Social Cognitive and Affective Neuroscience Advance Access originally published online on August 7, 2008
Social Cognitive and Affective Neuroscience 2008 3(3):279-289; doi:10.1093/scan/nsn023
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Differential involvement of the posterior temporal cortex in mentalizing but not perspective taking
1Department of Psychiatry and Psychotherapy, University of Cologne, Cologne, 2Department of Neurophysiology and Pathophysiology, University Medical Center Hamburg-Eppendorf, Hamburg, 3Department of Psychiatry and Psychotherapy, University Hospital Bonn, Bonn, Germany, 4C. & O. Vogt Institute for Brain Research, University of Düsseldorf, Düsseldorf, 5Institute of Neuroscience and Biophysics, Cognitive Neurology Section, Research Centre Jülich, Jülich, Germany, 6Brain Imaging Center West, Research Center Jülich, Jülich, 7Department of Philosophy, University of Bochum, Bochum and 8Department of Neurology, University of Cologne, Cologne, Germany
Correspondence should be addressed to Nicole David, Department of Neurophysiology and Pathophysiology, University Medical Center Hamburg-Eppendorf, Martinistr. 52, 20246 Hamburg, Germany. E-mail: ndavid{at}uke.uni-hamburg.de.
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ABSTRACT |
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 TOP ABSTRACT MATERIALS AND METHODSRESULTSDISCUSSIONSUPPLEMENTARY DATAREFERENCES |
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Understanding and predicting other people's mental states and behavior are important prerequisites for social interactions. The capacity to attribute mental states such as desires, thoughts or intentions to oneself or others is referred to as mentalizing. The right posterior temporal cortex at the temporal–parietal junction has been associated with mentalizing but also with taking someone else's spatial perspective onto the world—possibly an important prerequisite for mentalizing. Here, we directly compared the neural correlates of mentalizing and perspective taking using the same stimulus material. We found significantly increased neural activity in the right posterior segment of the superior temporal sulcus only during mentalizing but not perspective taking. Our data further clarify the role of the posterior temporal cortex in social cognition by showing that it is involved in processing information from socially salient visual cues in situations that require the inference about other people's mental states.
Keywords: posterior superior temporal sulcus; temporo–parietal junction; mentalizing; theory of mind; perspective taking; social cognition
Understanding what other people around us perceive, feel or think is important for predicting their behavior and vital to our social interactions as well as functioning in society. To this end, we take other people's perspective to judge what they perceive from their position in space in contrast to our own position (Piaget and Inhelder, 1967; Flavell et al., 1981; Newcombe, 1989; Vogeley and Fink, 2003; David et al., 2006) or try to make inferences about other people's mental states and find out what they think, believe or desire. This latter process has also been referred to as ‘mentalizing’ (Frith and Frith, 2006), ‘mind-reading’ or ‘theory of mind’ (Premack and Woodruff, 1978; Baron-Cohen, 1997). In contrast to mentalizing, perspective taking does not require an ascription of mental states to another person but rather an assessment of another person's visuospatial experiences. Nevertheless, perspective taking may be associated with mentalizing: ‘This ability to see the world from another's perspective enables us to realize that other people can have different knowledge from us and may have false beliefs about the world’ (Frith and Frith, 2006, p. 532).
