Reading the mind in cartoons and stories: an fMRI study of`theory of mind' in verbal and nonverbal tasks

H.L. Gallaghera, F. Happe b, N. Brunswicka, P.C. Fletchera, U. Frithc, C.D. Fritha,*aWellcome Department of Cognitive Neurology, Institute of Neurology, University College London, 12 Queen Square, London WC1N 3BG, UK

bSocial, Genetic and Developmental Psychiatry Research Centre, Institute of Psychiatry, Denmark Hill, London, UKcInstitute 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'Ðtheattribution 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 functionalmagnetic resonance imaging study (fMRI) involving two rather di�erent tasks both designed to tap theory of mind. Brainactivation during the theory of mind condition of a story task and a cartoon task showed considerable overlap, speci®cally inthe 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, chap-

ters 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 cog-

nitive mechanism [12,24]. This proposal gains particu-

lar support from studies of autism, a biologically-

based 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

the neural systems which are abnormal in people withautism, most of whom are unable to `mind-read'.

To date, there have been three published reports offunctional brain imaging studies of `theory of mind'.Baron-Cohen et al. [2] used single photon emissioncomputerised tomography (SPECT) and a regions ofinterest approach to isolate brain areas activatedduring recognition of mental state terms in a word list.They found that their normal adult volunteers showedincreased cerebral blood ¯ow during the mental staterecognition task in the right orbito-frontal cortex rela-tive to the left frontal-polar region. Goel et al. [15]used PET to scan adults engaged in a complex task inwhich subjects had to model the knowledge and infer-ence of another mind concerning the function of unfa-miliar and familiar objects. They found widespreadactivation associated with this task, including acti-vation of left medial frontal lobe and left temporallobe. Fletcher et al. [10] also used PET, and scannedvolunteers asked to read and answer questions aboutstories. Comparison of activation during `theory ofmind' stories (requiring mental state attribution) vs

Neuropsychologia 38 (2000) 11±21

0028-3932/99/$ - see front matter # 1999 Elsevier Science Ltd. All rights reserved.

PII: S0028-3932(99 )00053 -6

www.elsevier.com/locate/neuropsychologia

* Corresponding author. Tel.: +44-171-833-7472; fax: +44-171-

813-1420.

E-mail address: c.frith@®l.ion.ucl.ac.uk (C.D. Frith)

Fig. 1. Examples of story comprehension passages.

H.L. Gallagher et al. / Neuropsychologia 38 (2000) 11±2112

control `physical' stores revealed task-speci®c acti-vation in the left medial prefrontal gyrus, as well asincreased activation of the posterior cingulate cortex.Using the same technique with individuals with a formof autism (Asperger Syndrome), Happe et al. [20]found similar patterns of activation in all regionsexcept the medial frontal gyrus which had been linkedto theory of mind performance in the normal group.

The main aims of the current study were to investi-gate the neural correlates underlying theory of mindby exploiting the superior spatial resolution of fMRIcompared to PET and to examine anatomical conver-gence between theory of mind tasks presented in di�er-ent modalities. The story comprehension task used byFletcher et al. [10] was modi®ed for compatibility withfMRI to examine theory of mind in the verbal domain,while captionless cartoons provided a visual equival-ent. Previous behavioural studies have used both stor-ies [14,17,18] and visual jokes [7,18] to investigatetheory of mind impairments in adults and children andfound these tasks to be good markers of mentalisingabilities. We hypothesised that activation in medialprefrontal cortex would be associated with the attribu-tion of mental states independent of modality. Our®nal aim was to identify any modality speci®c regionsfor 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 30yr (range 23±36 yr). This study was approved by theInstitute of Neurology Ethics Committee. Informedwritten consent was obtained from all subjects prior toscanning.

2.2. Tasks

All stimuli were displayed on a monitor and pre-sented to the subject via a 458 angled mirror posi-tioned above the head coil; this mirror was adjusted tobe within the subjects ®eld of vision without having totilt his/her head. A test image was presented on thescreen prior to scanning to ensure that the image wasin focus and the subject could comfortably read thetext.

