Neuropsychologia 38 (2000) 11±21
www.elsevier.com/locate/neuropsychologia
Reading the mind in cartoons and stories: an fMRI study of
`theory of mind' in verbal and nonverbal tasks
H.L. Gallagher a, F. Happe b, N. Brunswick a, P.C. Fletcher a, U. Frith c, C.D. Frith a,*
a
Wellcome Department of Cognitive Neurology, Institute of Neurology, University College London, 12 Queen Square, London WC1N 3BG, UK
b
Social, Genetic and Developmental Psychiatry Research Centre, Institute of Psychiatry, Denmark Hill, London, UK
c
Institute of Cognitive Neuroscience & Department of Psychology, University College London, London, UK
Received 4 November 1998; received in revised form 31 March 1999; accepted 12 April 1999
Abstract
Previous functional imaging studies have explored the brain regions activated by tasks requiring `theory of mind'Ðthe
attribution of mental states. Tasks used have been primarily verbal, and it has been unclear to what extent di€erent results have
re¯ected di€erent tasks, scanning techniques, or genuinely distinct regions of activation. Here we report results from a functional
magnetic resonance imaging study (fMRI) involving two rather di€erent tasks both designed to tap theory of mind. Brain
activation during the theory of mind condition of a story task and a cartoon task showed considerable overlap, speci®cally in
the medial prefrontal cortex (paracingulate cortex). These results are discussed in relation to the cognitive mechanisms
underpinning our everyday ability to `mind-read'. # 1999 Elsevier Science Ltd. All rights reserved.
Keywords: Medial prefrontal cortex; Functional magnetic resonance imaging; Theory of mind
1. Introduction
Recent interest in the evolution, development, and
breakdown of social cognition (see, for example, chapters in Carruthers and Smith, [6]) has been re¯ected in
a number of functional imaging studies of this ability.
`Theory of mind', the ability to attribute independent
mental states to self and others in order to explain and
predict behaviour, has been suggested to arise from a
dedicated, domain-speci®c, and possibly modular cognitive mechanism [12,24]. This proposal gains particular support from studies of autism, a biologicallybased developmental disorder which appears to be
characterised by a selective impairment in theory of
mind [19]. Interest in the brain basis of normal theory
of mind, is ®red by the hope of better understanding
* Corresponding author. Tel.: +44-171-833-7472; fax: +44-171813-1420.
E-mail address: c.frith@®l.ion.ucl.ac.uk (C.D. Frith)
the neural systems which are abnormal in people with
autism, most of whom are unable to `mind-read'.
To date, there have been three published reports of
functional brain imaging studies of `theory of mind'.
Baron-Cohen et al. [2] used single photon emission
computerised tomography (SPECT) and a regions of
interest approach to isolate brain areas activated
during recognition of mental state terms in a word list.
They found that their normal adult volunteers showed
increased cerebral blood ¯ow during the mental state
recognition task in the right orbito-frontal cortex relative to the left frontal-polar region. Goel et al. [15]
used PET to scan adults engaged in a complex task in
which subjects had to model the knowledge and inference of another mind concerning the function of unfamiliar and familiar objects. They found widespread
activation associated with this task, including activation of left medial frontal lobe and left temporal
lobe. Fletcher et al. [10] also used PET, and scanned
volunteers asked to read and answer questions about
stories. Comparison of activation during `theory of
mind' stories (requiring mental state attribution) vs
0028-3932/99/$ - see front matter # 1999 Elsevier Science Ltd. All rights reserved.
PII: S 0 0 2 8 - 3 9 3 2 ( 9 9 ) 0 0 0 5 3 - 6
12
H.L. Gallagher et al. / Neuropsychologia 38 (2000) 11±21
Fig. 1. Examples of story comprehension passages.
H.L. Gallagher et al. / Neuropsychologia 38 (2000) 11±21
13
Fig. 1 (continued)
control `physical' stores revealed task-speci®c activation in the left medial prefrontal gyrus, as well as
increased activation of the posterior cingulate cortex.
Using the same technique with individuals with a form
of autism (Asperger Syndrome), Happe et al. [20]
found similar patterns of activation in all regions
except the medial frontal gyrus which had been linked
to theory of mind performance in the normal group.
