doi:10.1016/S1053 8119(03)00230 1

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1 www.elsevier.com/locate/ynimg NeuroImage 19 (2003) 1835–1842 Rapid Communication People thinking about thinking people The role of the temporo-parietal junction in “theory of mind” a, a,b R. Saxe * and N. Kanwisher a Department of Brain and Cognitive Sciences, Massachusetts Institute of Technology, Cambridge, MA, USA b McGovern Institute for Brain Research, Massachusetts Institute of Technology, Cambridge, MA, USA Received 15 January 2003; revised 12 March 2003; accepted 14 April 2003 Abstract Humans powerfully and flexibly interpret the behaviour of other people based on an understanding of their minds: that is, we use a “theory of mind.” In this study we distinguish theory of mind, which represents another person’s mental states, from a representation of the simple presence of another person per se. The studies reported here establish for the first time that a region in the human temporo-parietal junction (here called the TPJ-M) is involved specifically in reasoning about the contents of another person’s mind. First, the TPJ-M was doubly dissociated from the nearby extrastriate body area (EBA; Downing et al., 2001). Second, the TPJ-M does not respond to false representations in non-social control stories. Third, the BOLD response in the TPJ-M bilaterally was higher when subjects read stories about a character’s mental states, compared with stories that described people in physical detail, which did not differ from stories about nonhuman objects. Thus, the role of the TPJ-M in understanding other people appears to be specific to reasoning about the content of mental states. © 2003 Elsevier Science (USA). All rights reserved. fMRI; Social cognitive neuroscience; False belief; Mentalising; Superior temporal sulcus; EBA Keywords: have been implicated in a broad range of social cognition The remarkable human facility with social cognition depends on a fundamental ability to reason about other tasks (Allison et al., 2000; Gallagher and Frith, 2003; Green people. Specifically, we predict and interpret the behaviour and Haidt, 2003). Regions near the TPJ have preferential of people based on an understanding of their minds: that is, responses to human faces (e.g., Hoffman and Haxby, 2000), 1 we use a “theory of mind.” In this study we show that a bodies (e.g., Downing et al., 2001) and biological motion region of human temporo-parietal junction is selectively (e.g., Grossman et al., 2000). There is also some evidence involved in reasoning about the contents of other people’s that regions within human TPJ are involved in theory of minds. mind (ToM). A number of studies have reported increased Brain regions near the temporo-parietal junction (TPJ) responses in the TPJ when subjects read verbal stories or see pictorial cartoons that require inferences about a character’s (false) beliefs, compared with physical control stimuli (Fletcher et al., 1995; Brunet et al., 2000; Gallagher et al., * Corresponding author. Department of Brain and Cognitive Sciences, MIT, NE20-464, 77 Massachussetts Avenue, Cambridge, MA 02139, 2000; Castelli et al., 2000; Voegely et al., 2001. A number  USA. Fax: 1-617-258-8654. of other brain regions have also been implicated in theory of [email protected] (R. Saxe). E-mail address: mind; see reviews by Gallagher and Frith, 2003, and Greene 1 The term “theory of mind” has a more restricted sense, referring to the and Haidt, 2003). suggestion that the structure of knowledge in the mind is analogous to a scientific theory (e.g., Carey, 1985; Wellman and Gelman, 1992). For What is the role of the TPJ in these tasks? ToM reasoning discussions about the so-called theory-theory, see Carruthers and Smith, depends upon at least two kinds of representation: a repre- 1996, and Malle et al., 2001. In this study, we use the term theory of mind sentation of another person per se and a representation of in a broader sense, to refer to any reasoning about another person’s that other person’s mental states (see Leslie, 1999). While a representational mental states (also called “belief-desire psychology,” e.g., Bartsch and Wellman, 1995). representation of a person per se is a likely prerequisite for 1053-8119/03/$ – see front matter © 2003 Elsevier Science (USA). All rights reserved. doi:10.1016/S1053-8119(03)00230-1