2.2.1. Story comprehension taskThree types of verbal material were presented; these

were `theory of mind stories' (ToMS), `non-theory ofmind stories' (Non-ToMS) and `unlinked sentences'(US). Fig. 1. shows examples of these three types ofmaterial.

During each scanning epoch a passage of text waspresented on the screen for 21.6 s. The passage wasthen replaced by a question relating to the text (pre-sented for a further 11 s). Subjects were instructed toread the passage silently and to answer internally thequestion which followed. Each passage was precededby a relevant prompt to indicate either `theory ofmind', `unlinked sentences' or `non-theory of mind'.This prompt was displayed for 1 s. The question was

Fig. 1 (continued)

H.L. Gallagher et al. / Neuropsychologia 38 (2000) 11±21 13

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±2114

replaced by the next prompt indicating the start of anew epoch.

Following the scanning session each subject wasshown the same passages again and asked to give his/her answer to each question, to provide a measure ofbehavioural performance. Responses were scored 1 fora correct answer and 0 for an incorrect answer. Forthe theory of mind condition an answer was con-sidered correct only if an appropriate mental state wasattributed to one or more characters. The maximumscore was 4 per condition which was reached by allsubjects, thus indicating full understanding.

The passages used in these tasks have been used intwo previous experiments by this group [10,19]. Theresults from this study con®rmed that the tasks werematched for di�culty, as re¯ected in scores and read-ing times, with ToM stories taking if anything slightly(but nonsigni®cantly) less time to read than the Non-ToMS.

2.2.2. Cartoon taskThree 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 consideredto require theory of mind for its interpretation if attri-

bution of either false belief or ignorance to one ormore of the characters in the picture was vital forcomprehension. A cartoon was considered to be non-theory of mind if no mental state attribution wasneeded to understand the meaning. The `jumbled pic-tures' were constructed from images of randomly posi-tioned objects, animals and people, taken fromcartoons and children's colouring books. All imageswere captionless. Fig. 2 shows examples of the threestimulus types.

The cartoons were validated in a pilot study withtwenty naive normal subjects (age range 19±66 yr).Subjects were instructed to look at each cartoon andindicate to the experimenter as soon as they under-stood its meaning. Response time was recorded.Subjects then gave a brief explanation of the cartoon'smeaning. The explanations were scored 1 for a correctanswer and 0 for an incorrect answer. As for the storycondition, explanations in the theory of mind con-dition were considered correct only if an appropriatemental state was attributed to one or more characters.The explanation was recorded along with the timetaken. In addition, participants were asked to rate,from 1 to 5, how di�cult and how funny they thoughtthe cartoon to be (1 extremely easy, 5 extremely di�-cult; 1 meaning not funny and 5 extremely funny). No

Fig. 2 (continued)

H.L. Gallagher et al. / Neuropsychologia 38 (2000) 11±21 15

signi®cant di�erences were seen in any of the measuresbetween the two conditions.

During each scanning epoch four pictures were pre-sented on the screen each for 8.15 s. Subjects wereinstructed to look at each image during the theory ofmind and non-theory of mind epochs and to considerthe meaning silently. Subjects were also asked `just tolook' at the jumbled pictures in the control epochs.Each epoch was preceded by a relevant pictureprompt, lasting 1 s, of either a brain, indicating a the-ory of mind epoch, a brain with a large cross throughit, indicating a non-theory of mind epoch, or a face,indicating a jumbled picture epoch. The subjects wereshown the prompts prior to scanning to introducethem to this convention. The fourth image of eachepoch was replaced by the next prompt indicating thestart of a new epoch.