It is a matter of conjecture to which degree mentalizing and perspective taking are functionally independent processes. Evidence derived from developmental studies in children and experiments in monkeys suggests a dissociation (Flavell et al., 1978; Newcombe, 1989; Povinelli and Eddy, 1996; Meltzoff, 1999; Hare et al., 2000, 2001; Saxe et al., 2004a), whereas evidence from certain psychiatric conditions such as schizophrenia and autistic spectrum disorder showed common deficits (Yirmiya et al., 1994; Langdon and Coltheart, 2001). Surprisingly few experiments aimed to empirically compare mentalizing and perspective taking (Reed and Peterson, 1990; Langdon and Coltheart, 2001). Neuroimaging may help to elucidate the functional basis and relationship between mentalizing and perspective taking. Mentalizing has already been extensively investigated by functional magnetic resonance imaging (fMRI) experiments (Castelli et al., 2000; Gallagher et al., 2000, 2002; Vogeley et al., 2001; Gallagher and Frith, 2003; Saxe and Kanwisher, 2003; Ruby and Decety, 2003, 2004; Rilling et al., 2004; Saxe and Wexler, 2005; Völlm et al., 2006; Schulte-Rüther et al., 2007) as opposed to perspective taking (Zacks et al., 2003; Vogeley et al., 2004; Blanke et al., 2005; Aichhorn et al., 2006; Arzy et al., 2006; David et al., 2006; Jackson et al., 2006).
The fMRI studies on mentalizing converge on the recruitment of the medial prefrontal cortex (MPFC) as well as temporal cortex, specifically the superior temporal sulcus (STS), temporal–parietal junction (TPJ) and the temporal poles (Frith and Frith, 1999, 2006; Gallagher and Frith, 2003). These regions have, thus, been referred to as the neural ‘mentalizing network’ (Frith and Frith, 2003) or ‘human social cognition network’ (Saxe, 2006). Perspective taking studies (involving viewer rotations or egocentric perspective transformations; Vogeley and Fink, 2003) often reported activations of superior parietal and inferior parietal cortices, inferior frontal cortex and the cerebellum (Zacks et al., 1999; Vogeley et al., 2004; David et al., 2006; Jackson et al., 2006). Interestingly, the posterior STS at the TPJ was recruited both during mentalizing (Brunet et al., 2003; Gallagher et al., 2000; Ruby and Decety, 2003; Völlm et al., 2006; Schulte-Rüther et al., 2007) and perspective taking (Zacks et al., 1999, 2003; Vogeley et al., 2004; Blanke et al., 2005; Aichhorn et al., 2006; Arzy et al., 2006; David et al., 2006; Jackson et al., 2006).
Aichhorn and colleagues (2006) recently addressed the question whether perspective taking activates regions within the posterior STS region (or ‘STS/TPJ’ as referred to by Aichhorn et al., 2006) that have previously been associated with mentalizing (the authors used the term ‘theory of mind’). They found the ‘STS/TPJ’ to be recruited during perspective taking, however, only after a restricted search volume or region-of-interest (ROI) approach. Although Aichhorn and colleagues (2006) attempted to compare activation in the posterior temporal cortex during mentalizing vs perspective taking, they did not make a direct empirical comparison in the same set of subjects with comparable stimulus material. In fact, to date, no direct comparison between the neural correlates underlying the two kinds of perspective taking has been presented. Thus, the questions whether the posterior temporal cortex contributes to both mentalizing and perspective taking and on the exact nature of this contribution—by the processing of social, mental state conveying signals or by representation of different visual perspectives (Frith and Frith, 2006)—remain.
Thus, we sought to directly compare activation between mentalizing and perspective taking using fMRI and to specifically test whether and to which degree both processes recruit the same neural substrates, especially with respect to the posterior temporal cortex. Operationally, we defined mentalizing as the process of ascribing a preference for one of two objects to a virtual character. Mental states can be expressed in many different ways: here, we focused on non-verbal, socially salient signals expressed by the virtual character's face or body. Facial and other bodily cues represent relevant sources of information about somebody's inner states (Frith and Frith, 2006) and are often used in operationalizations of mentalizing (Baron-Cohen et al., 2001; Gallagher and Frith, 2004; Schulte-Rüther et al., 2007). Perspective taking was operationalized using the established own-body mental transformation task (Parsons, 1987; Zacks et al., 1999; Blanke et al., 2005; Arzy et al., 2006). Our results suggest that mentalizing and perspective taking can be dissociated within the posterior segment of the right STS with a stronger involvement of the STS in mentalizing for another person.
