to reach the
target object. Attending to the pursuer’s strategy induced
slightly greater activation in the left superior temporal gyrus
than attending to the outcome of the chase. Blakemore et al.
(2003) developed animations that varied both whether the
movement of abstract figures had animate properties and the
contingencies that held between the movement of 2 figures.
Viewers’ attention was either directed at the contingencies
that held between the 2 figures or left undirected. Right middle
frontal gyrus and the left STS were only engaged when
attention was directed to the contingent nature of the
movement patterns. However, Blakemore et al. (2003) found
no evidence of differential engagement of TPs, amygdala, right
STS, or DMPFC when attention was directed at salient
intentional contingencies. One possibility is that these areas
only respond when animations contain cues that support the
formation of schematically rich representations of mental states
of others interacting in social ways, and then only when
attention is directed towards salient features.
Given the ambiguous nature of Heider--Simmel animations,
and findings suggesting that ‘‘social’’ brain areas such as STS are
also activated in the processing of nonsocial motion processing
(Frith and Frith 2003; Dakin and Frith 2005), together with
uncertainty as to the relative roles of bottom-up versus top-
down social processing deficits in autism (Behrmann et al.
2006; Bird et al. 2006), there is a pressing need to acquire more
data on regional brain activation under circumstances where
identical animations are viewed from different observational
perspectives, conditions required to rule out simple stimulus-
driven effects. Our study therefore sought to extend previous
work by focusing on the effects of top-down attentional
perspective, in the context of relatively rich animation
scenarios of simple shapes that are open to interpretation in
terms of intentional and emotional attributes.
Our attentional manipulation was realized by directly cueing
the participants to pay attention to social or spatial aspects of
the figures’ behavior and then probing them after viewing the
animations with a question about either the social behavior of
the figures or their use of physical space. The animations were
developed to sample movement patterns characteristic of
affiliative, antagonistic, and indifferent behaviors (e.g., ap-
proach, avoidance, contact strength—Michotte 1950) in order
to ensure that when asked about social attributes they would
have attended to both interpersonal and affective dimensions.
In order to further clarify the role of top-down processes, we
scanned the participants while attending to the initial cue and
when responding to the final probe, in addition to ‘‘on-line’’
viewing of the animations. By drawing attention to the social
attributes of schematically rich behaviors, we predicted that,
the full spectrum of brain areas implicated in social cognition
would be engaged. By contrast, we predicted that tracking the
motion of such stimuli in space would activate parietal and
frontal areas recruited by diverse spatial processing demands.
Further, we anticipated some activation in areas involved in
motion and social processing when compared with baseline
during the processing of the initial social cue. We also
expected a good deal of overlap in the areas activated during
actually viewing the animations and when responding to the
verbal probes (Bargh 2006).
Method
During functional magnetic resonance imaging (fMRI), participants
were asked to watch a sequence of short animations. Each showed 2
abstract figures, or ‘‘sprites,’’ moving with properties that support
varying interpretations (indifferent, antagonistic, affiliative social
behaviors) in a structured environment or ‘‘sprite world.’’ Participants
were precued to attend either to the social behavior that could
underlie the movement of the sprites or to the spatial aspects of the
same movement. After each animation they were posed a simple true--
false question concerning social or spatial aspects of the animation just
viewed. Participants were scanned during all 3 phases. The baseline
condition involved presentation of a cue followed by a fixation cross for
a duration matched to that of the animations and a direct instruction to
make 1 of 2 responses.
Participants
Sixteen right-handed healthy female, native English speakers (mean age
=24 years, standard deviation [SD] =4 years) participated. The study
was approved by the Cambridgeshire Local Research Ethics Committee.
Written informed consent was obtained from all participants. Individ-
uals with a history of inpatient psychiatric care; neurological disease, or
head injuries were excluded, as were individuals on medication for
anxiety or depression.
Materials
The stimuli consisted of 14-s duration animations, displaying the
motion of 2 sprites—1 green circle and 1 blue circle. The sprite world
was 2-dimensional and included some obstacles to motion in a straight
line (see Fig. 1 for a schematic representation of the basic layout of
each animation). Three categories of animation were constructed, in
order to broadly sample different interpersonal situations: 1) affiliative,
friendly interactions between the 2 sprites; 2) antagonistic, hostile or
suspicious interactions between the 2 sprites; and 3) indifferent non-
interactions between the 2 sprites. Animations were created using the
Figure 1. Schematic representation of the sprites and sprite world.
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software package Blender version 2.1 (Blender Foundation, www.
blender.org).