The main aims of the current study were to investigate the neural correlates underlying theory of mind
by exploiting the superior spatial resolution of fMRI
compared to PET and to examine anatomical convergence between theory of mind tasks presented in di€erent modalities. The story comprehension task used by
Fletcher et al. [10] was modi®ed for compatibility with
fMRI to examine theory of mind in the verbal domain,
while captionless cartoons provided a visual equivalent. Previous behavioural studies have used both stories [14,17,18] and visual jokes [7,18] to investigate
theory of mind impairments in adults and children and
found these tasks to be good markers of mentalising
abilities. We hypothesised that activation in medial
prefrontal cortex would be associated with the attribution of mental states independent of modality. Our
®nal aim was to identify any modality speci®c regions
for theory of mind in the verbal and visual domains.
2. Methods
2.1. Subjects
Six right-handed volunteers with no neurological or
psychiatric history participated in this study. Of these,
®ve were male and one female, with a mean age of 30
yr (range 23±36 yr). This study was approved by the
Institute of Neurology Ethics Committee. Informed
written consent was obtained from all subjects prior to
scanning.
2.2. Tasks
All stimuli were displayed on a monitor and presented to the subject via a 458 angled mirror positioned above the head coil; this mirror was adjusted to
be within the subjects ®eld of vision without having to
tilt his/her head. A test image was presented on the
screen prior to scanning to ensure that the image was
in focus and the subject could comfortably read the
text.
2.2.1. Story comprehension task
Three types of verbal material were presented; these
were `theory of mind stories' (ToMS), `non-theory of
mind stories' (Non-ToMS) and `unlinked sentences'
(US). Fig. 1. shows examples of these three types of
material.
During each scanning epoch a passage of text was
presented on the screen for 21.6 s. The passage was
then replaced by a question relating to the text (presented for a further 11 s). Subjects were instructed to
read the passage silently and to answer internally the
question which followed. Each passage was preceded
by a relevant prompt to indicate either `theory of
mind', `unlinked sentences' or `non-theory of mind'.
This prompt was displayed for 1 s. The question was
14
H.L. Gallagher et al. / Neuropsychologia 38 (2000) 11±21
Fig. 2. Examples of theory of mind cartoons, non-theory of mind cartoons and jumbled pictures.
H.L. Gallagher et al. / Neuropsychologia 38 (2000) 11±21
15
Fig. 2 (continued)
replaced by the next prompt indicating the start of a
new epoch.
Following the scanning session each subject was
shown the same passages again and asked to give his/
her answer to each question, to provide a measure of
behavioural performance. Responses were scored 1 for
a correct answer and 0 for an incorrect answer. For
the theory of mind condition an answer was considered correct only if an appropriate mental state was
attributed to one or more characters. The maximum
score was 4 per condition which was reached by all
subjects, thus indicating full understanding.
The passages used in these tasks have been used in
two previous experiments by this group [10,19]. The
results from this study con®rmed that the tasks were
matched for diculty, as re¯ected in scores and reading times, with ToM stories taking if anything slightly
(but nonsigni®cantly) less time to read than the NonToMS.
2.2.2. Cartoon task
Three types of picture were presented in analogy to
the three types of text, `theory of mind cartoons'
(ToMC), `non-theory of mind cartoons' (Non-ToMC)
and `jumbled pictures' (JC). A cartoon was considered
to require theory of mind for its interpretation if attri-
bution of either false belief or ignorance to one or
more of the characters in the picture was vital for
comprehension. A cartoon was considered to be nontheory of mind if no mental state attribution was
needed to understand the meaning. The `jumbled pictures' were constructed from images of randomly positioned objects, animals and people, taken from
cartoons and children's colouring books. All images
were captionless. Fig. 2 shows examples of the three
stimulus types.
The cartoons were validated in a pilot study with
twenty naive normal subjects (age range 19±66 yr).
Subjects were instructed to look at each cartoon and
indicate to the experimenter as soon as they understood its meaning. Response time was recorded.
Subjects then gave a brief explanation of the cartoon's
meaning. The explanations were scored 1 for a correct
answer and 0 for an incorrect answer. As for the story
condition, explanations in the theory of mind condition were considered correct only if an appropriate
mental state was attributed to one or more characters.
The explanation was recorded along with the time
taken. In addition, participants were asked to rate,
from 1 to 5, how dicult and how funny they thought
the cartoon to be (1 extremely easy, 5 extremely dicult; 1 meaning not funny and 5 extremely funny). No
16
H.L. Gallagher et al. / Neuropsychologia 38 (2000) 11±21
signi®cant di€erences were seen in any of the measures
between the two conditions.