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MATERIALS AND METHODS |
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TOPABSTRACTMATERIALS AND METHODSRESULTSDISCUSSIONSUPPLEMENTARY DATAREFERENCES |
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Subjects
Seventeen subjects without psychiatric or neurological history participated in this study after having given informed written consent. Four subjects were later excluded due to poor behavioral performance. All analyzed subjects (N = 13; age 25 ± 3.5 years) were right-handed, as assessed by the Edinburgh Handedness Inventory (Oldfield, 1971). We studied only male volunteers based on evidence for sex differences in visuospatial and mentalizing tasks with respect to behavioral performance and hemispheric engagement (Parsons et al., 2004; Baron-Cohen et al., 2005). The study was approved by the ethics committee of the University of Cologne.
Design, stimuli and task
The present study employed a 2 x 2 factorial design with the factors Reference (first-person reference, 1P vs third-person reference, 3P) and Task (perspective taking, PERSP; mentalizing, MENT). Stimuli depicted a vis-à-vis positioned virtual character surrounded by two different objects of the same category (e.g. fruit: an apple and a banana; Figure 1). Object categories were randomly chosen. The two objects showed a subtle difference in position, with one object being slightly elevated compared with the other object. Virtual characters expressed preferences to one of the two objects. These preferences were indicated by systematically varied combinations of different facial expressions (positive, negative and neutral), gestures (positive, negative and no gesture) and head/body positions (turned towards or away from the object, neutral position; Supplementary Table S1). The salience of the character's preference depended on the congruence of these cues, which was not homogeneous throughout. Thus, ceiling effects and simple rule-based behavioral strategies were avoided (e.g. preferences could not be determined by paying attention to the character's face alone). Virtual characters were created and used in a previous project (Bewernick et al., 2005, meeting abstract), in which subjects performed certainty ratings on the virtual characters’ preference. For the purpose of the present experiment, stimuli associated with high uncertainty for preference judgments were excluded (Supplementary Table S1 shows the present distribution of social cue combinations).
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Subjects performed two different tasks from two different reference points. In a perspective taking task (PERSP), subjects indicated the elevated object from their own or first-person perspective (1P-PERSP) or the same from the virtual character's perspective or third-person (3P-PERSP). During the 3P-PERSP condition, subjects were instructed to do a mental transformation of their own, egocentric perspective onto the virtual character's body axis (Vogeley and Fink, 2003) similar to the ‘own-body mental transformation task’ (as used by Blanke et al., 2005; Parsons, 1987; Zacks et al., 1999) and were debriefed about that after the study. The 3P-PERSP condition (i.e. a viewer rotation) is different from mental rotation (i.e. an object rotation; Langdon and Coltheart, 2001; Zacks et al., 2003) and no subjects reported to have performed such a mental rotation of the scene. In the mentalizing task (MENT), subjects indicated the object they preferred themselves (condition 1P-MENT) or the one the virtual character preferred (condition 3P-MENT) based on its gestures, facial and bodily expressions. Subjects were instructed to follow a global impression of the virtual character's preference (i.e. not only pay attention to the face or gesture). During 3P-MENT subjects indicated the object as seen from their own perspective in order to avoid a confound with 3P-PERSP. All conditions involved left–right judgments by pressing a button with the index/middle finger of the left or right hand on an magnetic resonance imaging (MRI)-compatible button box (Lumitouch, Lightwave Medical Industries, CST Coldswitch Technologies, Richmond, CA, US). During 3P-PERSP, subjects were instructed to answer as if they were looking through the eyes of the avatar; thus, if a target object was to the right side of the avatar, subjects had to press the right button. Hands were counterbalanced across sessions and subjects. The present operationalization had the advantage of testing mentalizing and perspective taking without changing visual input: stimuli were the same for all four conditions which differed only by their instructions (Figure 1). In other words, facial or bodily expressions of the avatars were identical during MENT and PERSP trials.
