Social concepts are represented in the superior
anterior temporal cortex
Roland Zahn†‡, Jorge Moll†, Frank Krueger†, Edward D. Huey†, Griselda Garrido§, and Jordan Grafman†¶
†National Institutes of Health, National Institute of Neurological Disorders and Stroke, Cognitive Neuroscience Section, Bethesda, MD 20892-1440;
‡Department of Psychiatry and Psychotherapy, Albert Ludwigs University of Freiburg, 79104 Freiburg, Germany; and§Instituto Israelita de
Ensino e Pesquisa, Hospital Albert Einstein, 05651-901, Sao Paulo, Brazil
Edited by James L. McClelland, Carnegie Mellon University, Pittsburgh, PA, and approved February 18, 2007 (received for review August 15, 2006)
Social concepts such as ‘‘tactless’’ or ‘‘honorable’’ enable us to
describe our own as well as others’ social behaviors. The prevailing
view is that this abstract social semantic knowledge is mainly
subserved by the same medial prefrontal regions that are consid-
ered essential for mental state attribution and self-reflection.
Nevertheless, neurodegeneration of the anterior temporal cortex
typically leads to impairments of social behavior as well as general
conceptual knowledge. By using functional MRI, we demonstrate
that the anterior temporal lobe represents abstract social semantic
knowledge in agreement with this patient evidence. The bilateral
superior anterior temporal lobes (Brodmann’s area 38) are selec-
tively activated when participants judge the meaning relatedness
of social concepts (e.g., honor–brave) as compared with concepts
describing general animal functions (e.g., nutritious–useful). Re-
markably, only activity in the superior anterior temporal cortex,
but not the medial prefrontal cortex, correlates with the richness
of detail with which social concepts describe social behavior.
Furthermore, this anterior temporal lobe activation is independent
of emotional valence, whereas medial prefrontal regions show
enhanced activation for positive social concepts. Our results dem-
onstrate that the superior anterior temporal cortex plays a key role
in social cognition by providing abstract conceptual knowledge of
social behaviors. We further speculate that these abstract concep-
tual representations can be associated with different contexts of
social actions and emotions through integration with frontolimbic
circuits to enable flexible evaluations of social behavior.
functional MRI ? semantics ? social cognition ? temporal lobe ? frontal lobe
of a social or moral concept, such as honor, changes with cultural
context, we are nonetheless able to understand its core meaning
in a 16th-century play. Here, we explore the neuroanatomical
basis of this remarkably stable social domain of conceptual
knowledge. One hypothesis is that such abstract social semantic
knowledge necessary to describe psychological characteristics is
mainly subserved by the same medial prefrontal regions (1–3)
that are essential for attributing mental states (theory of mind)
and self-reflection (4–7). This study provides evidence for an
alternative view, which predicts separable abstract representa-
tions of social concepts (e.g., ‘‘ambitious,’’ ‘‘polite,’’ ‘‘tactless,’’
and ‘‘stingy’’) in the anterior temporal lobe.
Most of what we know about the neural organization of
conceptual knowledge is based on studies of names for living and
nonliving objects. These studies indicate that different concep-
tual domains [animals, fruits/vegetables, tools (8)] and features
is characterized by both domain-specific supramodal (verbal and
nonverbal) and unimodal feature-specific regions (8, 10). Pa-
tients with neurodegeneration of the anterior temporal lobe (11)
demonstrate not only verbal but also nonverbal semantic im-
pairments, which lead to the conclusion that this region repre-
sents supramodal semantic knowledge (12–15).
hat is honor?’’ asks Shakespeare’s Falstaff (The First Part
of King Henry the Fourth 5.1.133). Although the meaning
Contrary to concepts that are expressed by names for living
things (e.g., ‘‘dog’’), social concepts (e.g., ‘‘loyal’’) are names for
social behavior or properties of living things [e.g., ‘‘acting in a
loyal way’’ or ‘‘being loyal’’ (16)]. Therefore, as the most suitable
class of concepts for comparison with social concepts, we chose
concepts that are names for animal behavior or properties (e.g.,
‘‘being nutritious,’’ ‘‘useful,’’ ‘‘trainable,’’ and ‘‘healthy’’).
Social concepts can apply to humans as well as other animals
(e.g., ‘‘a loyal dog’’), and they rely on abstract functional, more
than on sensory, knowledge (16). In some instances, we associate
social concepts with socially relevant sensory cues, such as
biological motion, which depend on posterior temporal regions
(17), and social judgments based on such sensory cues also
involve prefrontal areas (18). Grasping the meaning of social
concepts, such as ‘‘honorable’’ or ‘‘tactless,’’ however, most
prominently requires abstract functional (i.e., nonsensory)
knowledge, which entails descriptions of social behavior rather
than sensory detail (16). For example, this knowledge enables us
to understand a person’s social behavior as tactless even when
sensory cues (e.g., a friendly facial expression and body posture)
would indicate otherwise. The neuroanatomical basis of this
abstract social conceptual knowledge is elusive.