During each scanning epoch four pictures were presented on the screen each for 8.15 s. Subjects were
instructed to look at each image during the theory of
mind and non-theory of mind epochs and to consider
the meaning silently. Subjects were also asked `just to
look' at the jumbled pictures in the control epochs.
Each epoch was preceded by a relevant picture
prompt, lasting 1 s, of either a brain, indicating a theory of mind epoch, a brain with a large cross through
it, indicating a non-theory of mind epoch, or a face,
indicating a jumbled picture epoch. The subjects were
shown the prompts prior to scanning to introduce
them to this convention. The fourth image of each
epoch was replaced by the next prompt indicating the
start of a new epoch.
Following the scanning session each subject was
shown the same cartoons again and asked to explain
the meaning of each to provide a measure of performance. Subjects scored 1 for a correct answer and 0 for
an incorrect answer. As in piloting, attribution of mental states was required for explanation of ToM cartoons to be considered correct. The maximum score
possible was 28 per condition. Once again the subjects
performance was close to ceiling on both the ToMC
and Non-ToMC conditions. Mean scores were:
ToMC, 26.5 (SD, 0.84), Non-ToMC, 26.5 (SD, 2.14).
2.3. Data acquisition
A Siemens VISION MRI system operating at 2
Tesla was used to acquire both T1 weighted anatomical and echo-planar T2 weighted image volumes with
blood oxygenation level-dependent (BOLD) contrast.
Functional images were acquired over two separate
runs, a story run and a cartoon run, the order of the
tasks and conditions was counterbalanced across subjects. Each image volume constituted 48 3 mm axial
slices with in-plane resolution of 3 3 mm positioned
to cover the whole brain. Volumes were acquired continuously every 4200 ms while subjects performed three
experimental tasks, each task epoch comprised 8 image
volumes. The story run comprised four theory of mind
epochs and four non-theory of mind epochs, the cartoon run included seven theory of mind epochs and
seven non-theory of mind epochs. Activation epochs
were interspersed with control (rest) conditions. Each
run began with six volumes which were discarded prior
to analysis to allow for T1 saturation e€ects. A total
of 364 volumes were acquired of which 352 were analysed. The duration of the scanning was approximately
40±45 min.
2.4. Data analysis
Data were analysed using Statistical Parametric
Mapping (SPM97, Wellcome Dept. of Cognitive
Neurology, London, UK) implemented in MATLAB
(Mathworks Inc., Sherborn, MA, USA) and run on a
SPARC workstation (Sun Microsystems Inc., Surrey,
UK). The imaging time series was realigned using the
®rst image and spatially normalised to the stereotactic
space of Tailarach and Tournoux [31] using MNI templates (Montreal Neurological Institute). These data
were subsequently smoothed with an isotropic
Gaussian kernel of 9 mm at full width half maximum.
Analysis was carried out using the general linear
model and a delayed boxcar waveform. Subject-speci®c
low-frequency drift in signal was removed by a high
pass ®lter and global signal changes were removed by
including a global covariate [21]. E€ects at each voxel
were estimated and regionally speci®c e€ects were
compared using linear contrasts. The resulting set of
voxel values for each contrast constituted a statistical
parametric map of the t statistic (SPM{t }) which was
subsequently transformed to the unit normal distribution, SPM{Z }. Statistical inferences were based on
the theory of random Gaussian ®elds [13]). We report
activations signi®cant at p < 0.05 corrected for multiple comparisons. In regions about which we had an a
priori hypothesis, activations are considered signi®cant
at p < 0.001 uncorrected [15].
The stereotactic coordinates of Talairach and
Tournoux [31] are used to report the observed activation foci. However, descriptions of the anatomical
localisation of the foci were determined using averaged
structural MRIs of the group and the atlas of
Duvernoy [9].
3. Results
3.1. Theory of mind stories vs non-theory of mind
stories
Signi®cant activations were seen in the medial prefrontal cortex, the temporal poles bilaterally and the
temporo-parietal junctions bilaterally (exact coordinates are given in Table 1). All these regions activated
in the comparison of ToMS vs control and, with the
exception of the medial prefrontal cortex, in NonToMS vs control. No signi®cant activations were seen
in the reverse contrast; Non-ToMS activation subtracting ToMS activation.