In the affiliative animations, the 2 sprites entered from different sides
of the sprite world. They approached each other and touched or moved
in a mutually coordinated way throughout. They then left the sprite
world together at a gentle pace. In the antagonistic animations, the 2
sprites entered from different sides of the sprite world and approached
each other. In this case, they initially kept at a small distance from each
other, while moving around slowly. In one-half of the animations, the
circles then repeatedly approached each other, made brief contact and
retreated rapidly, a pattern suggesting physical aggression. In the other
half they never actually touched each other, but moved in close
proximity to each other suggestive of suspicion. At the end of the
sequence, the 2 sprites left the scene in different directions.
In the indifferent animations, the 2 sprites again entered from different
sides of the sprite world and passed each other without interacting. The
sprites continued to move around, sometimes stopping and changing
direction, but never approaching each other or moving in any way that
might suggest an influence of one on the other. As with the antagonistic
animations, the sprites left the scene independently at a gentle pace.
Tasks
During scanning, each animation was preceded by a cue word, either
Behavioral or Spatial, specifying the nature of the probe that would
follow the next animation, thereby constraining the perspective from
which the animations were to be viewed. For the Behavioral cue,
participants were instructed to identify what kind of interaction might
be happening between the 2 circles. For the Spatial cue, participants
were instructed to pay attention to various aspects of the motion of the
2 circles, such as their speed, trajectory, position of entering/exiting
the scene, etc. After the presentation of each animation, a probe
statement appeared describing the contents of the animation, and the
participant had to decide (true/false) if the statement could appropri-
ately describe the behavior in the animation just viewed. Examples of
Behavioral statements included ‘‘Soldiers engaged in hand to hand
combat,’’ ‘‘People were shopping in a supermarket,’’ and ‘‘An old lady
was helped by a friend to carry her bags.’’ Examples of Spatial
statements included ‘‘The circles passed each other only once’’ and
‘‘The blue circle collided with one of the lines.’’ Behavioral and Spatial
probe statements were matched for number of words and syllables and
had been pretested to reliably elicit either a true or false response to
the particular animation to which it referred. Each animation and each
probe statement was presented only once, with 50% true and 50% false
responses to the probes. The cue word was presented for 1.8 s, the
animation for 14 s, and the probe statement for 7.5 s. In addition,
a baseline fixation condition was used. In this condition, the cue word
Cross appeared, followed by a fixation cross for 14 s, and then
a statement saying that the participants should press either the left or
right response button. Temporal jitter was introduced between the
cue, animation and probe statement phases to ensure they could be
modeled separately. There were 12 examples of each of the animations
in each of the 2 conditions (Behavioral or Spatial cue) plus baseline
condition, presented in pseudorandom order. The experiment took an
average of 16.3 min to complete.
Post-task Ratings
Participants were invited to return to do a follow-up ratings task. They
were asked to classify the behavior shown in each of the animations
they had seen in terms of the extent to which the sprites appeared
animate or alive, and the emotional valence associated with the
animation, using visual analogues scales with lines 11.5 cm long. For the
animacy judgment, ratings were made by marking a point on the line,
anchored by the terms ‘‘not alive’’ at one end and ‘‘fully alive’’ at the
other. For valence ratings, the scale ranged from ‘‘positive’’ at one end
through ‘‘indifferent’’ labeled at the center point of the line, to
‘‘negative’’ at the other end.
Image Acquisition
Blood oxygenation level--dependent contrast functional images were
acquired with echo-planar T
2
*-weighted (EPI) images using a Medspec
(Bruker, Ettlingen, Germany) 3-T MR system with a head coil gradient
set. Each image volume consisted of 21 interleaved 4-mm thick slices
(interslice gap: 1 mm; in-plane resolution: 2.2 32.2 mm; field of view:
20 320 cm; matrix size: 90 390; flip angle: 74°; echo time: 27.5 ms;
voxel bandwidth: 143 kHz; repetition time 1.6 s). Slice acquisition was
transverse oblique, angled to avoid the eyeballs, and covered most of
the brain. Six hundred and ninety-five volumes were acquired, and the
first 6 volumes were discarded to allow for T
1
equilibration.
Image Analysis
Neuroimaging data were analyzed using statistical parametric mapping
software (Wellcome Trust Centre for Neuroimaging, London, UK).