There have been several neuroimaging studies using socially
relevant words as stimuli. Most studies contrasted either a social
and nonsocial task condition (3) or two different social condi-
tions [e.g., self- vs. nonself-related (5, 19–23)] containing the
same set of words, thereby subtracting activations evoked by the
words and their respective semantic representations. It is well
established that the automatic activation of semantic represen-
tations can be elicited by the mere presentation of words even
when the task to be performed on the words does not require
explicit semantic knowledge [e.g., word/nonword decision (24)].
Two studies by Mitchell et al. (1, 2) used a different approach by
contrasting social and nonsocial words. They first looked at
containing fruit or clothing names (1). The authors concluded
that the medial prefrontal cortex represents social semantic
information about characteristics of persons and confined the
role of the detected left anterior temporal cortex activation to
sensory identification of socially relevant stimuli. It is not clear,
however, whether activity was related to the uniqueness of first
Author contributions: R.Z. and J.G. designed research; R.Z. and F.K. performed research;
G.G. contributed new reagents/analytic tools; R.Z., J.M., F.K., and G.G. analyzed data; and
R.Z., J.M., E.D.H., and J.G. wrote the paper.
The authors declare no conflict of interest.
This article is a PNAS Direct Submission.
Abbreviations: BA, Brodmann’s area; fMRI, functional MRI; HRF, hemodynamic response
function; ROI, region of interest.
Data deposition: The neuroimaging data have been deposited with the fMRI Data Center,
www.fmrdc.org (accession no. 2-2007-12248).
¶To whom correspondence should be addressed. E-mail: email@example.com.
This article contains supporting information online at www.pnas.org/cgi/content/full/
© 2007 by The National Academy of Sciences of the USA
April 10, 2007 ?
vol. 104 ?
names in comparison to nonunique object names, as proposed by
Grabowski et al. (25). In a subsequent study, Mitchell et al. (2)
reproduced medial prefrontal, but not anterior temporal acti-
vations for person-descriptive words when compared with body
part words; this finding supported their earlier conclusion that
semantic knowledge of psychological states is bound to the
medial prefrontal cortex. We argue, however, that none of the
above studies critically tested that prediction because when
socially relevant stimuli are categorically compared with less
by social cognitive processes other than domain-specific seman-
tic processing, such as mental state attribution.
Here, we use functional MRI (fMRI) to record brain activity
when participants make judgments about the meaning related-
ness of social concepts (e.g., honor–brave or tactless–impolite)
as compared with other animal function concepts (e.g.,
nutritious–useful, presented as word pairs), a task that requires
access to detailed conceptual knowledge. Animal function con-
cepts describe behaviors related to animal use and biological
function and can in principle apply to humans as well. Using this
categorical comparison, we are able to reproduce the network of
social cognition regions found in previous studies, including
medial prefrontal and anterior temporal cortex when comparing
socially relevant with less socially relevant words. However, as
the critical test of which of these regions is representing abstract
conceptual information, we applied two key measures of con-
ceptual knowledge as parameters in a separate regression anal-
ysis: (i) descriptiveness, and (ii) meaning relatedness of pre-
sented word pairs. By using these regression analyses, we provide
evidence that a superior sector of the anterior temporal lobe is
the only region in the brain to be selectively associated with
conceptual knowledge of social behaviors.
Descriptiveness of concepts is the richness of conceptual
knowledge detail (26). The more general a concept, the less
detail of description it conveys. A general concept (e.g., un-
friendly) is less descriptive than a more specific one (e.g.,
tactless) (27). More descriptive concepts require more detailed
conceptual knowledge and are thus predicted to increase neural
activity in conceptual brain regions. Because we specifically
investigated the detail of behavior descriptions (not sensory
detail) and adjusted for the effects of word imageability (highly
correlated with concreteness; Pearson r ? 0.91, P ? 0.0001), we
were able to determine whether regions code for abstract
functional (i.e., nonsensory) knowledge. Meaning relatedness is
an established measure of the degree to which two concepts are
similar in meaning, and the organization of conceptual knowl-
edge critically depends upon such information (14, 28).