3.2. Theory of mind cartoons vs non-theory of mind
cartoons
Signi®cant activations were seen in the medial pre-
H.L. Gallagher et al. / Neuropsychologia 38 (2000) 11±21
17
Table 1
Regions of increased brain activity associated with theory of mind compared with non-theory of mind for (i) story comprehension, (ii) cartoons,
and (iii) story comprehension and cartoons
Region
(i) ToM vs non-ToM stories
Medial prefrontal gyrus
L temporal pole
R temporal pole
L temporo-parietal junction
R temporo-parietal junction
(ii) ToM vs non-ToM cartoons
Medial prefrontal gyrus
R middle frontal gyrus
R temporo-parietal junction
Precuneus
Fusiform
(iii) ToM vs non-ToM stories and cartoons
Medial prefrontal gyrus
R middle frontal gyrus
L temporal pole
L temporal-parietal junction
R temporal-parietal junction
Precuneus
a
Putative Brodmann area
x
y
za
Z value
8/9
38
38
39/40
39/40
ÿ8
ÿ48
54
ÿ46
66
50
14
12
ÿ56
ÿ52
10
ÿ36
ÿ44
26
8
3.86
4.15
3.66
4.04
3.65
8
6
40
7/31
20/36
4
40
58
12
46
26
8
ÿ44
ÿ52
ÿ44
46
42
24
58
ÿ24
3.1
5.16
5.39
4.49
4.18
8/9
6
38
39/40
39/40
7/31
ÿ10
42
ÿ48
ÿ54
60
2
48
8
16
ÿ66
ÿ46
ÿ50
12
46
ÿ38
22
22
44
3.99
4.02
4.01
5.18
5.42
3.45
Coordinates are given with references to a standard stereotactic space (Tailarach and Tournoux [31]).
frontal cortex, the temporo-parietal junctions bilaterally, the right middle frontal gyrus, the precuneus and
the fusiform (Table 1). Once again, all these regions
activated in the comparison of ToMC vs control and,
with the exception of the medial prefrontal cortex,
Non-ToMC vs control. No signi®cant activations were
seen in the reverse contrast of Non-ToMC compared
with ToMC.
3.2.1. Conjunction: theory of mind stories vs non-theory
of mind stories and theory of mind cartoons vs nontheory of mind cartoons
Signi®cant activations were seen in the medial prefrontal cortex and the temporo-parietal junctions bilaterally, in the comparison of ToM and Non-ToM
stimuli combining both the story and cartoon tasks
(Table 1) (Fig. 3). Other regions that activated but did
not reach a corrected level of signi®cance were the
right middle frontal gyrus, the precuneus, and the left
temporal pole.
3.2.2. Interaction: (theory of mind vs non-theory of
mind) (story comprehension vs cartoons)
This interaction demonstrated regions of increased
activity associated with theory of mind, speci®c to the
story comprehension task. Only one region, the medial
prefrontal cortex was shown to be signi®cantly activated (Table 2).
3.2.3. Interaction: (theory of mind vs non-theory of
mind) (cartoons vs story comprehension)
This interaction demonstrated regions of increased
activity associated with theory of mind speci®c to the
cartoon task. Signi®cant activations were seen in the
right middle frontal gyrus the precuneus and the cerebellum (Table 2).
4. Discussion
Our study sought to clarify further the functional
anatomy of `theory of mind' using fMRI. We
attempted to examine anatomical convergence between
`theory of mind' tasks in two domains, verbal and
visual, and identify modality speci®c regions for `theory of mind' in these domains. The results of this experiment corroborate the evidence of the previous
study by Fletcher et al. [10] and also a di€erent study
by Goel et al. [15], suggesting that the ability to mentalise is mediated by the medial prefrontal cortex. In
addition, our results suggest that this region is activated by ToM tasks regardless of modality. A conjunction analysis of ToM vs Non-ToM activation common
to both tasks, also demonstrated increased activity of
the temporo-parietal junctions bilaterally. However,
the medial prefrontal cortex was the only region
uniquely activated in the theory of mind condition. All
other regions were also activated (albeit to a signi®cantly lesser extent) in the Non-ToM vs control contrast.