Standard preprocessing was conducted, comprising slice timing
correction, realignment, undistortion (Cusack et al. 2003), and masked
normalization of each participant’s EPI data to the Montreal Neurolog-
ical Institute (MNI) International Consortium for Brain Mapping
template (Brett, Leff, et al. 2001). Images were resampled into this
space with 2-mm isotropic voxels and smoothed with a Gaussian kernal
of 8-mm full-width at half-maximum. For the epochs of the animations
and probe statements, condition effects were estimated for each
participant at each voxel using boxcar regressors for the 14-s animation
period and 7.5-s probe statement period, respectively, convolved with
a canonical hemodynamic response function (HRF) in a general linear
model, with spatial realignment parameters included as regressors to
account for residual movement-related variance. Activation to the cue
word was modeled using a canonical HRF, plus time and dispersion
derivatives. A high-pass filter was used to remove low-frequency signal
drift, and the data were also low-pass filtered with the canonical HRF.
Activation contrasts between conditions were estimated for each
participant at each voxel, producing statistical parametric maps.
Random-effects analysis was conducted to analyze data at a group level.
We used small volume correction (SVC) (Worsley et al. 1996) for
multiple comparisons applied at P<0.05 (family-wise error), following
an initial thresholding of P<0.002 (uncorrected). Amygdala, TPs, and
PCC regions of interest (ROIs) were defined using structural templates
derived by automated anatomic labeling (Tzourio-Mazoyer et al. 2002).
The TP ROI resulted from the sum of the middle and superior TPs
structural templates. Spherical ROIs (10-mm radius spheres) were
created for the remaining ROIs. For the lateral FG (central coordinates
–42, –49, –19; 40, –48, –16), TPJ (–56, –56, 17; 57, –53, 17), DMPFC (BA 9/
10/32) (–4, 60, 32; 7, 55, 28), and ventromedial prefrontal cortex (±2,
48, –12), by computing the average of the reported activation
coordinates for these regions across previous imaging studies of
animate motion (Castelli et al. 2000, 2002; Martin and Weisberg 2003;
Schultz et al. 2003; Ohnishi et al. 2004). For the STS we used the
coordinates (±54, –34, 4) taken from Schultz et al. (2003). For the
dorsolateral prefrontal cortex (DLPFC) (±34, 36, 24), dorsal anterior
cingulate cortex (dACC) (±4, 14, 36), and parietal cortex (±37, –53, 40)
we used coordinates from Duncan and Owen (2000). For the frontal
eye fields (FEF) we used the coordinates (±32, –2, 47) from Silvanto
et al. (2006), and for the motion-selective homolog of macaque area MT
(hMT) we used the coordinates (±52, –60, 5) from Blakemore et al.
(2001).
For anatomical labeling purposes, activation coordinates were
transformed into the Talairach and Tournoux (1988) coordinate system
using an automated nonlinear transform (Brett, Christoff, et al. 2001)
and labeled with reference to the Talairach Demon database (http://
ric.uthscsa.edu/projects/talairachdaemon.html). For visualizing activa-
tions, group maps are overlaid on the ICBM 152 structural template, an
average T
1
-weighted image of 152 individuals coregistered to MNI
space. Activations are reported using (x,y,z) coordinates in MNI
standardized space.
Results
Behavioral and Rating Data
Participants took an average of 3.3 s to respond to the probes
and the times to respond showed no main effects of scenario
(affiliative, antagonistic, and indifferent) F
1,12
=0.84, P=0.46,
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task (Behavioral,Spatial)F
1,12
=0.03, P=0.87, or response
(true, false) F
1,12
=0.20, P=0.66 and there were no interactions
between category, type of statement, and task. Participants
provided the expected response to 92% of the probes and error
rates were comparable across the types of animations
(affiliative, antagonistic, and indifferent) v
2
=2.9, df =2,
P=0.24, for the 2 viewing tasks (Behavioral,Spatial)v
2
=
0.072, df =1, P=0.79 and for the 2 responses (true, false) v
2
=
0.129, df =1, P=0.72. There was therefore no evidence to
indicate that the Behavioral and Spatial probe questions
differed in difficulty.
The post-task visual analogue scale based valence ratings
(see methods section under ‘‘post-task ratings’’) confirmed that
the films were rated as intended (affiliative—positive; antago-
nistic—negative; indifferent—neutral). There was however,
some evidence that the animated materials differed in the
nature of their most likely interpretation. The indifferent set of
animations was rated as less alive than the affiliative (Z=–3.01,
P<0.01) and antagonistic animations (Z=–3.01, P<0.01),
which did not differ from each other (affiliative mean 9.5 SD
2.24; antagonistic mean 9.3 SD 2.19, indifferent mean 5.2 SD
2.09).