Although there is no direct evidence on the anatomical locus
of abstract social concept knowledge, indirect evidence suggests
that anterior temporal lobe regions might be involved. Some
patients with penetrating head injuries to the temporal lobes
incurred during World War I were selectively unable to give
examples of social behaviors to define concepts that describe
character attributes (29). In addition, patients with anterior
temporal lobe neurodegeneration not only exhibit gross concep-
tual impairments (12), but also display changes in social behavior
lobe activations in such diverse tasks as moral cognition (31),
understanding others’ mental states (4) or emotions (32), when
tasks used persons’ first names (1) or famous faces and names
(33), as well as retrieval of famous name–face associations (34).
Despite this indirect evidence for the importance of the anterior
temporal lobe in social cognition, its exact contribution remains
obscure because the common cognitive component across these
different tasks has not been identified. In a recent model (31),
we therefore hypothesized that specific anterior temporal lobe
regions represent conceptual knowledge of social behaviors,
which would be an essential underlying cognitive component
shared by these social cognition tasks. A critical test of this
prediction is whether there are distinct anterior temporal lobe
regions selective for social concepts, and whether activity in
these regions correlates with (i) the degree of detail with which
concepts describe social behavior and (ii) the relatedness in
meaning of two concepts in a word pair.
The number of concept pairs judged as being related or unre-
lated in meaning during fMRI was equal across conditions [see
quickly to related word pairs in all conditions. Overall response
time was significantly slower for social than animal function
concepts (see SI Fig. 3). Therefore, we tested the effects of
response time on the observed brain activations for social
concepts. There was no association of increased response time
with temporal lobe activation for social concepts, ruling out
effects of task difficulty.
The categorical subtraction analysis for social compared with
animal function concepts revealed a cluster of activation (sig-
nificant at P ? 0.05, family-wise error-corrected for multiple
comparisons; Table 1) within right superior temporal [Brodma-
nn’s area (BA)38] and lateral orbitofrontal/inferior frontal cor-
tex (BA47/45). In this categorical comparison, additional regions
implicated in social cognition (4, 7, 31) were also activated,
including the dorsomedial prefrontal cortex (BA8) and the left
posterior fusiform activations comparable with those reported in
studies of sensory social semantics and face recognition were
detected (17, 35, 36). Although temporal lobe activation did not
differ between positive and negative social concepts, positive
social concepts engaged a more anterior sector of the medial
prefrontal cortex (BA10/32; see Fig. 1 and SI Fig. 4), indicating
that anterior temporal lobe activation is independent of emo-
The anterior temporal region of interest (ROI) analysis com-
paring social vs. animal concepts revealed bilateral activation of
the superior anterior temporal lobe (BA38) and less pronounced
signal increases in the anterior middle temporal cortex (BA21;
Fig. 2a). The reverse comparison (animal vs. social concepts)
revealed no significant effects. Animal function concepts com-
pared with fixation engaged anterior middle temporal cortex
(BA21; Fig. 2b). The same area was activated by social concepts
vs. fixation, indicating that the anterior middle temporal areas
are shared by both classes of concepts.
The prior demonstration of regional activity for social con-
cepts by categorical subtraction of activation for less socially
relevant concepts, however, does not reveal whether activity
specific to social concepts is elicited by conceptual knowledge of
social behaviors or by other social cognitive processes (e.g.,
attribution of mental states or self-reflection). The critical test of
this hypothesis was an independent analysis that searched for
regions in which activity was not only higher for social than for
animal concepts, but was independently correlated with the
degree of descriptiveness of social behavior and with meaning
relatedness (conjunction analysis for these three effects). The
right superior anterior temporal region (BA38) was the only
region surviving this conjunction analysis on a whole-brain basis.
ROI analysis revealed additional activation in homologous left-
hemispheric cortex (BA38; Fig. 2c). In remarkably close agree-
ment with our predictions, activity in the right superior anterior
temporal region (BA38) showed a significantly stronger corre-
lation with descriptiveness of social behavior than activity in
2). There was no correlation of neural activity with descriptive-
ness of social behavior in the nonspecific right anterior middle
temporal region (BA21; data not shown). Furthermore, indi-
vidual case analyses confirmed consistent anatomical separation
Zahn et al.PNAS ?
April 10, 2007 ?
vol. 104 ?
no. 15 ?
of activations related to social concepts (superior temporal) and
animal function concepts (middle temporal) within the anterior
temporal cortex, particularly within the right hemisphere (Fig.
2e; right hemisphere, P ? 0.0001; left hemisphere, P ? 0.001;
Fisher’s two-sided exact test; see SI Methods).