Stimuli were designed using the commercially available software Poser (Version 6, Curious Labs, Inc., Santa Cruz, CA, US), Photo Objects (Vol. 1, Hemera Technologies, Inc., Gatineau, Quebec, Canada) and Adobe Photoshop CS (Version 8.0.1, Adobe Systems, Inc., San Jose, CA, US). In total, there were nine different male virtual characters, each placed in front of a neutral white background. The use of virtual stimuli in social cognitive neuroscience or psychology has proven to be useful as virtual characters are perceived as social agents evoking similar responses or impressions in observers as human stimuli (Bente et al., 2001; Schilbach et al., 2006). The side of the elevated object and the side of preference of the virtual character were counterbalanced as much as possible and systematically varied.
The experiment was conducted as a block design with four sessions (
8 min each), the order of which was randomized between subjects. The four experimental conditions were repeated twice per session (eight experimental blocks in total) and alternated with baseline blocks (cross-hair for 20 s). Each session contained 72 stimuli (nine per experimental block). Each stimulus was displayed for 4245 ms followed by a 200 ms inter-stimulus interval. Instructions were shown for 5 s at the beginning of each block. The experiment was programmed and presented on a light gray background using Presentation (Version 9.13, Neurobehavioral Systems, Albany, CA, US).
Analyses of behavioral data
Statistical analyzes were performed using the Statistical Package for the Social Sciences (SPSS for Windows; Version 12.0). Dependent variables were reaction times (RT) and accuracy (percentage of correct responses). Accuracy was not determined for 1P-MENT as subject's responses reflected subjective preferences in this condition. For 3P-MENT, accuracy for stimuli with relative ambiguity was estimated based on which reply the majority of subjects had given. Means and standard errors of the mean (s.e.m.) are reported. RT differences between conditions were analyzed using a two-way repeated-measures analysis of analysis of variance (ANOVA) with Reference (1P vs 3P) and Task (MENT vs PERSP) as within-subject factors. Differences in accuracy were tested for significance non-parametrically (Friedman test). The significance level for all analyses was set at P < 0.05 two-tailed.
Image acquisition
Functional and structural MRI was performed on a Siemens 3T MRI whole body scanner (SIEMENS Trio) using a standard head coil and a custom-built head holder to reduce head movement. Functional images were obtained using a single-shot gradient echo, echoplanar imaging (EPI) sequence (TR: 2500 ms, TE: 30 ms, 90° flip angle, FOV: 200 mm, matrix: 64 x 64, voxel size: 3.1 x 3.1 x 3 mm). Each EPI volume contained 40 axial slices (distance factor: 10), acquired in ascending order, covering the whole brain. Each session contained 193 functional images. In addition, a high-resolution T1-weighted magnetization-prepared rapid gradient-echo imaging (MP-RAGE) 3D MRI sequence was acquired from each subject (TR: 2250 ms, TE: 3.93 ms, 9° flip angle, FOV: 256 mm, matrix: 256 x 256, voxel size: 1 x 1 x 1 mm).
Image preprocessing and analyses
Functional images were corrected for head movement between scans by an affine registration (Ashburner and Friston, 2003) using the Statistical Parametric Mapping software (SPM5; Wellcome Department of Imaging Neuroscience, London, UK) implemented in Matlab (Mathworks Inc., Sherborn, MA, USA). To allow localization of functional activation on the subjects’ structural MRIs, T1-scans were coregistered to each subject's mean image of the realigned functional images. The mean functional image was subsequently normalized to the Montreal Neurological Institute (MNI) single-subject template (Evans et al., 1992; Collins et al., 1994) using linear proportions and a non-linear sampling as derived from a segmentation algorithm (Ashburner and Friston, 2005). Normalization parameters were then applied to the functional images and coregistered to the T1-image. Images were resampled at a 2 x 2 x 2 mm voxel size and spatially smoothed using a 8 mm full width half maximum Gaussian kernel. The data were analyzed using a general linear model (Kiebel and Holmes, 2003). Each experimental condition was modeled using a boxcar reference vector convolved with a canonical hemodynamic response function and its first-order temporal derivative. Low-frequency signal drifts were filtered using a cutoff period of 128 s. Parameter estimates were subsequently calculated for each voxel using weighted least squares to provide maximum likelihood estimates based on the non-sphericity assumption of the data (Kiebel and Holmes, 2003) in order to get identical and independently distributed error terms. For each subject, main effects were computed by applying appropriate baseline contrasts (simple effects). These first-level individual contrasts were then fed into a second-level group analysis using an ANOVA (factor: condition, blocking factor subject), thus employing a random-effects model (Penny and Holmes, 2003).