Following the scanning session each subject wasshown the same cartoons again and asked to explainthe meaning of each to provide a measure of perform-ance. Subjects scored 1 for a correct answer and 0 foran incorrect answer. As in piloting, attribution of men-tal states was required for explanation of ToM car-toons to be considered correct. The maximum scorepossible was 28 per condition. Once again the subjectsperformance was close to ceiling on both the ToMCand 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 2Tesla was used to acquire both T1 weighted anatom-ical and echo-planar T2� weighted image volumes withblood oxygenation level-dependent (BOLD) contrast.Functional images were acquired over two separateruns, a story run and a cartoon run, the order of thetasks and conditions was counterbalanced across sub-jects. Each image volume constituted 48 3 mm axialslices with in-plane resolution of 3 � 3 mm positionedto cover the whole brain. Volumes were acquired con-tinuously every 4200 ms while subjects performed threeexperimental tasks, each task epoch comprised 8 imagevolumes. The story run comprised four theory of mindepochs and four non-theory of mind epochs, the car-toon run included seven theory of mind epochs andseven non-theory of mind epochs. Activation epochswere interspersed with control (rest) conditions. Eachrun began with six volumes which were discarded priorto analysis to allow for T1 saturation e�ects. A totalof 364 volumes were acquired of which 352 were ana-lysed. The duration of the scanning was approximately40±45 min.

2.4. Data analysis

Data were analysed using Statistical ParametricMapping (SPM97, Wellcome Dept. of CognitiveNeurology, London, UK) implemented in MATLAB(Mathworks Inc., Sherborn, MA, USA) and run on aSPARC workstation (Sun Microsystems Inc., Surrey,UK). The imaging time series was realigned using the®rst image and spatially normalised to the stereotacticspace of Tailarach and Tournoux [31] using MNI tem-plates (Montreal Neurological Institute). These datawere subsequently smoothed with an isotropicGaussian kernel of 9 mm at full width half maximum.

Analysis was carried out using the general linearmodel and a delayed boxcar waveform. Subject-speci®clow-frequency drift in signal was removed by a highpass ®lter and global signal changes were removed byincluding a global covariate [21]. E�ects at each voxelwere estimated and regionally speci®c e�ects werecompared using linear contrasts. The resulting set ofvoxel values for each contrast constituted a statisticalparametric map of the t statistic (SPM{t }) which wassubsequently transformed to the unit normal distri-bution, SPM{Z }. Statistical inferences were based onthe theory of random Gaussian ®elds [13]). We reportactivations signi®cant at p < 0.05 corrected for mul-tiple comparisons. In regions about which we had an apriori hypothesis, activations are considered signi®cantat p < 0.001 uncorrected [15].

The stereotactic coordinates of Talairach andTournoux [31] are used to report the observed acti-vation foci. However, descriptions of the anatomicallocalisation of the foci were determined using averagedstructural MRIs of the group and the atlas ofDuvernoy [9].

3. Results

3.1. Theory of mind stories vs non-theory of mindstories

Signi®cant activations were seen in the medial pre-frontal cortex, the temporal poles bilaterally and thetemporo-parietal junctions bilaterally (exact coordi-nates are given in Table 1). All these regions activatedin the comparison of ToMS vs control and, with theexception of the medial prefrontal cortex, in Non-ToMS vs control. No signi®cant activations were seenin the reverse contrast; Non-ToMS activation subtract-ing ToMS activation.

3.2. Theory of mind cartoons vs non-theory of mindcartoons

Signi®cant activations were seen in the medial pre-

H.L. Gallagher et al. / Neuropsychologia 38 (2000) 11±2116

frontal cortex, the temporo-parietal junctions bilater-ally, the right middle frontal gyrus, the precuneus andthe fusiform (Table 1). Once again, all these regionsactivated in the comparison of ToMC vs control and,with the exception of the medial prefrontal cortex,Non-ToMC vs control. No signi®cant activations wereseen in the reverse contrast of Non-ToMC comparedwith ToMC.

3.2.1. Conjunction: theory of mind stories vs non-theoryof mind stories and theory of mind cartoons vs non-theory of mind cartoons

Signi®cant activations were seen in the medial pre-frontal cortex and the temporo-parietal junctions bilat-erally, in the comparison of ToM and Non-ToMstimuli combining both the story and cartoon tasks(Table 1) (Fig. 3). Other regions that activated but didnot reach a corrected level of signi®cance were theright middle frontal gyrus, the precuneus, and the lefttemporal pole.