In summary, social concepts consistently activated a selective
superior anterior temporal lobe region (BA38), and both animal
and social concepts shared a nonspecific anterior middle tem-
(orbitofrontal, medial prefrontal cortex, and temporoparietal
junction) known to be crucial for social cognition (1, 4, 31). Only
activity in the superior anterior temporal cortex, however,
robustly correlated with the richness of detail with which social
concepts describe social behavior. This finding corroborates our
prediction that specific anterior temporal lobe regions represent
conceptual knowledge of social behaviors (31). Activity in the
superior temporal pole (BA38) agrees with selective connections
region for social cognition (4, 6, 7, 31, 38). On the contrary,
middle and inferior temporal lobes are primarily connected to
the orbital network, which integrates information from sensory
systems and rewards (37). The exact anatomical location of our
superior anterior temporal lobe region according to recent
human anatomical studies (39) is at the posterior border of the
temporal pole (BA38) reaching into the anterior superior tem-
poral gyrus (BA22), which is highly connected to the superior
temporal pole [BA38 (37)].
The independence of temporal lobe activation from emotional
valence is in line with our prediction that abstract social con-
ceptual representations in the anterior temporal lobe are
valence-independent and can be dynamically associated with
different emotionally relevant contexts encoded in frontolimbic
circuits (31). This independence from emotional valence may
explain why neuroimaging investigations of emotional word
connotations did not find consistent anterior temporal lobe
activation (40, 41).
These results cannot be attributed to confounding differences
between social and animal function words because we meticu-
lously controlled for all relevant psycholinguistic differences
(including lexical frequency and familiarity) in the categorical
subtractions and confirmed our results by independent para-
metric regression analyses. Higher frequency of adjectives in the
social concept condition cannot explain effects within the ante-
rior temporal lobe because lesions in this region lead to con-
ceptual impairment irrespective of word class (12, 14). Also,
there was no association of increased response time with tem-
Our results are in agreement with the central role of the
anterior temporal lobes for representing abstract conceptual
knowledge (12–15, 28, 42), concepts denoted by composite
expressions (43–45), and the importance of the right temporal
domains (e.g., tools, animals, and faces) were demonstrated in
modality-specific posterior temporal regions (35, 36, 46). This
study demonstrates that specialized subregions for different
conceptual domains also exist within the anterior temporal lobe.
It has been argued that subdivisions for different object cate-
gories in the posterior temporal cortex do not necessarily reflect
Table 1. Social vs. animal function concepts
L and R
Lateral orbitofrontal/anterior temporal cluster**
Anterior superior temporal gyrus**
Lateral orbitofrontal/inferior frontal gyrus**
Dorsomedial prefrontal cortex*
Middle frontal gyrus*
Inferior frontal gyrus*
Lateral inferior temporal gyrus*
Lateral fusiform gyrus*
Medial occipital gyrus*
Social concepts (sum of all effects of interest: i ? ii ? iii) vs. animal function concepts (sum of all effects of interest: i ? ii ? iii) inclusively masked with social
concepts (all effects of interest: i ? ii ? iii) vs. fixation. Effects of interest: (i) partial regression effect of event-related hemodynamic response function (HRF),
voxel-level threshold (minimum cluster size ? 10 voxels) in the whole-brain analysis are reported. Subclusters ?8 mm apart are italic.*, Areas surviving
stringent correction (family-wise error) or those predicted by an a priori anatomical hypothesis are discussed in the text (see SI Methods). MNI, Montreal
Neurological Institute Standard Brain coordinates. L, left; R, right.
coordinate of the displayed anterior medial prefrontal (BA10/32) region is
?12, 54, 18, Z ? 3.64. Regions not displayed here, which were additionally
detected: left lateral orbitofrontal cortex (BA47/11), ?42, 30, ?15, Z ? 4.44;
dorsal anterior cingulate (BA24), ?3, 0, 24, Z ? 3.51. No temporal lobe
differences emerged on a whole-brain and anterior temporal ROI basis, also
at P ? 0.05 uncorrected. See also SI Fig. 4 for parameter estimates for positive
and negative social concepts.
Positive vs. negative social concepts. Whole-brain analysis at a
www.pnas.org?cgi?doi?10.1073?pnas.0607061104Zahn et al.
modular specialization for a given category, and that for most
categories of objects category effects can be explained by a
continuous topographical representation of attributes (i.e., fea-
be applied to our finding of topographic differences within the
anterior temporal lobe for conceptual representations of social
and general animal behavior. They could be equally well ex-
plained by domain-specific as well as conceptual similarity-based
topographic organization of underlying cortical representations.