2 Rapid Communication / NeuroImage 19 (2003) 1835–1842 1836 mental states is ToM, achieving a representation of others his/her false belief. Descriptions of human actions required ’ the core responsibility of a ToM. Some authors suggest that analysis of mental causes, in the absence of false beliefs. the TPJ is involved only in the preliminary stages of social We compared these conditions to two non-social control “ aid ” cognition that ToM, not in ToM reasoning itself (e.g., conditions, (1) mechanical inference control stories, which required the subject to infer a hidden physical (as opposed Gallagher and Frith, 2003). We here provide evidence to mental) process, such as melting or rusting (for examples, against this suggestion, and argue on the contrary that a see Appendix 1), and (2) descriptions of nonhuman objects. region of the TPJ is selectively involved in representation other peoples Unlike previous studies, we did not cue or instruct sub- ’ mental states. cally to mental states. With this design Neuroimaging studies have followed developmental psy- jects to attend speci fi we were able to look for regions of cortex in individual stories as the prototypical ” false belief “ chology in using problem for ToM reasoning (Fletcher et al., 1995; Gallagher subjects that are selectively and spontaneously involved in understanding the mental (as opposed to physical) causes of et al., 2000; see also Vogeley et al., 2001). In these scenar- s false ’ ’ s action is based on the character events. ios, a character To test whether the response to ToM stories was a re- belief (Wimmer and Perner, 1983). False beliefs provide a sponse to the presence of a person in the stimulus, we useful behavioural test of a ToM, because when the belief is presented still photographs of people, and nonhuman ob- false, the action predicted by the belief is different from the action that would be predicted by the true state of affairs jects. Downing et al. (2001) reported a bilateral region near (Dennett, 1978). Note, though, that everyday reasoning the posterior superior temporal cortex that responds prefer- entially to the visual appearance of human bodies, compared about other minds, by adults and children, depends on with a range of control objects (the extrastriate body area, attributions of mostly true beliefs (e.g., Dennett, 1996; Bar- tsch and Wellman, 1995). EBA). We tested directly the functional and anatomical relationship between the EBA and the (proposed) TPJ-M. Previous investigations of the neural correlates of ToM (Fletcher et al., 1995; Gallagher et al., 2000) have compared false belief ( “ theory of mind ” ) stories with two control Methods which describe ” non-theory of mind stories, “ conditions: actions based on the character ” control “ s true beliefs, and ’ stories, consisting of unrelated sentences. These authors ve healthy right-handed adults (12 women) fi Twenty- volunteered or participated for payment. All subjects had found that the TPJ response was high during theory of mind stories, but was also high during non-theory of mind stories. normal or corrected-to-normal vision and gave informed consent to participate in the study. They concluded (see also Gallagher and Frith, 2003) that the Subjects were scanned in the Siemens 1.5-(9 subjects) TPJ is not selectively involved in ToM. This conclusion and 3.0-T (16 subjects) scanners at the MGH-NMR center does not follow. Because the non-theory of mind stories in Charlestown, MA, using a head coil. Standard echoplanar s (true) beliefs, a re- ’ invite inferences about the character imaging procedures were used [TR gion involved in reasoning about other minds should show 2s,TE   40 (3 T) or 30 (1.5 T) ms, fl ip angle 90 ° ]. Twenty 5-mm-thick near- a high response to these stories, as well as to the so-called coronal slices (parallel to the brainstem) covered the occip- theory of mind stories. (For an argument against the use of ital lobe and the posterior portion of the temporal and unrelated sentences as the baseline condition, see Ferstl and parietal lobes. von Cramon, 2002.) Stimuli consisted of short center-justi fi ed stories, pre- We propose two basic tests for a region selectively in- volved in ToM reasoning. First, it must show increased sented in 24-point white text on a black background (aver- age number of words  36). Stories were constructed to fi response to tasks/stimuli that invite ToM reasoning (about t four categories: false belief, mechanical inference, human true or false beliefs) compared with logically similar non- action, and nonhuman descriptions (Appendix 1). Each social controls. Second, the region must respond not just story was presented for 9500 ms, followed by a 500-ms cally when a person is present in the stimulus, but speci fi interstimulus interval. Each scan lasted 260 s: four 40-s ’ s mental states. Be- when subjects reason about the person epochs, each containing four stories (one from each condi- low, we provide evidence that a subregion of the TPJ, here called the TPJ-M, passes both these criteria for a selective fi tion), and 20 s of xation between epochs. The order of role in