3.2.2. Interaction: (theory of mind vs non-theory ofmind) � (story comprehension vs cartoons)

This interaction demonstrated regions of increasedactivity associated with theory of mind, speci®c to thestory comprehension task. Only one region, the medialprefrontal cortex was shown to be signi®cantly acti-vated (Table 2).

3.2.3. Interaction: (theory of mind vs non-theory ofmind) � (cartoons vs story comprehension)

This interaction demonstrated regions of increased

activity associated with theory of mind speci®c to thecartoon task. Signi®cant activations were seen in theright middle frontal gyrus the precuneus and the cer-ebellum (Table 2).

4. Discussion

Our study sought to clarify further the functionalanatomy of `theory of mind' using fMRI. Weattempted to examine anatomical convergence between`theory of mind' tasks in two domains, verbal andvisual, and identify modality speci®c regions for `the-ory of mind' in these domains. The results of this ex-periment corroborate the evidence of the previousstudy by Fletcher et al. [10] and also a di�erent studyby Goel et al. [15], suggesting that the ability to men-talise is mediated by the medial prefrontal cortex. Inaddition, our results suggest that this region is acti-vated by ToM tasks regardless of modality. A conjunc-tion analysis of ToM vs Non-ToM activation commonto both tasks, also demonstrated increased activity ofthe temporo-parietal junctions bilaterally. However,the medial prefrontal cortex was the only regionuniquely activated in the theory of mind condition. Allother regions were also activated (albeit to a signi®-cantly lesser extent) in the Non-ToM vs control con-trast.

The verbal ToM task activated a broader region ofthe medial prefrontal cortex which extended anteriorlyand inferiorly. According to the atlas of Tailarach andTournoux [31] activation elicited by ToM cartoons

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 Putative Brodmann area x y za Z value

(i) ToM vs non-ToM stories

Medial prefrontal gyrus 8/9 ÿ8 50 10 3.86

L temporal pole 38 ÿ48 14 ÿ36 4.15

R temporal pole 38 54 12 ÿ44 3.66

L temporo-parietal junction 39/40 ÿ46 ÿ56 26 4.04

R temporo-parietal junction 39/40 66 ÿ52 8 3.65

(ii) ToM vs non-ToM cartoons

Medial prefrontal gyrus 8 4 26 46 3.1

R middle frontal gyrus 6 40 8 42 5.16

R temporo-parietal junction 40 58 ÿ44 24 5.39

Precuneus 7/31 12 ÿ52 58 4.49

Fusiform 20/36 46 ÿ44 ÿ24 4.18

(iii) ToM vs non-ToM stories and cartoons

Medial prefrontal gyrus 8/9 ÿ10 48 12 3.99

R middle frontal gyrus 6 42 8 46 4.02

L temporal pole 38 ÿ48 16 ÿ38 4.01

L temporal-parietal junction 39/40 ÿ54 ÿ66 22 5.18

R temporal-parietal junction 39/40 60 ÿ46 22 5.42

Precuneus 7/31 2 ÿ50 44 3.45

a Coordinates are given with references to a standard stereotactic space (Tailarach and Tournoux [31]).

H.L. Gallagher et al. / Neuropsychologia 38 (2000) 11±21 17

was restricted to Brodmann's area 8, while that associ-ated with ToM stories extended into the adjacent area9. The cartoon and story tasks in the present studywere not equated for di�culty, and may have di�eredin the level of theory of mind or degree of mental stateembedding which they elicited from participants.Whilst involvement of the medial prefrontal cortex wasdemonstrated in both, the extent of the activation wasgreater in the verbal task. This may have re¯ectedpossible di�erences in the subtracted Non-ToM task;in viewing cartoons, participants may try to work outwhat the cartoonist intended the joke to meanÐandthis may lead to a degree of theory of mind activityeven during viewing of cartoons without mental statecontent. The cartoon task was associated withincreased activity in additional regions; the precuneus(BA7), the middle frontal gyrus (BA6) and the cerebel-lum. Once again, these regions activated in Non-ToMC compared to the control condition (JC) andtherefore cannot be regarded as speci®c to theory ofmind.