Further studies are necessary to address the exact role of more
inferior anterior temporal lobe regions that were reported in
addition to superior anterior temporal cortex in neuroimaging
studies of social cognition (31–33) and in what way famous face
naming relates to conceptual knowledge of social behaviors
studied here. Famous face and proper name processing were
used as measures of person-specific semantic knowledge in
patient lesion and functional imaging studies (33, 47–52). Neu-
ropsychological cases exhibited extensive lesions of the left (49)
or right (50) anterior temporal lobe, with relative sparing of the
superior sector (51). The specificity of famous face-naming
impairment after temporal pole lesions has been questioned by
findings of equal impairments for other unique entities [e.g.,
temporal pole is involved in lexical retrieval of unique object
names. Other authors have stressed the distributed nature of
lesions leading to impairments of retrieving proper names (53).
In any case, our findings cannot be attributed to lexical (i.e.,
word-form) retrieval of proper names because we used non-
unique concepts as stimuli, and our task did not involve lexical
One possible relation between more sensory semantic infor-
mation related to persons (as measured by famous face-naming
tasks) and the abstract conceptual knowledge investigated here
was proposed by Burton and colleagues (54). They suggest
dissociable cognitive representations for multimodal informa-
tion necessary to identify a person (e.g., face image, proper
name) and more abstract information about a person (e.g.,
occupation). The anatomical locus of both systems has not been
by social concepts vs. fixation] and less strongly anterior middle temporal cortex (BA21; right ? left; 57, ?3, ?21, Z ? 2.17). (b) Animal function concepts vs.
fixation engaged the same anterior middle temporal area (BA21, right ? left; 57, ?3, ?21; Z ? 3.62, section at x ? 60). This region was also activated by social
concepts vs. fixation (BA21, right ? left; 54, ?3, ?21; Z ? 4.83, corrected P ? 0.001 by using the total 914 voxel volume of the bilateral anterior temporal ROI
mask; data not shown). (Note that Z scores for clusters with P values that are ? 0.05 and were small-volume-corrected by using a 12-mm sphere around the peak
voxel in the whole-brain analysis are bold.) (c) Regions in which activity was higher for social than for animal concepts and that were independently correlated
with descriptiveness of social behavior and meaning relatedness: conjunction null analysis for all of these three effects (ROI for display, sagittal at x ? 57, and
axial views at z ? ?6) revealed a selective right superior anterior temporal region on a whole-brain basis (BA38; right vs. left; 51, 15, ?12; Z ? 2.9). (d) Parameter
estimates and SEs for descriptiveness of social behavior, meaning relatedness, and domain-specificity (social vs. animal) at right superior anterior temporal,
aTL, n ? 0; right middle aTL, n ? 22; other, n ? 4). Fisher’s two-sided exact tests showed that type of contrast (social vs. animal or animal vs. fixation) influenced
how many subjects had activations located within the superior aTL or middle aTL, respectively (right, P ? 0.0001; left, P ? 0.001; see SI Methods).
Zahn et al. PNAS ?
April 10, 2007 ?
vol. 104 ?
no. 15 ?
identified yet. Following this scheme, the abstract conceptual
representations detected in our study would be part of the latter
necessary nor sufficient to identify a person. Formal testing of
more abstract social semantic knowledge is usually not reported
in cases describing famous face-naming impairments in anterior
temporal lobe lesions. In a patient with bilateral inferior tem-
poral pole lesions, however, normal spontaneous use of abstract
knowledge about social values was described, which contrasted
with severe impairments on famous face naming (55). Together
with the relative sparing of the superior anterior temporal lobe
in another case of famous face-naming impairment (51), this
evidence points to a possible inferior–superior gradient for
multisensory versus abstract person-specific knowledge. The
exact topographic relation of both types of person-related se-
mantic systems needs to be addressed in future studies.
Previous functional neuroimaging studies comparing abstract
with concrete concepts have, among other areas, reported
activations in comparable superior temporal pole regions, as the
one identified here (38, 56–58). In principle, one could derive
two different conclusions from this anatomical convergence: (i)
Our findings can be explained on the basis of the abstractness of
representations, and (ii) the reported superior anterior temporal
activations in studies on abstract concepts are due to the
incidental use of socially relevant concepts as stimuli. We argue
that the latter conclusion is strongly supported. Because in all
our analyses partial effects of social concepts are adjusted for
effects of imageability, which is highly correlated with concrete-
ness, differences in abstractness/concreteness cannot explain the
differences in activations between social and animal function
concepts. Furthermore, our regression analyses demonstrated
that the degree of activation in the superior anterior temporal
region was increased with the degree to which concepts de-
scribed social behavior. This effect was again independent of
differences in imageability because it was adjusted for in the
multiple-regression model. In summary, our experimental de-
sign carefully rules out a confounding effect of general abstract-
ness to explain our data. Thus, the comparable activation sites
within the superior anterior temporal lobe reported in previous
studies on abstract concepts can be reinterpreted as due to the
social relevance of used concepts. This conclusion is corrobo-
rated by looking at provided listings of stimuli, which, to a large
proportion, contained socially relevant concepts (38, 56, 58). For
example, in the study by Sabsevitz et al. (38), concepts such as
‘‘courage’’ and ‘‘disgrace’’ were mixed with less socially relevant
abstract concepts such as ‘‘lesson’’ and ‘‘riddle.’’ The degree of
social relevance, however, was not controlled.