fMRI Data
Preparatory Cues
When the response in the ROIs to each of the Behavioral and
Spatial cues was compared with fixation, there was significant
activation in hMT (Behavioral: 48, –68, 2, Z=3.56, P
SVC
=
0.024; Spatial:–50, –68, 6, Z=3.75, P
SVC
=0.012, and 48, –68, 2,
Z=3.53, P
SVC
=0.023); in the posterior parietal ROI
(Behavioral:–28, –52, 46, Z=3.26, P
SVC
=0.053; Spatial:–30,
–52, 48, Z=4.08, P
SVC
=0.004, and 30, –56, 44, Z=3.19, P
SVC
=
0.055); and in FEFs (Behavioral:–26, –2, 54, Z=5.07, P
SVC
<
0.001 and 24, –4, 52, Z=4.72, P
SVC
<0.001; Spatial:–24, –4, 52,
Z=4.95, P
SVC
<0.001, and 24, –4, 52, Z=4.55, P
SVC
<0.001)
(Fig. 2).
When the response in the ROIs to the Behavioral cue was
compared with that of the Spatial cue, there was no significant
activation in any of our ROIs.
For the opposite contrast (Spatial >Behavioral cue), there
was increased activity in the posterior parietal ROI (–30, –50,
42, Z=3.85, P
SVC
=0.01); DLPFC (–34, 46, 24, Z=4.07, P
SVC
=
0.005 and 30, 42, 24, Z=3.89, P
SVC
=0.009); dACC (0, 16, 44,
Z=3.57, P
SVC
=0.022); and FEFs (–40, 2, 52, Z=3.21, P
SVC
=
0.055).
Animation Viewing
Regions showing increased activation during viewing of
interactive animations (affiliative and antagonistic) rated
as strongly animate, relative to noninteractive (indifferent)
animations rated as more ambiguously animate. When
contrasting activation for the highly interactive scenarios, rated
as highly animate (affiliative and antagonistic) relative to the
indifferent scenarios, rated as more weakly animate, there was
increased activity in FG (bilaterally) (–42, –40, –24, Z=3.84,
P
SVC
=0.011 and 48, –48, –20, Z=3.89, P
SVC
=0.009), right TPJ
(54, –46, 12, Z=3.54, P
SVC
=0.029), right STS (56, –40, 2,
Z=4.64, P
SVC
=0.001), amygdala (–18, 0, –20, Z=4.06, P
SVC
=
0.002 and 24, 0, –24, Z=3.19, P
SVC
=0.026), and hMT (48, –60, 6,
Z=3.81, P
SVC
=0.012 and –48, –62, 6, Z=3.46, P
SVC
=0.032).
Regions showing increased activation during film viewing
when animations are preceded by the Behavioral cue,
compared with the Spatial cue. All of the social brain ROIs
showed increased activation when animation viewing followed
the Behavioral cue relative to the Spatial cue: right FG,
amygdalae (bilaterally), PCC (bilaterally), right TPJ, right STS,
DMPFC (bilaterally), and TP (bilaterally) (Fig. 3, Table 1).
Activation in right TPJ, right STS, right TP, right amygdala,
bilateral FG, and right PCC was also increased when comparing
animation viewing following the Behavioral cue relative to
baseline, as was activation in posterior parietal cortex, DLPFC,
hMT, and FEFs (see online Supplementary Table 1).
Does the modulatory effect of task cue (Behavioral vs.
Spatial) depend on the manifest emotionality of the
animations? When comparing the emotionally valenced
(affiliative and antagonistic) animations relative to the neutral,
indifferent animations for the Behavioral >Spatial contrast,
there was no significantly increased activity in any of our
predefined ROIs, suggesting that increased activation for the
Behavioral >Spatial contrast cannot be due to the indirect
influence of the manifest emotionality of the animations.
Figure 2. Regional activations to the Behavioral and Spatial cue events relative to the fixation cue. (A) Activation in the FEF. (B) Activation in hMT. Cross-hair centered on group-
average location of hMT reported in Downing et al. (2007).
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Regions showing increased activation during film viewing
when films are preceded by the Spatial cue, compared with
the Behavioral cue. For the contrast Spatial >Behavioral,
increased activity was seen in the posterior parietal cortex
(–32, –48, 42, Z=3.87, P
SVC
=0.009 and 38, –44, 46, Z=3.09,
P
SVC
=0.07), DLPFC (–34, 38, 32, Z=3.37, P
SVC
=0.037 and 38,
40, 32, Z=4.03, P
SVC
=0.005), dACC (2, 18, 44, Z=3.60,
P
SVC
=0.02), and FEFs (–30, 2, 46, Z=5.47, P
SVC
<0.0001 and 32,
–4, 54, Z=3.48, P
SVC
=0.027), but not in hMT. This difference
did not vary as a function of animation type. Activation in all
these regions, and also in the vicinity of hMT was increased
when comparing animation viewing following the Spatial cue
relative to fixation (data not presented).