We were interested, first, in comparing mentalizing with a third person as opposed to first-person reference (3P-MENT vs 1P-MENT) and, second, perspective taking with a third person as opposed to a first-person reference (3P-PERSP vs 1P-PERSP) (Table 1, for the reader's convenience, also contains the opposite contrasts 1P-MENT vs 3P-MENT and 1P-PERSP vs 1P-PERSP). Third, we aimed for the direct comparison of mentalizing and perspective taking especially with respect to a third-person reference (3P-MENT vs 3P-PERSP; Table 1). Common activations of third-person mentalizing and third-person perspective taking were identified via a conjunction analysis using the Global Null, a less conservative approach testing the combined null hypothesis for both contrasts of interest (3P-MENT vs 1P-MENT and 3P-PERSP vs 1P-PERSP; Table 2; Price and Friston, 1997; Friston et al., 2005). To investigate the interaction between the factors Reference and Task (e.g. for the combination of the factor levels 3P and MENT), and to formally test differences in simple effects, the following interaction contrast was calculated: [(3P-MENT vs 1P-MENT) vs (3P-PERSP vs 1P-PERSP)]. Note that main effects were not of primary interest because they are collapsing across conditions of interest (Supplementary Table S1).
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All activations were identified using a height threshold of P < 0.05, family-wise corrected (FWE), and an extent threshold of k = 10 contiguous voxels. Resulting activation peaks were superimposed on a normalized high-resolution MP-RAGE image (averaged across subjects) and anatomically localized using an atlas of the human brain (Duvernoy, 1999). For a priori expected activations, ROI analyses were carried out based on previously published coordinates. Such an analysis is restricted to a smaller number of voxels so that the number of multiple comparisons is reduced (Eickhoff et al., 2006), increasing power to detect an effect compared with a whole-brain analysis (reducing potential Type II errors). For example, the hypothesis on posterior temporal activation during the interaction of Reference and Task (e.g. the difference between 3P and 1P is bigger for MENT than PERSP) was tested with a spherical ROI (10 mm radius; Figure 2A), which was generated using Marsbar and centered around averaged, previously reported coordinates (Castelli et al., 2000; Gallagher et al., 2000; Saxe and Kanwisher, 2003; Gallagher and Frith, 2004: x, y, z = 57 ± 4, –52 ± 4, 15 ± 5). Within this ROI, activation was tested for using the WFU Pick Atlas (Maldjian et al., 2003) as implemented in SPM5 (Figure 2B and C). Furthermore, ROIs were used to extract each subject's parameter estimates from the peak activation obtained in the whole-brain analyses for each condition as compared to baseline. These parameter estimates were then averaged across subjects and means and s.e.m. plotted (Figure 3B and D). In addition, individual parameter estimates were extracted within ROIs centered around previously published coordinates in the posterior temporal cortex associated with other social-cognitive tasks, averaged and plotted (Figure 4). If these coordinates were in Talairach and Tournoux (1988) space, they were transformed into MNI space using the algorithm tal2mni (Matthew Brett; http://imaging.mrc-cbu.cam.ac.uk/downloads/MNI2tal/tal2mni.m) implemented in Matlab.