ToM. conditions was counterbalanced within and across runs. Subjects were asked to press a button to indicate when they had fi nished reading each story. Subjects read a total of 8 (4 Experiment 1 subjects) or 12 (21 subjects) stories per condition. Fourteen of the subjects from Experiment 1 (7 women) were also scanned on an EBA localizer in the same scan We devised a new version of the false belief stories task (Fletcher et al., 1995) to compare reasoning about true and session, all at 3.0 T. Stimuli consisted of 20 grayscale photographs of whole human bodies (including faces) in a false beliefs to reasoning about non-social control situa- ’ range of postures, standing and sitting, and 20 photographs tions. ToM stories described a character s action caused by

3 1842 Rapid Communication / NeuroImage 19 (2003) 1835 1837 – Table 1 eses, we concentrated on this region in the subsequent a Experiment 1 analyses. ned TPJ-M regions of interest (ROI) in the left fi We de No. of voxels MNI coordinate Region Z ( 0.05, corrected)  (max voxel) P and right TPJ in each individual subject as contiguous voxels in each hemisphere that were more active ( P  54 5.88 LTPJ-M [   60 21] 63 0.0001) during false belief than mechanical inference sto- LaSTS 5.40 12]  55  57  [ 27 [ Prec   9 5.20 41 51 33] ries. The TPJ-M bilaterally generalized beyond false beliefs, [51  54 27] R TPJ-M 10 5.10 responding signi cantly more to human action (HA) stories fi 18  15] 4.91 2 R aSTS [66  t than to nonhuman descriptions [N-H D; paired samples a Five regions showed increased signal during theory of mind, compared tests, right: HA average percent signal change from fi xation with mechanical inference, stories (random effects, 0.05): left  n  P 25,  0.0001; left: (PSC): 0.22, N-H D average PSC: 0.02, P and right temporo-parietal junction (TPJ-M), left and right anterior superior  HA average PSC: 0.35, N-H D average PSC: 0.10, P temporal sulcus (aSTS), and precuneus (Prec). All coordinates are accord- 0.0001]. ing to the Montreal Neurological Institute standard brain. In the 14 subjects who also had an EBA localizer scan, fi EBA ROIs were de ned as the cluster of contiguous voxels of easily recognizable inanimate objects (e.g., car, drum, in extrastriate cortex (bilaterally in 13 subjects and right- tulip). (Two other conditions, cropped faces and scrambled 0.0001) during  P only in 1 subject) that was more active ( objects, were included in the scan but were not analyzed pictures of human bodies than during pictures of nonhuman here). objects in each individual subject (following Downing et al., Image presentation followed the blocked design de- 2001). Both right and left EBA ROIs failed to discriminate scribed in Tong et al. (2000; Experiment 1) except that P between any story conditions (paired samples  t tests, all images were presented at a rate of one every 800 ms (stim- 0.4, all story PSCs below 0.01; Fig. 2).  ulus duration 300 ms), 500 ms, interstimulus interval  TPJ-M response to photographs was lateralized. The left and each scan lasted 336 s. Subjects performed a one-back TPJ-M did not discriminate between photographs of people matching task (Tong et al., 2000). (PSC: 0.09, paired samples  0.04) and of objects (PSC:  MRI data were analyzed using SPM 99, FS-fast, and 0.4). The right TPJ-M showed a trend toward a t test, P  in-house software. greater response to photographs of people (PSC: 0.24) than 0.10). A of objects (PSC: 0.10, paired samples t test, P  repeated-measures ANOVA of content (person versus ob- Results ject) by stimulus modality (stories versus photograph) by hemisphere (right versus left) revealed a main effect of Average reading times for theory of mind and mechan- photographs person object (  P  0.001) and of stories  fi cantly (ToM  ical inference stories did not differ signi  P ( 0.05) modulated by an interaction between stimulus 0.2).  P 6.5 s, 6.4 s, MI  modality and hemisphere (response to photographs only on ve loci fi Random effects analyses of 25 subjects revealed P 0.005) and a trend toward a three-way  the right, of greater activation during the theory of mind compared interaction (the right TPJ-M response distinguishes photo- with mechanical inference stories ( 0.05 corrected for P   graphs of bodies and objects more than the left TPJ-M, P multiple spatial hypotheses): left and right TPJ-M, left and 0.1; Fig. 2). right anterior superior temporal sulcus (aSTS), and precu- neus (Table 1, Fig. 1). [Consistent with many previous studies (e.g., Gusnard and Raichle, 2001; Raichle et al., Discussion 2000) the precuneus was deactivated (BOLD signal less fi than xation baseline) during all of our story conditions. Experiment 1 thus shows an increased BOLD response The ToM stories deactivated