Common to both the story comprehension and thecartoon tasks we found widespread increased acti-vation of the temporo-parietal junctions bilaterally. Leftsided activations of this region are often seen in ima-ging studies of language processing and have beenattributed to semantic knowledge of single words, (x,y, and z, coordinates: ÿ40, ÿ70, 24) [25,32,33]. This isborne out by lesion data [1,8]. Bilateral activations ofa portion of this same region were elicited by Puce etal. [26] (x, y, and z, coordinates: 51, ÿ49, 5) when sub-jects perceived movements of the eyes and mouth com-pared to non-facial movement in the same part of thevisual ®eld or movement of a radial background. Anarea posterior to that of Puce's [26] but within theboundaries of the region of the current study, was alsoactivated by Bonda et al. [4] (x, y, and z, coordinates:56, ÿ54, 8) by the perception of simulated handactions and body movements compared to object andrandom motion. These two studies have been inter-preted as showing a role for this region in the percep-tion of biological motion. Our theory of mind

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.

H.L. Gallagher et al. / Neuropsychologia 38 (2000) 11±2118

materials, in which there was no motion, also activatedthese regions. This may suggest that the region of thetemporo-parietal junction is sensitive not merely tobiological motion but, more generally, to stimuli whichsignal intentions or intentional activity.

The critical area activated in association with theoryof mind was medial prefrontal cortex. By examiningeach individual subject's activations plotted onto his/her own T1 weighted structural images we pinpointedthis region to the medial convexity labelled BA 8/9 byTailarach and Tournoux [31], but closely associatedwith the anterior cingulate region of BA 32 (Fig. 4).This is consistent with previous PET studies of theoryof mind that have referred to the region as BA 8/9,predominantly on the left [10,15]. Since the cytoarchi-tecture of this region is not well de®ned, the area ofactivation would be more appropriately labelled asparacingulate cortex (R. Passingham, personal com-munication). In those subjects with two or more cingu-late sulci the activity was seen in the paracingulatesulcus. A similar region was shown to be activated byBottini et al. [5] when processing of metaphorical sen-tences was compared with processing of literal sen-tences. In addition, they found activation in a regionof right middle frontal gyrus that corresponds closelyto the region (BA 6) activated in the current studyduring the consideration of cartoon meaning, and par-ticularly theory of mind cartoon meaning. Thissuggests a role for attribution of mental states duringinterpretation of metaphorical utterances. Indeed,Happe [16] has demonstrated a strong theoretical andempirical relation between theory of mind capacitiesand understanding of ®gurative language (metaphor,sarcasm) in normal children and those with autism. Arole for right hemisphere regions in understanding ofmental states is supported by neurological studiesshowing impairments on theory of mind tasks follow-ing right hemisphere stroke [18,30].

The medial parietal ( precuneus ) region was activatedin all the conditions when compared with the appro-priate baseline control. However, this region wasshown to be signi®cantly more activated in the ToMCthan the Non-ToMC task. Previous functional imagingstudies 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 usedonly the story comprehension task, reported a deacti-vation of the precuneus in the comparison of unlinkedsentences vs stories. By lowering the threshold in thiscurrent study we saw activity associated with mentalis-ing in the story comprehension task. This may indicatethat the deactivation in the original paper resultedfrom a subthreshold activation in the ToMS vsunlinked sentences comparison, and suggests increaseduse of mental imagery in theory of mind tasks.

The interaction condition showed activations in theleft ¯occulus of the cerebellum. There is a growingbody of literature suggesting a role for the cerebellumin higher cognitive functions such as non-motor learn-ing, executive function, spatial cognition, language andemotional regulation of behaviour [23,27,28].However, in this instance we deduct the use of di�er-ential strategies for scanning pictures and text, whichis consistent with previous studies associating the ¯o-cullus with the control of eye movements [22]. Finally,increased activity of the fusiform was shown to be as-sociated with understanding the meaning of visualjokes in this study, particularly during the theory ofmind task. This region is known to be associated withthe processing of faces and objects [3].