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RESULTS |
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TOPABSTRACTMATERIALS AND METHODSRESULTSDISCUSSIONSUPPLEMENTARY DATAREFERENCES |
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Behavioral data
There was a significant interaction effect on RTs between the factors Reference and Task (F1,12 = 35.86, P < 0.001). Subjects were significantly faster when making visuospatial judgments (i.e. detecting the elevated object) from their own perspective (mean ± s.e.m. for RTs on 1P-PERSP: 997 ± 41 ms). This was indicated by pairwise comparisons [1P-PERSP vs 3P-PERSP (1548 ± 148 ms): t12 = 4.64, P < 0.001; 1P-PERSP vs 1P-MENT (1833 ± 104 ms): t12 = 1.86, P < 0.001; 1P-PERSP vs 3P-MENT (1659 ± 139 ms): t12 = 5.32, P < 0.001]. Conditions also differed with respect to accuracy (Friedman
2 = 10.6; Figure 5). Subjects were significantly more accurate on 3P-PERSP (mean ± s.e.m.: 94 ± 2% correct) vs 3P-MENT (88 ± 2% correct; Wilcoxon Z = –2.1, P < 0.05) and on 1P-PERSP (96 ± 1% correct) vs 3P-MENT (Z = –3.0, P < 0.01). Nothing could be inferred about an interaction of Reference and Task on accuracy because subjects’ replies during 1P-MENT reflected subjective judgments. In postexperimental debriefings, 11 subjects reported to have primarily used the virtual character's facial expression and hand gesture in order to judge the character's preferences, one subject preferably used gesture and another subject facial expression together with body orientation as primary cues for preference judgments.
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Whole-brain analyses
Inferring the virtual character's preference compared to one's own preference (3P-MENT vs 1P-MENT) was associated with significantly (P < 0.05, FWE corrected) greater activation of the right middle frontal gyrus, right posterior STS region and the right middle temporal gyrus, bilateral supramarginal gyrus and right precuneus (Table 1; Figure 3A and B).
Taking a third person in contrast to a 1P-PERSP (3P-PERSP vs 1P-PERSP) recruited left middle/inferior frontal and right middle/superior frontal gyri, the left supramarginal gyrus and the cerebellum bilaterally (Table 1; Figure 3). In contrast to previous studies on perspective taking, this contrast did not reveal any activation in the TPJ or posterior STS.
The direct comparison of third-person mentalizing vs third-person perspective taking (3P-MENT vs 3P-PERSP) yielded a significant cluster in the right posterior STS (Table 1; Figure 3A and B).
The analysis of common activation patterns between mentalizing and perspective taking (3P-MENT vs 1P-MENT in conjunction with 3P-PERSP vs 1P-PERSP) revealed overlapping activation patterns in the inferior parietal lobule bilaterally and in the right middle frontal gyrus (BA 6; Table 2).
There was no significant activation associated with the interaction [(3P-MENT vs 1P-MENT) vs (3P-PERSP vs 1P-PERSP)].
ROI analyses
Two kinds of ROI analyses were performed: (i) expected activation, that is, during the interaction of Reference and Task, was tested for in a more restricted search volume comprising the posterior STS based on previously published coordinates, (ii) parameter estimates were extracted from presently and previously reported coordinates in order to show the strength of activation during each condition and to demonstrate the differential recruitment of the posterior temporal cortex during mentalizing vs perspective taking.
The first ROI analysis confirmed our hypothesis about an interaction Reference and Task, specifically a third-person reference and mentalizing, within the posterior STS (x = 60, y = –50, z = 13; Z = 3.79; FWE P < 0.05; 60 voxel; Figure 2).