the precuneus less than me- in a region of the TPJ bilaterally, here called the TPJ-M, chanical inference stories. It was therefore unclear whether during ToM compared with mechanical inference stories. this effect should be considered a response to ToM or to This activation is robust and reliable across individual sub- mechanical inference stories, and the precuneus response nding replicates the earlier reports with a new jects. This fi was not analyzed further.] set of stimuli, a less biased task (no cues), and with more The same pattern of results was apparent in individual stringent statistical tests (both individual subject analyses xed effects fi subjects ( 0.0001, uncorrected for all re-  P fi rm that and random effects group analyses). Our results con sults reported here). Voxels more responsive during ToM the TPJ-M response to verbal descriptions generalizes to than mechanical inference stories were observed at the TPJ human actions based on true beliefs. in 22 of 25 subjects (bilaterally in 14, left in 5, and right in Importantly, we distinguished the TPJ-M from its neigh- fi cant at this level 3 subjects). The aSTS activation was signi bour, the EBA, which did not respond to any verbal story in 10 of 25 subjects. Because the TPJ-M was most consis- tent across subjects and was the focus of our prior hypoth- conditions. However, the TPJ-M response to nonverbal so-

4 1842 1838 – Rapid Communication / NeuroImage 19 (2003) 1835 Experiment 2 The results of Experiment 1 established that bilateral regions near the TPJ show a greater increase in BOLD ’ signal when subjects reason about others mental states, than when they reason about nonhuman objects. However, in Experiment 1, stories involving people and mental states were compared with stories that involve neither people nor mental states. In Experiment 2, we asked which of these two components was responsible for the observed activation. We directly compared the response of the TPJ-M to stories about people that did (desires) or did not (physical people) require inferences based on mental states. culty and causal Also, while they were controlled for dif fi structure, the logical structure of the ToM stories used in Experiment 1 (and previous studies) differed systematically from the control stories: only the false belief stories require the notion of a false representation, in this case a false belief. This confounding factor was perceived by develop- mental psychologists, who invented its solution: false pho- “ Fig. 1. Experiment 1. Random effects analysis, 0.05, corrected, n  P   mechanical inference stories. Crosshair marks the 25. Theory of mind fi most signi cant voxel in the left TPJ (1). Also visible are activations in right TPJ (2), left aSTS (3), and precuneus (4). TPJ, temporo-parietal junction; aSTS, anterior superior temporal sulcus. Fig. 3. (a) Experiments 1 and 2. Activation overlap within an individual subject showing bilateral temporo-parietal junction (bilateral TPJ) and xed effects, P precuneus regions ( 0.001). Red  theory of mind  fi  xation in (a) Fig. 2. Experiment 1. Average percent signal change from fi false photo (Exp. 2). false belief mechanical inference (Exp. 1). Blue   ned in individual fi left and right TPJ-M and (b) left and right EBA, de Green both. (b) Single subject time course of response during Experi-  14). The EBA consisted of contiguous voxels in bilateral  subjects ( n ment 2 to false belief (dark gray) and false photograph (white) stories in the extrastriate cortex that responded signi cantly more to pictures of human fi same subject ned by a greater response to fi s TPJ-M, independently de ’ bodies than pictures of nonhuman objects ( P  0.0001, uncorrected). The  theory of mind than to mechanical inference stories in Experiment 1; P TPJ-M consisted of contiguous voxels near the temporo-parietal junction 0.0001, uncorrected. Medium gray indicates fi xation. Time course aver- that responded signi fi cantly more to theory of mind (ToM) stories than to aged over four runs. mechanical inference (MI) stories ( P  0.0001, uncorrected). (Response magnitudes for the conditions that were used to de ne the regions of fi interest are illustrative only.) The EBA did not respond to story stimuli. The right TPJ-M differentiated between pictures of bodies and of objects cial stimuli appeared to be lateralized. The left TPJ-M P theory 0.05, paired samples t test), but the left TPJ-M did not. ToM  (  response was selective for verbal descriptions, while the   mechanical inference stories, Body of mind (false belief) stories; MI right TPJ-M activation may generalize to nonverbal stimuli,  photographs of human bodies, Obj photographs of nonhuman objects; such as photographs. EBA, extrastriate body area.