Taken together, our findings indicate that a superior sector of
the anterior temporal cortex plays a key role in social cognition
by representing abstract conceptual knowledge of social behav-
iors, and that these representations are independent of emo-
tional valence. Furthermore, we demonstrated that, although
medial prefrontal cortex is involved in processing socially rele-
vant information, it does not represent abstract social semantic
knowledge. We further speculate that abstract conceptual rep-
resentations in the anterior temporal lobe can be associated with
different contexts of social actions and emotions through inte-
gration with frontolimbic circuits to enable flexible evaluations
of social behavior (31).
Materials and Methods
Subjects. Twenty-six healthy participants (13 men; age mean ?
29.4 ? 9.0; years of education mean ? 17.5 ? 2.5) took part in
to be excluded before the statistical analysis (n ? 2, no response
times recorded; n ? 1, MR-scanner failure; n ? 1, head motion;
n ? 1, temporal lobe signal loss). All participants were right-
handed (59) and native English speakers. All participants un-
derwent a neurological examination by a board-certified neu-
rologist and a clinical MRI during the previous 12 months, had
normal or corrected-to-normal vision, had no history of psychi-
atric or neurological disorders, and were not taking centrally
active medications. Informed consent was obtained according to
procedures approved by the National Institute of Neurological
compensated for their participation according to the National
Institute of Neurological Disorders and Stroke’s standards.
Measures of self-esteem and trait affective style were collected
mean ? 36.1 ? 3.7; PANAS (61) positive affect score, mean ?
36.3 ? 6.8; negative affect score, mean ? 13.6 ? 3.7].
fMRI Paradigm. Participants decided whether written word pairs
were related in meaning by pressing one of two response keys.
Three different types of word pairs or a visual fixation pattern
were presented: (i) animal function concepts [used with kind
permission of the authors of ref. 62; e.g., nutritious–useful, n ?
authors of ref. 27; e.g., honor–brave, n ? 75], and (iii) negative
social concepts (27) (e.g., tactless–impolite, n ? 75; see SI Fig.
5 and SI Methods). In two independent prestudies, we asked
participants to rate the degree of detail with which each word
described social behavior (social concepts) or animal behavior
(animal function concepts) and how related in meaning both
words within a pair were (i.e., meaning relatedness; see SI
Methods). Relevant psycholinguistic variables were matched
across conditions (word familiarity, frequency, difference in
category breadth and social desirability within word pair, asso-
ciativity, and meaning relatedness; see SI Methods).
Image Acquisition. Echo-planar T2*-weighted images with blood
oxygenation-level-dependent contrast were acquired (311 vol-
umes per run) on a 3 Tesla General Electric scanner (GE
high-order manual shimming to temporal and ventral frontal
lobes [3-mm slice thickness, 64 ? 64 matrix, 37 slices, repetiton
time ? 2.3 sec, field of view: 220 ? 220, parallel to the anterior
to posterior commissural line, whole-brain coverage (not cere-
bellum)]. The first five volumes were discarded to allow for T1
equilibration effects. The combination of high-field MRI, thin-
ner slices, and high-order manual shimming optimized the signal
in anterior temporal and ventral frontal lobes. All participants
had full coverage of the anterior temporal lobes upon inspection
of normalized images (see SI Fig. 6). One subject was excluded
before statistical analysis because of signal dropout within
predefined critical regions (anterior temporal lobe, BA38/22,
BA21, BA20; ventromedial prefrontal, BA11, BA25, BA24,
BA32; ventrolateral prefrontal, BA11/47; and frontopolar cor-
tex, BA10). In addition, high-resolution (?1 mm3) T1-weighted
3D magnetization-prepared rapid acquisition gradient echo
structural images were collected (1-mm slice thickness, 128
slices, matrix: 224 ? 224, field of view: 220 ? 222). Head motion
was restricted by using vacuum bags fitted to the participant’s
Image Analysis. Imaging data were analyzed by using statistical
spm5) and a general linear model (63). The mean degree of
descriptiveness and meaning relatedness were modeled as para-
metric predictors of interest for each stimulus condition. Im-
ageability, number of syllables, and social desirability for social
concepts were modeled as covariates of no interest. A separate
model was set up including all above variables, with the addition
of response time for each stimulus condition to test whether
domain-specific effects were due to response time effects.
www.pnas.org?cgi?doi?10.1073?pnas.0607061104 Zahn et al.