5. Conclusions

We have reported a study of the functional anatomyof theory of mind, using tasks in visual and verbalmodalities. Story and cartoon tasks requiring mentalstate attribution engaged speci®c networks of corticalregions, and showed common areas of increased acti-vation in the medial prefrontal gyrus and the temporo-parietal junctions bilaterally. An area of medial pre-frontal cortex (the paracingulate cortex) was the onlyregion uniquely activated by theory of mind tasks, andnot activated above baseline in the comparison storiesor cartoons. This provides further evidence that theability to attribute mental states is mediated by thishighly circumscribed brain system, and that such acti-vation is independent of modality. These results have

Table 2

Regions of increased brain activity associated with theory of mind compared with non-theory of mind for (i) story comprehension and (ii) car-

toons

Region Putative Brodmann area x y z Z value

(i) Interaction: (ToM vs non-ToM) � (stories vs pictures)

Medial prefrontal gyrus 9 10 50 30 3.29

(ii) Interaction (ToM vs non-Tom) � (pictures vs stories)

R middle frontal gyrus 6 54 8 34 3.94

Precuneus 7/31 2 ÿ56 50 3.33

Flocculus ÿ24 ÿ30 ÿ42 4.05

H.L. Gallagher et al. / Neuropsychologia 38 (2000) 11±21 19

implications for our understanding of developmentaldisorders of theory of mind, speci®cally autism.

References

[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 orbito-

frontal 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

jokes in people with schizophrenia: a study of `mentalising' abil-

ity. Schizophrenia Research 1997;24:319±27.

[8] Damasio H, Grabowski TJ, Hichwa RD, Damasio AR. A

neural basis for lexical retreival. Nature 1996;380:499±505.

[9] Duvernoy H. The human brain: surface, three-dimensional sec-

tional anatomy and MRI. Wien, New York: Springer-Verlag,

1991.

[10] Fletcher PC, Happe F, Frith U, Baker SC, Dolan RJ,

Frackowiak RSJ, Frith CD. Other minds in the brain: a func-

tional imaging study of `theory of mind' in story comprehen-

sion. Cognition 1995;57:109±28.

[11] Fletcher PC, Frith CD, Baker SC, Shallice T, Frackowiak RS,

Dolan RJ. The minds eyeÐprecuneus activation in memory re-

lated imagery. Neuroimage 1995;2:195±200.

[12] Fodor JA. A theory of the child's theory of mind. Cognition

1992;44:283±96.

[13] Friston KJ, Holmes AP, Worsley KJ, Poline J-P, Frith CD,

Frackowiak RSJ. Statistical parametric maps in functional ima-

ging: a general linear approach. Human Brain Mapping

1995;2:189±210.

[14] Frith CD, Corcoran R. Exploring `theory of mind' in people

with schizophrenia. Psychological Medicine 1996;26:521±30.

[15] Goel V, Grafman J, Sadato N, Hallett M. Modelling other

minds. Neuroreport 1995;6:1741±6.

[16] Happe FGE. Communicative competance and theory of mind

in autism: a test of relevance theory. Cognition 1993;48:101±19.

[17] 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.

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 func-

tional and structural scans for this subject show that the activation lies in the paracingulate cortex.

H.L. Gallagher et al. / Neuropsychologia 38 (2000) 11±2120

[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, BuÈ chel C, Friston KJ. Statistical model-

ling 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 cerebel-

lum. Human Brain Mapping 1995;2:244±54.

[24] Leslie AM, Thaiss L. Domain speci®city in conceptual develop-

ment: 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.

[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�ec-

tive 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 prag-

matic 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 sub-

jects: studies with PET. Brain 1996;119:159±79.

H.L. Gallagher et al. / Neuropsychologia 38 (2000) 11±21 21