The verbal ToM task activated a broader region of
the medial prefrontal cortex which extended anteriorly
and inferiorly. According to the atlas of Tailarach and
Tournoux [31] activation elicited by ToM cartoons
18
H.L. Gallagher et al. / Neuropsychologia 38 (2000) 11±21
Fig. 3. A statistical parametric map (SPM{Z }) as a maximum intensity projection showing the areas where there was greater activation to theory
of mind stories and cartoons compared to non-theory of mind stories and cartoons. Views are from the right, top and behind.
was restricted to Brodmann's area 8, while that associated with ToM stories extended into the adjacent area
9. The cartoon and story tasks in the present study
were not equated for diculty, and may have di€ered
in the level of theory of mind or degree of mental state
embedding which they elicited from participants.
Whilst involvement of the medial prefrontal cortex was
demonstrated in both, the extent of the activation was
greater in the verbal task. This may have re¯ected
possible di€erences in the subtracted Non-ToM task;
in viewing cartoons, participants may try to work out
what the cartoonist intended the joke to meanÐand
this may lead to a degree of theory of mind activity
even during viewing of cartoons without mental state
content. The cartoon task was associated with
increased activity in additional regions; the precuneus
(BA7), the middle frontal gyrus (BA6) and the cerebellum. Once again, these regions activated in NonToMC compared to the control condition (JC) and
therefore cannot be regarded as speci®c to theory of
mind.
Common to both the story comprehension and the
cartoon tasks we found widespread increased activation of the temporo-parietal junctions bilaterally. Left
sided activations of this region are often seen in imaging studies of language processing and have been
attributed to semantic knowledge of single words, (x,
y, and z, coordinates: ÿ40, ÿ70, 24) [25,32,33]. This is
borne out by lesion data [1,8]. Bilateral activations of
a portion of this same region were elicited by Puce et
al. [26] (x, y, and z, coordinates: 51, ÿ49, 5) when subjects perceived movements of the eyes and mouth compared to non-facial movement in the same part of the
visual ®eld or movement of a radial background. An
area posterior to that of Puce's [26] but within the
boundaries of the region of the current study, was also
activated by Bonda et al. [4] (x, y, and z, coordinates:
56, ÿ54, 8) by the perception of simulated hand
actions and body movements compared to object and
random motion. These two studies have been interpreted as showing a role for this region in the perception of biological motion. Our theory of mind
H.L. Gallagher et al. / Neuropsychologia 38 (2000) 11±21
19
Table 2
Regions of increased brain activity associated with theory of mind compared with non-theory of mind for (i) story comprehension and (ii) cartoons
Region
(i) Interaction: (ToM vs non-ToM) (stories vs pictures)
Medial prefrontal gyrus
(ii) Interaction (ToM vs non-Tom) (pictures vs stories)
R middle frontal gyrus
Precuneus
Flocculus
Putative Brodmann area
x
y
z
Z value
9
10
50
30
3.29
6
7/31
54
2
ÿ24
8
ÿ56
ÿ30
34
50
ÿ42
3.94
3.33
4.05
materials, in which there was no motion, also activated
these regions. This may suggest that the region of the
temporo-parietal junction is sensitive not merely to
biological motion but, more generally, to stimuli which
signal intentions or intentional activity.
The critical area activated in association with theory
of mind was medial prefrontal cortex. By examining
each individual subject's activations plotted onto his/
her own T1 weighted structural images we pinpointed
this region to the medial convexity labelled BA 8/9 by
Tailarach and Tournoux [31], but closely associated
with the anterior cingulate region of BA 32 (Fig. 4).
This is consistent with previous PET studies of theory
of mind that have referred to the region as BA 8/9,
predominantly on the left [10,15]. Since the cytoarchitecture of this region is not well de®ned, the area of
activation would be more appropriately labelled as
paracingulate cortex (R. Passingham, personal communication). In those subjects with two or more cingulate sulci the activity was seen in the paracingulate
sulcus. A similar region was shown to be activated by
Bottini et al. [5] when processing of metaphorical sentences was compared with processing of literal sentences. In addition, they found activation in a region
of right middle frontal gyrus that corresponds closely
to the region (BA 6) activated in the current study
during the consideration of cartoon meaning, and particularly theory of mind cartoon meaning. This
suggests a role for attribution of mental states during
interpretation of metaphorical utterances. Indeed,
Happe [16] has demonstrated a strong theoretical and
empirical relation between theory of mind capacities
and understanding of ®gurative language (metaphor,
sarcasm) in normal children and those with autism. A
role for right hemisphere regions in understanding of
mental states is supported by neurological studies
showing impairments on theory of mind tasks following right hemisphere stroke [18,30].