Categorical contrasts were formed by summing up all effects Download full-text
of interest per condition: (i) condition-specific hemodynamic
response function (HRF), (ii) effect of behavior descriptiveness
convolved with HRF, and (iii) effect of meaning relatedness
convolved with HRF. Reported statistics were performed on the
second level by using a random-effects model.
specific effect (social vs. animal), descriptiveness of social be-
havior, and meaning relatedness of social concepts were detect-
able in conjunction (conjunction null analysis), we set up a
separate factorial model at the second level. The factorial model
included the following contrasts: (i) condition-specific HRF
compared with fixation HRF, (ii) effect of behavior descriptive-
ness, and (iii) meaning relatedness of each condition convolved
with the respective HRF.
We inclusively masked each reported categorical contrast
(e.g., social vs. animal) with a contrast against the low-level
control condition (e.g., social vs. fixation; see SI Methods).
Whole-brain analyses were based on an uncorrected voxel level
of P ? 0.005 (10 voxels minimum cluster size) in a priori
predicted regions known from the social and semantic neuro-
science literature (see SI Methods). Results from anterior tem-
poral lobe ROI analyses were displayed on an uncorrected voxel
level of P ? 0.05 (10 voxels minimum cluster size) to show the
extent of activation and corroborate regional specificity. All
reported coordinates are in Montreal Neurological Institute
We thank Katherine O’Leary for help with data acquisition, John Bartko
for statistical advice, Eric Wassermann for performing neurological
exams, and Kris Knutson and several statistical parametric mapping
experts from the discussion list for imaging analysis advice. This study
was supported by German Academy of Natural Scientists Leopoldina
Grant BMBF-LPD 9901/8–122 (to R.Z.); the National Institute of
Neurological Disorders and Stroke Intramural Research Program (to
J.G.); and Fundac ¸a ˜o de Amparo a ´ Pesquisa do Estando de Sa ˜o Paulo
Grant 03/11794-6 (to G.G.).
1. Mitchell JP, Heatherton TF, Macrae CN (2002) Proc Natl Acad Sci USA
2. Mitchell JP, Banaji MR, Macrae CN (2005) NeuroImage 28:757–762.
3. Mason MF, Banfield JF, Macrae CN (2004) Cereb Cortex 14:209–214.
4. Blakemore SJ, Winston J, Frith U (2004) Trends Cogn Sci 8:216–222.
5. Northoff G, Heinzel A, de Greck M, Bermpohl F, Dobrowolny H, Panksepp
J (2006) Neuroimage 31:440–457.
6. Gusnard D, Akbudak E, Shulman G, Raichle M (2001) Proc Natl Acad Sci USA
7. Amodio DM, Frith CD (2006) Nat Rev Neurosci 7:268–277.
8. Caramazza A, Mahon BZ (2003) Trends Cogn Sci 7:354–361.
9. McCarthy RA, Warrington EK (1988) Nature 334:428–430.
10. Zahn R, Garrard P, Talazko J, Gondan M, Bubrowski P, Juengling F, Slawik
H, Dykierek P, Koester B, Huell M (2006) J Cogn Neurosci 18:2138–2151.
11. Davies RR, Hodges JR, Kril JJ, Patterson K, Halliday GM, Xuereb JH (2005)
12. Bozeat S, Lambon Ralph MA, Patterson K, Garrard P, Hodges JR (2000)
13. Garrard P, Carroll E (2006) Brain 129:1152–1163.
14. Rogers TT, Lambon Ralph MA, Garrard P, Bozeat S, McClelland JL, Hodges
JR, Patterson K (2004) Psychol Rev 111:205–235.
15. Jefferies E, Lambon Ralph MA (2006) Brain 129:2132–2147.
16. Hampson S, John O, Goldberg L (1986) J Pers Soc Psychol 51:37–54.
17. Martin A, Weisberg J (2003) Cogn Neuropsychol 20:575–587.
18. Heberlein AS, Saxe RR (2005) NeuroImage 28:770–777.
19. Kircher TT, Senior C, Phillips ML, Benson PJ, Bullmore ET, Brammer M,
Simmons A, Williams SC, Bartels M, David AS (2000) Brain Res Cogn Brain
21. Fossati P, Hevenor SJ, Graham SJ, Grady C, Keightley ML, Craik F, Mayberg
H (2003) Am J Psychiatry 160:1938–1945.