5 1842 – Rapid Communication / NeuroImage 19 (2003) 1835 1839 stories (Zaitchik, 1990), which require subjects to tograph ” question (4 s)]. There were two blocks per condition per run. represent the (false) content of a physical representation Fixations of 12 s were interleaved between blocks. The such as a photograph or map. ve new sets of order of conditions was counterbalanced across runs. Be- fi For Experiment 2, we therefore created stories (for examples, see Appendix 2): (1) false belief havioural data were collected during the scan. stories, (2) false photograph stories, (3) desires, (4) inani- mate descriptions, and (5) physical people. Desire stories Results described a character ’ s goals or intentions and thus rely on ToM. Nonhuman description stories consisted of short de- Subjects were slower when responding to questions scriptions of nonhuman objects such as plants, cars, or about false photograph than false belief stories (FB: 2.6 vs. planets. Physical people stories were short descriptions of people from a purely physical perspective: clothing, hair FP: 2.8 s, P  0.01), making it unlikely that false belief cult. fi inferences were simply more dif colour, facial markings, and so on. cally involved in ToM As predicted, a random effects analysis on the 21 sub- fi We predicted that regions speci jects who underwent whole brain scanning revealed regions should have a equally low response in the nonhuman de- of increased BOLD signal to false belief compared with scription and (critically) physical people conditions, and a false photograph stories ( P  higher BOLD response in the desire condition. By contrast, 0.0001, uncorrected) at the ), left: ( 18 51  TPJ bilaterally [right: (54 63 33)],  regions involved in processing any other representation of  48 other people would show a high BOLD signal for the precuneus/posterior cingulate [(3  54 30)], right anterior superior temporal sulcus [(54 physical people condition. 18  15)], and in medial  superior frontal gyrus [(6 57 18)] in the frontal pole (Fig. 2). Medial prefrontal cortex has repeatedly been implicated in Methods ToM processing, both in neuroimaging and in lesion studies (e.g., Rowe et al., 2001; Stuss et al., 2001). For the 7 subjects who were scanned in both Experi- Twenty-one naive right-handed subjects (11 women) ments 1 and 2, two additional analyses were conducted to were scanned at 1.5 T, using twenty 5-mm-thick axial slices con fi rm that the TPJ-M was consistent across experiments. that covered the whole brain. An additional 7 subjects from Experiment 1 (4 women) also participated in part of Exper- First, in all 7 subjects the TPJ-M de fi ned in Experiment 1 ned by the contrast of fi overlapped strikingly with TPJ-M de iment 2. All were scanned at 3.0 T using twenty 5-mm-thick false belief (FB) versus false photograph (FP) stories in near-coronal slices (parallel to the brainstem) covering most of the occipital lobe and the posterior portion of the tem- Experiment 2. Fig. 3a shows the overlap in a typical indi- ned by these two tasks. fi vidual subject of the TPJ-M de poral and parietal lobes. rmed with a functional ROI fi Second, this overlap was con Story stimuli consisted of 70 stories (12 each of false belief, false photograph, desire, physical description, and analysis. Voxels near the TPJ are more active during ToM nonhuman description, average number of words  32; see than mechanical inference stories in these individual sub- 0.0001, uncorrected) were jects in Experiment 1 ( P Appendix 2). After each story a two-alternative forced  probed for their response during Experiment 2. This inde- “fi choice question was presented for 4 s. ” ll-in-the-blank The question consisted of a single sentence with a word pendent ROI showed a much greater response to false belief missing, presented above two alternative completions on the than false photograph stories in Experiment 2 (mean FB t 0.7,  1.6, mean FP PSC  PSC left and right side of the screen. Subjects pressed the left- 0.02; Fig. 3b).  P test The reliability of the TPJ-M across experiments makes it hand response button if the word on the left completed the unlikely that the results of Experiment 1 were the result of sentence to fi t the story, and the right-hand button to choose the word on the right. Fifty percent of the false belief, false stimulus confounds or logical differences between conditions. ve conditions, the fMRI photograph, and desire story questions probed the charac- fi For the 14 subjects who saw all ter ’ s mental states; the other 50% probed the actual out- data were further analyzed within individually de fi ned func- tional regions of interest (ROI) that included all voxels that come, to prevent formulaic response preparation. Subjects cantly more active in met two criteria, i.e., they were signi were given three practice trials before going into the scan- fi ner: two false belief trials, and one false photograph trial. at least half of the individual subjects during false belief Fourteen subjects (including the 7 from Experiment 1) than false photograph stories ( P  0.0001, uncorrected), were tested on only false belief and false photograph stories. and they fell within a sphere of 15-mm radius centered on For these subjects, each run lasted 204 s and consisted of six cant voxel of clusters identi fi ed in the ran- the most signi fi blocks [each containing 1 story (10 s) and 1 question (4 s)], dom effects group analysis ( P  0.0001, uncorrected) of the ed ROIs in alternating between the two conditions; there were three same contrast. Using these criteria, we identi fi the TPJ-M bilaterally and right aSTS. blocks per condition per run. The remaining 14 subjects ve conditions. Each run lasted 272 and In the TPJ-M and the right aSTS, the BOLD signal were tested on all fi change during desire stories was signi fi cantly greater than consisted of 10 blocks [each containing 1 story (10 s) and 1