The medial parietal ( precuneus ) region was activated
in all the conditions when compared with the appropriate baseline control. However, this region was
shown to be signi®cantly more activated in the ToMC
than the Non-ToMC task. Previous functional imaging
studies have suggested a role for the precuneus in men-
tal imagery at the retrieval stage of episodic memory
[11,29]. The study by Fletcher et al. [10] which used
only the story comprehension task, reported a deactivation of the precuneus in the comparison of unlinked
sentences vs stories. By lowering the threshold in this
current study we saw activity associated with mentalising in the story comprehension task. This may indicate
that the deactivation in the original paper resulted
from a subthreshold activation in the ToMS vs
unlinked sentences comparison, and suggests increased
use of mental imagery in theory of mind tasks.
The interaction condition showed activations in the
left ¯occulus of the cerebellum. There is a growing
body of literature suggesting a role for the cerebellum
in higher cognitive functions such as non-motor learning, executive function, spatial cognition, language and
emotional regulation of behaviour [23,27,28].
However, in this instance we deduct the use of di€erential strategies for scanning pictures and text, which
is consistent with previous studies associating the ¯ocullus with the control of eye movements [22]. Finally,
increased activity of the fusiform was shown to be associated with understanding the meaning of visual
jokes in this study, particularly during the theory of
mind task. This region is known to be associated with
the processing of faces and objects [3].
5. Conclusions
We have reported a study of the functional anatomy
of theory of mind, using tasks in visual and verbal
modalities. Story and cartoon tasks requiring mental
state attribution engaged speci®c networks of cortical
regions, and showed common areas of increased activation in the medial prefrontal gyrus and the temporoparietal junctions bilaterally. An area of medial prefrontal cortex (the paracingulate cortex) was the only
region uniquely activated by theory of mind tasks, and
not activated above baseline in the comparison stories
or cartoons. This provides further evidence that the
ability to attribute mental states is mediated by this
highly circumscribed brain system, and that such activation is independent of modality. These results have
20
H.L. Gallagher et al. / Neuropsychologia 38 (2000) 11±21
Fig. 4. Area of activation in the medial frontal cortex of a single subject elicited by theory of mind stories and cartoons. Co-registration of functional and structural scans for this subject show that the activation lies in the paracingulate cortex.
implications for our understanding of developmental
disorders of theory of mind, speci®cally autism.
[8]
[9]
References
[10]
[1] Alexander MP, Hiltbrunner B, Fischer RS. Distibuted anatomy
of transcortical sensory aphasia. Archives of Neurology
1989;46:885±92.
[2] Baron-Cohen S, Ring H, Moriarty J, Schmitz B, Costa D, Ell
P. The brain basis of theory of mind: the role of the orbitofrontal region. British Journal of Psychiatry 1994;165:640±9.
[3] Bly BM, Kosslyn SM. Functional anatomy of object recognition
in humans: evidence from positron emission tomography and
functional magnetic resonance imaging. Current Opinion in
Neurology 1997;10:5±9.
[4] Bonda E, Petrides M, Ostry D, Evans A. Speci®c involvement
of human parietal systems and the amygdala in the perception
of biological motion. The Journal of Neuroscience
1996;16:3737±44.
[5] Bottini G, Corcoran R, Sterzi R, Paulesu E, Schenone P,
Scarpa P, Frackowiak RSJ, Frith CD. The role of the right
hemisphere in the interpretation of ®gurative aspects of
language: a positron emission tomography activation study.
Brain 1994;117:1241±53.
[6] Carruthers P, Smith PK, editors. Theories of theories of mind.
Cambridge: Cambridge University Press, 1996.
[7] Corcoran R, Cahill C, Frith CD. The appreciation of visual
[11]
[12]
[13]
[14]
[15]
[16]
[17]
jokes in people with schizophrenia: a study of `mentalising' ability. Schizophrenia Research 1997;24:319±27.
Damasio H, Grabowski TJ, Hichwa RD, Damasio AR. A
neural basis for lexical retreival. Nature 1996;380:499±505.
Duvernoy H. The human brain: surface, three-dimensional sectional anatomy and MRI. Wien, New York: Springer-Verlag,
1991.