22. Johnson S, Baxter L, Wilder L, Pipe J, Heiserman J, Prigatano G (2002) Brain
23. Schmitz T, Kawahara Baccus T, Johnson S (2004) NeuroImage 22:941–947.
24. Mummery CJ, Shallice T, Price CJ (1999) NeuroImage 9:516–525.
25. Grabowski TJ, Damasio H, Tranel D, Ponto LL, Hichwa RD, Damasio AR
(2001) Hum Brain Mapp 13:199–212.
26. Rosch E (1975) J Exper Psychol Gen 104:192–233.
27. John OP, Hampson SE, Goldberg LR (1991) J Pers Soc Psychol 60:348–361.
28. McClelland JL, Rogers TT (2003) Nat Rev Neurosci 4:310–322.
29. Von Kleist K (1922) in Geistes und Nervenkrankheiten, ed Bonhoeffer K
(Verlag von Johann Ambrosius Barth, Leipzig).
30. Liu W, Miller BL, Kramer JH, Rankin K, Wyss-Coray C, Gearhart R,
Phengrasamy L, Weiner M, Rosen HJ (2004) Neurology 62:742–748.
31. Moll J, Zahn R, de Oliveira-Souza R, Krueger F, Grafman J (2005) Nat Rev
32. Wicker B, Perrett DI, Baron-Cohen S, Decety J (2003) Neuropsychologia
33. Gorno-Tempini ML, Price CJ, Josephs O, Vandenberghe R, Cappa SF, Kapur
N, Frackowiak RS (1998) Brain 121:2103–2118.
34. Kikyo H, Miyashita Y (2004) NeuroImage 23:1348–1357.
35. Chao LL, Haxby JV, Martin A (1999) Nat Neurosci 2:913–919.
36. Kanwisher N, McDermott J, Chun MM (1997) J Neurosci 17:4302–4311.
37. Kondo H, Saleem KS, Price JL (2003) J Comp Neurol 465:499–523.
38. Sabsevitz DS, Medler DA, Seidenberg M, Binder JR (2005) NeuroImage
39. Insausti R, Juottonen K, Soininen H, Insausti AM, Partanen K, Vainio P,
Laakso MP, Pitkanen A (1998) AJNR Am J Neuroradiol 19:659–671.
40. Cato MA, Crosson B, Gokcay D, Soltysik D, Wierenga C, Gopinath K, Himes
42. Spitsyna G, Warren JE, Scott SK, Turkheimer FE, Wise RJ (2006) J Neurosci
43. Xu J, Kemeny S, Park G, Frattali C, Braun A (2005) NeuroImage 25:1002–
44. Vandenberghe R, Nobre AC, Price CJ (2002) J Cogn Neurosci 14:550–560.
45. Sharp DJ, Scott SK, Wise RJ (2004) Ann Neurol 56:836–846.
46. Ishai A, Ungerleider LG, Martin A, Schouten JL, Haxby JV (1999) Proc Natl
Acad Sci USA 96:9379–9384.
47. Damasio H, Grabowski T, Tranel D, Hichwa R, Damasio A (1996) Nature
48. Tranel D (2006) Neuropsychology 20:1–10.
49. Giovanello KS, Alexander M, Verfaellie M (2003) Neurocase 9:15–26.
50. Thompson SA, Graham KS, Williams G, Patterson K, Kapur N, Hodges JR
(2004) Neuropsychologia 42:359–370.
51. Evans JJ, Heggs AJ, Antoun N, Hodges JR (1995) Brain 118:1–13.
52. Ellis AW, Young AW, Critchley EM (1989) Brain 112:1469–1483.
53. Semenza C, Mondini S, Zettin M (1995) Neurocase 1:183–188.
54. Burton AM, Bruce V, Johnston RA (1990) Br J Psychol 81:361–380.
55. Sirigu A, Duhamel JR, Poncet M (1991) Brain 114:2555–2573.
56. Perani D, Cappa SF, Schnur T, Tettamanti M, Collina S, Rosa MM, Fazio F
(1999) Brain 122:2337–2344.
57. Kiehl KA, Liddle PF, Smith AM, Mendrek A, Forster BB, Hare RD (1999)
Hum Brain Mapp 7:225–233.
58. Noppeney U, Price CJ (2004) NeuroImage 22:164–170.
59. Oldfield RC (1971) Neuropsychologia 9:97–113.
60. Rosenberg M (1989) Society and the Adolescent Self-Image (Wesley Univ Press,
61. Watson D, Clark LA, Tellegen A (1988) J Pers Soc Psychol 54:1063–1070.
62. McRae K, Cree GS, Seidenberg MS, McNorgan C (2005) Behav Res Methods
63. Friston KJ, Frith CD, Turner R, Frackowiak RS (1995) NeuroImage 2:157–165.
Zahn et al. PNAS ?
April 10, 2007 ?
vol. 104 ?
no. 15 ?