6 1842 1840 – Rapid Communication / NeuroImage 19 (2003) 1835 culty or stimuli, and is not merely an effect of the dif fi logical structure of false belief stories, since the TPJ-M did cult and logically similar false fi not respond to the more dif photograph stories. We asked whether the TPJ-M represents the simple pres- ence of another person (possibly via detecting a human body and/or biological motion) or is involved speci fi cally in ToM. We found that the TPJ-M was anatomically and functionally distinct from the nearby EBA (Downing et al., 2001), which responded preferentially to the visual appearance of human bodies, suggesting the presence of at least two distinct regions involved in social information processing. Fig. 4. Experiment 2. Average percent signal change in left and right A key innovation of this study over previous studies was 14) as voxels that respond  n ned in individual subjects ( fi TPJ-M, de the inclusion in Experiment 2 of physical people stories, signi P cantly more to false belief (FB) than to false photo (FP) stories (  fi 0.0001, uncorrected. Response magnitude for these two conditions is il- which described the physical appearance of human bodies. lustrative only, since these data were used to determine the region of Previous studies (Fletcher et al., 1995; Gallagher et al., interest). In the TPJ-M bilaterally the BOLD response to physical people 2000) have included physical “ stories describing acting ” P 0.05), and not  cantly lower than to desire stories ( fi stories was signi people, which produced greater activation in the TPJ than a cantly different from nonhuman description stories ( fi 0.1,  signi P scrambled sentence control. Our data show that the TPJ-M repeated-analysis of variance). Response decreases are commonly ob- served in the TPJ vicinity during demanding nonsocial tasks (Shulman et response was no greater to stories that described other al., 1997; Gusnard and Raichle, 2001). people in physical detail than that to stories describing the physical details of nonhuman objects fi and was signi — - cantly lower than to stories that did invite a mental state during either physical people or nonhuman description sto- interpretation (desire stories). t P  0.05), which did not tests ries (both paired samples fl Could the TPJ-M activation re ect mental imagery of the differ from each other (Fig. 4). The left and right TPJ-M did biological motion or goal-directed action described in the not differ. Thus, these regions are not involved in the de- false belief, human action, and desire stories? We think this tection of any person in verbal stories, but respond selec- is unlikely. Saxe, R., Xiao, D.K., Kovacs, G., Perrett, D., ’ tively to stories in which describe (or imply) characters and Kanwisher, N. (unpublished data) found that the TPJ-M le of fi mental states. Did any regions show the predicted pro response to a movie of a walking person was much lower a response to a person per se? At a lower threshold, a than its response to false belief stories. If the response of the  separate whole brain analysis ( 0.001, uncorrected) of P TPJ-M to verbal stories was merely a consequence of sub-  physical people nonhuman descriptions revealed regions imagining biological motion, we would predict the ’ jects  3 57 39), and of frontal cortex [dorsal medial prefrontal ( opposite. Also the TPJ-M was doubly dissociated from its right lateral frontal cortex (39 15 54)]. neighbour, the pSTS-VA (visual analysis of action), which responded more to the movies than to verbal stories. 2 is In all, our results show that a region of the TPJ Discussion involved in reasoning about other minds, not just in under- standing stories involving people per se (Gallagher and fi The results of Experiment 2 con rm that the TPJ-M Frith, 2003; p 80). But critically, neighbouring subregions shows an increased response to stimuli that invite ToM fi les, highlighting the of cortex have different functional pro reasoning compared with logically similar nonsocial con- necessity of careful within-subject comparisons. The trols (false photograph stories). Second, the TPJ-M does not TPJ-M, identi fi ed here by responses to (false) belief stories, show an increased response to the mere presence of a person in the stimulus (physical people stories). The right and left 2 TPJ-M responses to physical people stories did not differ, What is the relationship between the TPJ-M and attention? Selective thus resolving the ambiguity of the apparently lateralized attention leads to increases in regions of the TPJ during social perception tasks (e.g., Narumoto et al., 2001; Winston et al., 2002), and to decreases response to photographs of bodies in Experiment 1. in regions of the TPJ during visual attention tasks (Shulman et al., 1997; Gusnard and Raichle, 2001; Jiang Y., Kanwisher, N., unpublished data). General discussion Downar et al. (2001) proposed “ a role for the TPJ in detecting behav- iourally relevant events in the sensory environment ” (p. 1256) that is interfered with by demanding visual attention. One possibility is that the In two experiments we found greater BOLD response in mental states of other people constitute a particular category of such a region within the TPJ bilaterally (here called TPJ-M) “ ect behaviourally relevant ” stimuli. Alternatively, these results may re fl while subjects read stories that describe or imply a charac- functionally and anatomically distinct subregions within the TPJ. Direct s goals and beliefs than during stories about nonhuman ’ ter testing of the relationship between the TPJ-M and selective attention is an objects. This pattern is robust across subjects, tasks, and important avenue for future work.