Probe Statements
Regions showing increased activation in response to
Behavioral probe statements, compared with the Spatial
probe statements. All social brain ROIs showed increased
activation in response to Behavioral probes relative to Spatial
probes: right FG, amygdala (bilaterally), PCC (bilaterally), STS
(bilaterally), right TPJ, TPs (bilaterally), and DMPFC (bilaterally)
(Fig. 4, Table 2). There were no significant differences in these
ROIs when comparing probes related to the emotional versus
the indifferent, neutral animations.
Table 1
Neural regions showing increased activation to the animations during selective attention to
Behavioral versus Spatial properties
Brain regions MNI coordinates Zscore P
SVC
PCC 652 22 3.33 0.029
12 52 30 3.28 0.026
DMPFC 8 60 24 4.06 0.005
4 56 20 4.25 0.003
TPJ 54 50 14 4.52 0.001
STS 50 30 4 4.08 0.005
TP 44 8 30 4.43 0.004
40 26 24 3.74 0.042
Amygdala 26 2 26 3.46 0.01
26 0 24 3.34 0.016
FG 42 54 18 3.86 0.009
Note: NB for abbreviations see Figures 3 and 4.
Figure 3. Regional activations to the animations following the Behavioral relative to
the Spatial cue. (A) PCC and DMPFC. (B) Right TPJ and anterior STS. (C) Amygdala.
(D) FG. (E) Parameter estimates.
Figure 4. Regional activations to the Behavioral relative to the Spatial probe
statements. (A) PCC and DMPFC. (B) Right TPJ and anterior STS/TPs. (C) Amygdala.
(D) FG. (E) Parameter estimates.
Table 2
Neural regions showing increased activation to Behavioral relative to the Spatial probes
Brain regions MNI coordinates Zscore P
SVC
PCC 0 50 26 4.52 \0.001
250 28 4.45 \0.001
DMPFC 8 56 34 5.44 \0.0001
2 58 28 5.26 \0.001
TPJ 52 64 22 3.19 0.06
58 60 22 4.16 0.004
STS 52 32 2 3.65 0.018
50 36 4 4.34 0.002
TP 44 26 20 3.95 0.019
44 20 30 4.53 0.003
Amygdala 20 816 4.33 0.001
30 4 28 3.92 0.003
FG 42 48 22 4.31 0.002
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Amygdala, left FG, left STS, TPJ, TP, and DMPFC were also
more active when comparing response to the Behavioral
probes relative to the control condition (online Supplementary
Table 2).
Regions showing increased activation in response to Spatial
probe statements, compared with the Behavioral probe
statements. For the opposite contrast (Spatial >Behavioral
probes) only the left posterior parietal cortex ROI showed
increased activation (–36, –48, 44, Z=4.72, P
SVC
=0.001).
Relative to baseline, both the Spatial and Behavioral probes
produced increased activation in posterior parietal cortex,
dACC and FEFs (online Supplementary Table 3).
For each of the 3 phases (cue, animation, probe statements),
regions surviving a threshold of P<0.05, whole brain corrected
(family-wise error), with a cluster extent minimum of 5 voxels,
for the Behavioral versus Spatial and Spatial versus Behav-
ioral contrasts are presented in Table 3. The only region
outside our predefined ROIs more active for the Behavioral
versus Spatial contrast was the right parahippocampal gyrus,
during the response to the probe statements.
Discussion
We examined the influence of manipulating observers’ selec-
tive attention, on brain activity in preparation for, during, and in
response to probe questions concerning animations of simple
geometric shapes that could be interpreted as animate agents
engaged in meaningful social interactions. This was achieved by
precueing participants to attend either to the social behavior
underlying their interaction or to the spatial aspects of their
movement patterns. Our design allowed us to analyze
separately brain activity to the cue, to the animations, and to
the probe statements. Several novel findings emerged.
Activations during the Preparatory Cue Period
Relative to the control Cross condition, presentation of both
the Behavioral and Spatial cue words lead to increased
activation in the FEF and posterior parietal cortex, as well as in
motion sensitive area hMT. Relative to the Behavioral cue, the
Spatial cue produced increased activation in posterior parietal
cortex, the vicinity of the FEF, the DLPFC, and dACC. These
results are consistent with prior studies on the preparatory
deployment of spatial attention and attention to motion (e.g.,
Luks and Simpson 2004; Caplan et al. 2006). The same
frontoparietal regions have also been implicated in nonspatial
orienting, for example to semantic categories (Cristescu et al.