Fletcher PC, Happe F, Frith U, Baker SC, Dolan RJ,
Frackowiak RSJ, Frith CD. Other minds in the brain: a functional imaging study of `theory of mind' in story comprehension. Cognition 1995;57:109±28.
Fletcher PC, Frith CD, Baker SC, Shallice T, Frackowiak RS,
Dolan RJ. The minds eyeÐprecuneus activation in memory related imagery. Neuroimage 1995;2:195±200.
Fodor JA. A theory of the child's theory of mind. Cognition
1992;44:283±96.
Friston KJ, Holmes AP, Worsley KJ, Poline J-P, Frith CD,
Frackowiak RSJ. Statistical parametric maps in functional imaging: a general linear approach. Human Brain Mapping
1995;2:189±210.
Frith CD, Corcoran R. Exploring `theory of mind' in people
with schizophrenia. Psychological Medicine 1996;26:521±30.
Goel V, Grafman J, Sadato N, Hallett M. Modelling other
minds. Neuroreport 1995;6:1741±6.
Happe FGE. Communicative competance and theory of mind
in autism: a test of relevance theory. Cognition 1993;48:101±19.
Happe F. An advanced test of theory of mind: understanding of
story characters' thought and feelings by able autistics, mentally
handicapped and normal children and adults. Journal of
Autism and Developmental Disorders 1994;24:129±54.
H.L. Gallagher et al. / Neuropsychologia 38 (2000) 11±21
[18] Happe F, Brownell H, Winner E Acquired `theory of mind'
impairments following stroke. (submitted).
[19] Happe F, Ehlers S, Fletcher P, Frith U, Johansson M, Gillberg
C, Dolan R, Frackowiak R, Frith C. `Theory of Mind' in the
brain. Evidence from a PET scan study of Asperger syndrome.
Neuroreport 1996;8:197±201.
[20] Happe F, Frith U. Theory of mind in autism. In: Schopler E,
Mesibov GB, editors. Learning and cognition in autism. New
York: Plenum, 1996.
[21] Holmes AP, Josephs O, BuÈchel C, Friston KJ. Statistical modelling of low-frequency confounds in fMRI. Neuroimage
1997;5:s480.
[22] Krauzlis RJ, Lisberger SG. Directional organisation of eye
movement and visual signals in the ¯occular lobe of the monkey
cerebellum. Experimental Brain Research 1996;109:289±302.
[23] Leiner HC, Leiner AL, Dow RS. The underestimated cerebellum. Human Brain Mapping 1995;2:244±54.
[24] Leslie AM, Thaiss L. Domain speci®city in conceptual development: neuropsychological evidence from autism. Cognition
1992;43:225±51.
[25] Price C, Moore C, Humphreys GW, Wise R. Segregating
semantic from phonological processes during reading. Journal
of Cognitive Neuroscience 1997;9:727±33.
21
[26] Puce A, Allison T, Bentin S, Gore JC, McCarthy G. Temporal
cortex activation in humans viewing eye and mouth movements.
Journal of Neuroscience 1998;18:2188±99.
[27] Raichle ME, Fiez JA, Videon TO, MacLeod AM, Pardo JV,
Fox PT, Peterson SE. Practise related changes in human brain
functional anatomy during non-motor learning. Cerebral Cortex
1994;4:8±26.
[28] Schmahmann JD, Sherman JC. The cerebellar cognitive a€ective syndrome. Brain 1998;121:561±79.
[29] Shallice T, Fletcher P, Frith CD, Grasby P, Frackowiak RSJ,
Dolan R. Brain regions associated with acquisition and retrieval
of verbal episodic memory. Nature 1994;368:633±5.
[30] Siegal M, Carrington J, Radel M. Theory of mind and pragmatic understanding following right hemisphere damage. Brain
and Language 1996;53:40±50.
[31] Tailarach J, Tournoux P. Coplanar stereotactic atlas of the
human brain. Stuttgart: George Thieme Verlag, 1988.
[32] Vandenbergh R, Price C, Wise R, Josephs O, Frackowiak RSJ.
Functional anatomy of a common semantic system for words
and pictures. Nature 1996;383:254±6.
[33] Warburton E, Wise RJS, Price C, Weiller C, Hadar U, Ramsay
S, Frackowiak RSJ. Noun and verb retreival by normal subjects: studies with PET. Brain 1996;119:159±79.