7 1842 – Rapid Communication / NeuroImage 19 (2003) 1835 1841 may play a broad role in social and even moral cognition False belief (FB) sample story (Moll J et al., 2002; Greene and Haidt, 2003). John told Emily that he had a Porsche. Actually, his car is a Ford. Emily doesn ’ t know anything about cars though, so she believed John. Acknowledgments — ’ s car she When Emily sees John This work was funded by grants NEI 13455 and NIHM thinks it is a 66696. Our thanks especially to Yuhong Jiang for com- porsche ford ments and conversation, and to Ben Balas, Robb Rutledge, Miles Shuman, and Amal Dorai for help with data collection False photograph (FP) sample story and analysis. A photograph was taken of an apple hanging on a tree branch. The fi lm took half an hour to develop. In the meantime, a strong Appendix wind blew the apple to the ground. — Experiment 1 The developed photograph shows the apple on the ground branch Instructions: “ Read each story silently to your self. Desire sample story Please make sure you understand what is happening; it is more important that you understand the story, than that you For Susie s birthday, her parents decided ’ go as fast as possible. When you are done reading the story, to have a picnic in the park. They wanted ponies and games on the lawn. If it rained, press the button. ” the children would have to play inside. — Theory of mind (ToM) sample story ’ s parents wanted to have her birthday Susie A boy is making a paper mache project inside outside for his art class. He spends hours ripping newspaper into even strips. Physical people sample story fl our. His Then he goes out to buy Emily was always the tallest kid in her mother comes home and throws all the class. In kindergarten she was already newspaper strips away. over 4 feet tall. Now that she is in  4  . She is a head taller college she is 6 Mechanical inference (MI) sample story than the others. — A pot of water was left on low heat In kindergarten Emily was over yesterday in case anybody wanted tea. 4ft 6ft The pot stayed on the heat all night. . . .tall Nobody did drink tea, but this morning, the water was gone. Nonhuman description sample story Nine planets and their moons, plus various Human action sample story lumps of debris called asteroids and Jane is walking to work this s solar system. comets, make up the sun ’ morning through a very industrial The earth is one of four rocky planets area. In one place the crane is in the inner solar system. taking up the whole sidewalk. To — get to her building, she has to The solar system has take a detour. nine four . . .planets. Experiment 2 References Please read each story carefully. After Instructions: “ fi each story, you will be given one ll-in-the-blanks question Allison, T., Puce, A., et al., 2000. Social perception from visual cues: role about the story. Underneath will be two words that could fi ll 278. of the STS region. Trends Cogn. Sci. 4, 267 – in the blank. Choose the correct word (to make the sentence Bartsch, K., Wellman, H., 1995. Children Talk about the Mind. Oxford true in the story) by pressing the left button to choose the University Press, New York. left-hand word, and the right button to choose the right-hand Brunet, E., Sarfati, Y., et al., 2000. A PET investigation of the attribution 166. of intentions with a nonverbal task. Neuroimage 11, 157 word. ” –