2006), and may underpin a general purpose attentional
orienting network (Corbetta and Shulman 2002).
Notably, there was a lack of Behavioral cue-related activity
in social brain regions. This relative lack of preparatory activity
might relate to Raichle and colleagues’ notion of a default
brain state (Raichle and Gusnard 2005)—in particular the idea
that social brain regions form part of a default mental state
involved in constantly monitoring for self-relevant events
(Iacoboni et al. 2004). If this is indeed the default state then
there is little need for a ‘‘baseline shift’’—these systems are
constantly ‘‘primed,’’ ready for action, as part of our default
mode of interacting with the world. Alternatively, the
Behavioral cue might be more indeterminate than the Spatial
cue, and hence not trigger specific preparatory processes
beyond those involved in motion processing and perceptual
event segmentation (Zacks et al. 2001) (i.e., hMT, FEF), which
have been argued to play important roles in social perception
(Michotte 1950; Newtson 1980; Premack and Premack 1995;
Baldwin and Baird 2001).
Activations during Animation Viewing
Again, during film viewing, and following both cues, there was
increased activity in regions implicated in motion processing,
event segmentation, sustained attentive tracking, and spatial
working memory, including the posterior parietal cortex, hMT,
FEFs, and DLPFC (Culham et al. 1998). Increased activity
following the Spatial relative to the Behavioral cue was found
not only in the posterior parietal cortex (Schultz et al. 2003),
but also in DLPFC, FEF, and dACC. This suggests an increased
demand on sustained attentional tracking and spatial working
memory processes in the Spatial condition, presumably as
participants could rely on ‘‘gist’’-based knowledge to answer
the Behavioral probe statements, rather than encoding the full
detail of sprite motion. Notably, there was no difference in
activity in area hMT between the 2 conditions, consistent with
a role for motion cues in the perception of intention and social
behaviors.
When comparing activation to the highly interactive,
strongly animate (antagonistic and affiliative), relative to the
more weakly animate (indifferent) animations, regardless of
attentional set, we saw activity in regions previously implicated
in processing biological motion and animacy (see introduc-
tion). These included TPJ, STS, FG and amygdala, as well as
activity in motion processing areas including hMT, but not in
DMPFC, PCC, or TP, consistent with the former, but not the
latter, areas in detecting animacy/social interactivity from
perceptual/motion cues (Allison et al. 2000; Schultz et al. 2003;
Heberlein and Adolphs 2004; Frith and Frith 2006). However,
the animations also differed in emotional valence, and so this
comparison cannot unambiguously disentangle responses to
animacy from those due to manifest emotion.
More importantly, similar to Schultz et al. (2003), when
comparing animation viewing following the Behavioral versus
the Spatial cue, there was increased activity in all our social
brain ROIs: the amygdala, PCC, right FG, right TPJ, right STS,
TPs, and DMPFC, demonstrating that activity in all these areas is
Table 3
Neural regions showing increased activation at P\0.05 whole brain corrected, cluster minimum
5 voxels for key contrasts
Brain regions MNI coordinates Zscore P
Cue
Behavioral versus Spatial
No suprathreshold voxels
Spatial versus Behavioral
Cerebellum 10 61 10 5.42 0.007
Animations
Behavioral versus Spatial
DMPFC 12 50 36 5.00 0.036
Spatial versus Behavioral
Middle frontal gyrus 30 0 42 5.47 0.005
Phrases
Behavioral versus Spatial
Amygdala 26 12 11 5.38 0.006
DMPFC 8 56 29 5.44 0.004
2 57 23 5.26 0.012
Parahippocampal Gyrus 22 20 17 5.48 0.004
16 33 7 5.54 0.002
Spatial versus Behavioral
No suprathreshold voxels
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strongly influenced by selective attention to the social mean-
ings of the movements. In Schultz et al. (2003), different films
were used for the social and physical decisions, and so an
influence of stimulus material on those findings cannot be
excluded. Our results extend those findings by showing that,
for identical stimulus materials, a manipulation of selective
attention towards social meaning increases activity in essen-
tially all areas implicated in social processing. This strikingly
demonstrates the widespread influence of goal-directed selec-
tive attention, and supports the idea of a distributed social brain
network (Brothers 1990).
Wheatley et al. (2007) have very recently reported similar
widespread increases in activity throughout the social brain
when people were biased to interpret the same moving shapes
as either animate (e.g., ice-skater) or inanimate (e.g., spinning
top), although they did not find differences in the TPs. Their
study manipulated the background context in which the
shapes appeared. It is logically possibly, however, that the
background contexts themselves contained certain stimuli that
could drive areas of the social brain in a bottom-up fashion. In
our study, all perceptual inputs were identical in both social
and nonsocial conditions.