Parameter estimates extracted at the afore-mentioned peak coordinates within the posterior temporal cortex for each of the four experimental conditions revealed that only 3P-MENT evoked a strong activation within the posterior STS, while the remaining three conditions were associated with activation levels under baseline (Figure 3B and E), being also suggestive of an interaction effect. Indeed a repeated measures ANOVA on these parameter estimates yielded a significant interaction between Reference and Task for both peaks (F1,12 = 7.48, P < 0.05 for x, y, z = 56, –32, 3; F1,12 = 15.4, P < 0.005 for x, y, z = 64, –52, 1), driven by the 3P-MENT condition (pairwise comparisons of 3P-MENT against 1P-MENT, 3P-PERSP and 1P-PERSP: all t12 > 3.6; all P < 0.001).
Plotting the effects of our task in previously published coordinates in the posterior temporal cortex (reported as ‘posterior STS’ or ‘TPJ’) associated with mentalizing (Saxe and Kanwisher, 2003; Figure 4A) or perspective taking (or own-body mental transformations, Blanke et al., 2005; Figure 4B), and related processes such as the recognition of expressive gestures (Gallagher and Frith, 2004; Figure 4C), the observation of intentional human actions (Saxe et al., 2004b; Figure 4D), the perception of expressive faces (Narumoto et al., 2000; Figure 4E) and of biological motion (Bonda et al., 1996; Figure 4F) also revealed the highest activation during 3P-MENT compared with all other conditions.
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DISCUSSION |
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TOPABSTRACTMATERIALS AND METHODSRESULTSDISCUSSIONSUPPLEMENTARY DATAREFERENCES |
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The two main objectives of the present experiment were to compare mentalizing and perspective taking in a single sample using identical stimulus material and to evaluate the role of the posterior temporal cortex for the two processes. We found distinct activation of the posterior segment of the STS during mentalizing but not perspective taking. More specifically, the right posterior STS was significantly activated when subjects made inferences about a virtual character's preference as indicated by its facial or bodily expressions.
The human posterior STS region has been described as the cortex within the STS as well as adjacent superior or middle temporal and angular gyri (e.g. Pelphrey et al., 2004a). This definition goes beyond the posterior segment of the STS per se and blurs the anatomical border to the TPJ, which can be assumed to be located more dorsally. In fact, the posterior STS and TPJ are often referred to synonymously (Castelli et al., 2000; Aichhorn et al., 2006) despite reports on a functional differentiation (Saxe, 2006). Here, we found the posterior segment of the sulcus itself—more inferior to the TPJ—to be significantly activated during mentalizing, that is, when subjects inferred a virtual character's preference based on non-verbal social cues expressed by the character. Several authors have associated activity in the posterior STS with tasks as different as the detection of animate entities (Castelli et al., 2000; Blakemore et al., 2003; Schultz et al., 2004, 2005), a non-verbal mentalizing task (Brunet et al., 2003), perception of expressive faces (Narumoto et al., 2000) or expressive gestures (Gallagher and Frith, 2004). Often tasks that activate the posterior STS, including our own, used stimuli that involved the perception of social cues and the interpretation of such cues for the understanding of other people's behavior and mental states. Recent data obtained from a patient with a lesion to the posterior STS confirmed the present key finding on differential STS activation for mentalizing and perspective taking: this patient was unable to infer the virtual character's preferences (i.e. during 3P-MENT) but performed well above chance on all other conditions (Cohen et al., in press). Interestingly, intracranial recordings from the posterior STS surface in four other patients showed specifically enhanced neural oscillations in the gamma frequency range in association with the 3P-MENT condition—possibly also underlying the present fMRI activation (Cohen et al., in press).