8 1842 – Rapid Communication / NeuroImage 19 (2003) 1835 1842 Carey, S., 1985. Conceptual Change in Childhood. MIT Press, Cambridge, ” Theory of mind “ Leslie, A., 1999. as a mechanism of selective attention, in: Gazzanigal, M. (Ed.), The New Cognitive Neurosciences. MIT MA. Carruthers, P., Smith, P., 1996. Theories of Theories of Mind. Cambridge Press, Cambridge, MA. University Press, New York. Malle, B.F., Moses, L.J., Baldwin, D.A. (Eds.), 2001. Intentions and Intentionality: foundations of Social Cognition. MIT Press, Cambridge, Castelli, F., Happe, F., et al., 2000. Movement and mind: a functional MA. imaging study of perception and interpretation of complex intentional Moll, J., et al., 2002. The neural correlates of moral sensitivity: a functional movement patterns. Neuroimage 12, 314 – 325. Dennett, D., 1978. Beliefs about beliefs. Behav. Brain Sci. 1, 568 570. magnetic resonance imaging investigation of basic and moral emotions. – – Dennett, D., 1996. Kinds of Minds: Toward an Understanding of Con- 2736. J. Neurosci 22, 2730 sciousness. Basic Books New York, NY. Narumoto, J., Okada, T., et al., 2001. Attention to emotion modulates fMRI activity in human right superior temporal sulcus. Brain Res. Cogn. Downar, J., Crawley, A.P., et al., 2001. The effect of task relevance on the 231. – Brain Res. 12, 225 cortical response to changes in visual and auditory stimuli: an event- Rowe, A.D., Bullock, P.R., et al., 2001. ” Theory of mind “ 1267. impairments and related fMRI study. Neuroimage 14, 1256 – Downing, P.E., Jiang, Y., et al., 2001. A cortical area selective for visual their relationship to executive functioning following frontal lobe exci- sions. Brain 124 (Pt 3), 600 616. 2473. – processing of the human body. Science 293, 2470 – Ferstl, E.C., von Cramon, D.Y., 2002. What does the frontomedian cortex Saxe, R., Xiao, D.K., et al., 2003. Distinct representations of bodies, actions and thoughts in posterior superior temporal sulcus, submitted. contribute to language processing: coherence or theory of mind? Neu- Shulman, G.L., Corbetta, M., et al., 1997. Top-down modulation of early roimage 17, 1599 – 1612. sensory cortex. Cereb Cortex 7, 193 – 206. Fletcher, P.C., Happe, F., et al., 1995. Other minds in the brain: a func- in story comprehension. Stuss, D.T., Gallup Jr., G.G., et al., 2001. The frontal lobes are necessary ” tional imaging study of “ theory of mind theory of mind. – – 128. “ Cognition 57, 109 ” Brain 124 (Pt 2), 279 286. for Tong, F., Nakayama, N., et al., 2000. Response properties of the human Frith, C.D., Frith, U., 1999. Interacting minds — a biological basis. Science 1695. – fusiform face area. Cogn. Neuropsychol. 17, 257 – 279. 286, 1692 Gallagher, H.L., Frith, C.D., 2003. Functional imaging of theory of “ Vogeley, K., Bussfeld, P., et al., 2001. Mind reading: neural mechanisms of theory of mind and self-perspective. Neuroimage 14 (1 Pt 1), 83. Trends Cogn. Sci. 7, 77 ” mind. – Gallagher, H.L., Happe, F., et al., 2000. Reading the mind in cartoons and 170 – 181. ” “ stories: an fMRI study of theory of mind Wellman, H.M., Gelman, S., 1992. Cognitive development: foundational in verbal and nonverbal 21. theories of core domains. Annu. Rev. Psychol. 43, 337 – 375. tasks. Neuropsychologia 38, 11 – Greene, J., Haidt, J., 2003. How (and where) does moral judgement work? Wimmer, H., Perner, J., 1983. Beliefs about beliefs: representation and constraining function of wrong beliefs in young children – s understand- ’ Trends Cogn. Sci. 6, 517 523. Grossman, E., Donnelly, M., et al., 2000. Brain areas involved in percep- ing of deception. Cognition 13, 103 – 128. 720. tion of biological motion. J. Cogn. Neurosci 12, 711 – Winston, J.S., Strange, B.A., et al., 2002. Automatic and intentional brain Gusnard, D.A., Raichle, M.E., 2001. Searching for a baseline: functional responses during evaluation of trustworthiness of faces. Nat. Neurosci. 5, 277 imaging and the resting human brain. Nat. Rey. Neurosci. 2, 685 694. 283. – – Hoffman, E.A., Haxby, J.V., 2000. Distinct representations of eye gaze and ict with reality: the pre- fl Zaitchik, D., 1990. When representations con ’ s problem with false beliefs and ” false schooler photographs. Cogni- identity in the distributed human neural system for face perception. “ – 68. – tion 35, 41 84. Nat. Neurosci. 3, 80

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