Our results and those of Wheatley et al. (2007; see also
Kumaran and Maguire 2005) contrast with previous studies
manipulating social attention reporting relatively focal top-
down influences, notably in DMPFC (see Introduction). Sugase
et al. (1999) argue that the ability of neurons in visual temporal
cortex to make fine social perceptual distinctions depends on
feedback from more anterior regions including prefrontal
cortex and amygdala. Frith and Frith (2006) argue that activity
in DMPFC is concerned with the ‘‘set’’ appropriate to social
tasks and is the source of top-down signals that modify signal
processing in more posterior brain regions such as STS,
concerned with the analysis of social signals. Although our
results are consistent with these suggestions, they suggest
there may be multiple neural sources of social attention.
One issue with interpreting the main effect of cue
(Behavioral vs. Spatial) when viewing the animations is that
some of the scenarios depicted were emotional—one could
argue that differences in brain activity for this contrast activity
reflect a ‘‘side-effect’’ of increased attention to emotion in the
social condition. Importantly, we found no significant differ-
ences in activity in any ROI, for the manifestly emotional
scenarios relative to the neutral scenarios, suggesting that
manifest emotion per se is not driving differential activation in
these regions under the Behavioral cue (see also Martin and
Weisberg 2003).
Activations during Probe Statements
A key feature of our experimental design was the investigation
of the brain responses to probe statements concerning the
social interactions depicted in the animations, in the absence of
activity triggered by the stimuli themselves. Relative to a low-
level decision baseline, activation was seen to both Behavioral
and Spatial probe statements in the posterior parietal cortex,
FEF and dACC. These joint activations likely reflect activity in
a system responsible for the planning and execution of reading
saccades (Leff et al. 2000). As with the processing of the cue
and when viewing the animations, there was increased activity
in the posterior parietal cortex for the Spatial probes relative
to the Behavioral probes, consistent with the well known
involvement of parietal regions in spatial cognition, including
spatial imagery, reasoning, problem solving, and memory
(Cabeza and Nyberg 2000).
With regard to our primary focus, the Behavioral versus
Spatial contrast, with exception of the left TPJ and left FG, all
remaining social brain ROIs showed increased activation. This
suggests modulation of all these regions in the absence of any
direct perceptual input, including regions linked to bottom-up
elements of social perception (Allison et al. 2000; Frith and
Frith 2006) and social behavior (Amaral 2003), when partic-
ipants respond to probe statements concerning the social
meaning of the interactions. That is, merely thinking about the
social properties of a remembered stimulus seems to be
sufficient to engage all these brain regions (Bargh 2006). These
activations were also seen relative to a low-level baseline
condition, suggesting they are not the result of relative de-
activations to the Spatial probes, but are true activations
(Pessoa et al. 2005).
A notable feature of the probe statements is that they did not
ask explicitly about complex mental states, such as beliefs, but
rather required participants to consider the interactions of the
figures in terms of rather generic situations involving social
agents sampled across quite diverse domains (e.g., shopping,
playing, fighting, visiting an art gallery, business activities, etc).
Activation in certain brain regions, for example, TPJ could
result from the spontaneous use of component Theory of Mind
processes (Saxe 2006). Additionally, to answer the rather
generic Social probe statement, as opposed to the much more
referentially specific Spatial probe statements, participants
presumably utilize higher-order knowledge structures such as
models, scripts, schemas, narratives, or context frames that
represent generic knowledge. People understand their social
worlds via schemas for personalities, selves, roles, groups
(stereotypes), social events, emotional episodes, and narra-
tives, which guide encoding, remembering, and responding
(Parkinson 1995; Fiske 2004).
The anterior TPs have been argued to play a critical role in
social concept processing (Zahn et al. 2007), in narrative
processing (Maguire et al. 1999), in social schemas (Frith and
Frith 2003), and in emotional context frames (together with
the parahippocampal cortex) (Mobbs et al. 2006). The
amygdala has been linked to emotional gist memory (Adolphs
et al. 2005). The DMPFC has also been implicated in schema-
level knowledge representation. For example, Heberlein and
Saxe (2005) found that DMPFC was activated when partic-
ipants made personality judgments, whereas Teasdale et al.
(1999) found increased DMPFC activity when viewing pairs of
pictures and captions which when combined formed a co-
herent (emotional) schematic model of ‘‘implicational mean-
ings’’ (Teasdale and Barnard 1993). DMPFC and PCC have also
been implicated in autobiographical memory retrieval and
prospection (Svoboda et al. 2006; Buckner and Carroll 2007).
There are strong narrative and schematic influences on both
autobiographical memory recall and prospective memories
(Eldridge et al. 1994), suggesting an interesting reason for such
overlap. We are currently examining how individual differences
in schema elaboration modulate activity in these regions to
social animations.
Our data confirm a role for TPs, DMPFC, and PCC in higher-
level aspects of social cognition, perhaps related to represen-
tation of complex social knowledge structures, but additionally
suggest that a rather broad network of structures is activated
when participants access and utilize social knowledge,
Cerebral Cortex Page 7 of 10
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including FG and STS, regions often argued to support social
perception rather than social knowledge (Allison et al. 2000),
as well as social behavior (e.g., amygdala) (Amaral 2003). As
mentioned above, it is likely that activation in sensory regions
results from top-down feedback from structures like DMPFC,
PCC, TP, and amygdala. According to certain models of
semantic cognition (McLelland and Rogers 2003) semantic
knowledge arises from the interactive activation of modality-
specific representations of perceptual attributes (such as
patterns of movement, etc.)—these different modality-specific
representations communicate with one another through
a common set of amodal representations coded in the anterior
temporal lobes. Our results are also consistent with the
influence of priming social knowledge structures on percep-
tion, motivation, and behavior (Bargh 2006).
Although related studies of ‘‘spontaneous’’ social understand-
ing based on animate motion have not shown engagement of
TPs (Wheatley et al. 2007), it is likely that the increased
complexity and richness of the animations we used played
a key role. Our animations were not only relatively rich in
perceptual terms (Mar et al. 2007) but also in terms of the
potential for assigning highly varied meanings to the scenarios
in our sprite world (Iacoboni et al. 2004), and this likely
resulted in our findings of activation throughout so many areas
of the social brain, including anterior TPs, when participants’
were instructed to attend to the social meaning of the
interactions.
An important topic for future research will be to determine
whether activity in ‘‘social’’ brain regions is specific to social
stimuli, or could be similarly active for certain nonsocial stimuli
(Mitchell 2007). For example, it has been proposed that
schematic representations of self, others, and world are all
encoded at an abstract, generic level of representation that
integrates over both cognitive and affective dimensions derived
from multimodal inputs and the interpretation of referentially
specific meanings (Teasdale and Barnard 1993). Such a position
would be consistent with extensive evidence that key areas of
the social brain integrate over multimodal sources (Pandya and
Yeterian 2001). Similarly, Ochsner et al. (2004) argue that
regions such as DMPFC respond to stimuli ‘‘for which an
attributed meaning is a meta-level emergent property of
multiply interpretable inputs, that in and of themselves, do
not directly imply a single interpretation’’.
Summary and Conclusions
In an fMRI study we manipulated the viewers’ perspective to
assess the part played by top-down attentional influences on
neural activity to animations of simple geometric shapes that
can be interpreted as animate agents engaged in social
interactions with meaningful properties. Our results highlight
that widespread structures throughout the social brain are
strongly influenced by social processing goals, consistent with
them operating as a common processing network. Such
influences did not appear to depend on there being manifest
emotional content in the animations. Notably, very similar
patterns of neural activation were seen when viewing films and
when remembering and responding to them. This indicated
that activity in regions, such as the FG, linked to perceptual
processing of social stimuli, does not need to be perceptually
triggered, but can be re-activated as a result of input from top-
down social attention and knowledge systems, consistent with
several current ‘‘interactionist’’ theoretical models of social
knowing. Our results also emphasize the close relationship
between social knowledge structures, perception, motivation,
and behavior. Finally, our use of stimuli supporting the
generation of rich and diverse social narratives in this study
likely enhanced the influence of top-down processes on neural
activity in key areas of the social brain, such as the TPs.
Supplementary Material
Supplementary material can be found at: http://www.cercor.
oxfordjournals.org/
Funding
PT would like to thank Fundacxa
˜o para a Cie
ˆncia e Tecnologia
(Portugal) for their support in the form of a fellowship
(PRAXISXXI/BD/21369/1999).
Notes
This study was conducted at the University of Cambridge Wolfson Brain
Imaging Centre (WBIC). Our thanks go to the staff of the WBIC for
assistance with fMRI data collection. Conflict of Interest: None declared.
Address correspondence to Paula Tavares, Instituto de Biofı
´sica e
Engenharia Biome
´dica, Faculdade de Cie
ˆncias da Universidade de
Lisboa, Campo Grande, 1749-016 Lisboa, Portugal. Email: ptavares@
fc.ul.pt.
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