Although perspective taking may also depend on the detection of visual social cues such as eye gaze or pointing gestures, it does not necessarily require the representation of other people's mental states such as beliefs, desires or preferences. Empirical evidence on visuospatial perspective taking is limited; yet, it was suggestive of a posterior STS/TPJ involvement during perspective transformation (Zacks et al., 1999; Vogeley et al., 2004; Blanke et al., 2005; Aichhorn et al., 2006; Arzy et al., 2006). We could not replicate the finding of increased TPJ activation during perspective taking. However, we found other areas previously associated with perspective transformations, such as the middle frontal or premotor cortex, the cerebellum and inferior parietal lobule (Ruby and Decety, 2001; Vogeley and Fink, 2003; Vogeley et al., 2004; David et al., 2006). As a potential caveat, subjects—against instructions—may have performed 3P-PERSP by first computing 1P-PERSP and then inverting the response. RT patterns, however, do not favor this alternative hypothesis: inverting a response should have been easier resulting in RTs and accuracy closer to the 1P-PERSP, which was not the case. The validity of our task was further demonstrated by postexperimental debriefings, in which subjects indicated that they actively adopted the character's perspective. A closer inspection of studies that implicated the TPJ in perspective taking are less contradictive. For example, Aichhorn et al. (2006) reported posterior temporal activation for perspective taking but only after employing an ROI approach. Zacks et al. (1999) reported activity in ‘areas near the posterior-temporal-occipital junction’, predominantly in the left hemisphere and not necessarily comprising the TPJ. Blanke and coworkers (2005; Arzy et al., 2006), however, showed quite solid electrophysiological evidence on the right TPJ's role for perspective transformations, although the authors rather propose a specific role in embodiment (not necessarily an aspect of our task). In one of our previous studies, we also reported activation extending to the posterior STS region at the TPJ for perspective taking; however, we used dynamic stimuli that depicted biological motion (David et al., 2006). Further studies are needed to reconcile such discrepant findings.
Plotting the effects of our task in superior temporal coordinates, previously associated with other social-cognitive tasks, also revealed interesting observations with respect to a differential function of posterior superior temporal cortex for mentalizing and perspective taking (Figure 4). First, similar functional profiles across the posterior STS region (including coordinates reported as ‘TPJ’ and ‘STS’) could be observed with highest activity for mentalizing (i.e. 3P-MENT). Second, activity in response to our manipulation in coordinates associated with biological motion or action observation was similar to activity in coordinates associated with the perception of expressive faces, gestures and mentalizing. Third, within almost all coordinates there was a relative increase of activity during perspective taking with a third person compared to a first-person reference (3P-PERSP vs 1P-PERSP). However, this difference in activity was small compared with the increase of activity associated with third-person mentalizing demonstrating the advantage of our design: an investigation of perspective taking alone would have revealed posterior temporal activation, however, not to the same degree as mentalizing.
It is noteworthy that a crucial contrast for showing a differential involvement of the posterior STS in third-person mentalizing vs perspective taking—the interaction effect—did not result in significant posterior STS activation at the whole-brain threshold (Schulte-Rüther et al., 2007). Several other aspects of our data, however, point to a differential involvement. First, a ROI analysis confined to the posterior STS revealed a significant interaction effect. Second, parameter estimates extracted from the posterior STS also showed a significant interaction. Third, a conjunction analysis for third-person mentalizing vs perspective taking did not reveal common posterior STS activation and, thus, also supports a differential involvement of the posterior STS for either mentalizing or perspective taking. Finally, a patient with a lesion in the posterior STS showed a selective impairment for third-person mentalizing only (i.e. during 3P-MENT; Cohen et al., in press).
It is a well-established finding that the posterior STS responds to the perception of expressive faces or biological motion (Allison et al., 2000). Despite constant visual input for all conditions, attention may have been more oriented towards, for example, facial expressions of the avatar during the 3P-MENT condition. Did subjects infer the virtual characters’ preference in a non-mentalizing way by simply attending to which object the avatar, for example, was pointing to? Did the mere processing of biological motion or bodily stimuli drove activation of the STS? Our task, however, more than the processing of bodily cues implied mental processes on the side of the avatar (e.g. a preference, decision making, etc.)—as subjects were asked for by the instructions. The question remains to which degree mentalizing and the perception of social—communicative content conveying—stimuli could be dissociated. To this question, Frith and Frith (2006, p. 531) say: ‘Many cues in different modalities can trigger the process of mentalizing as long as they originate from an agent [...] faces, in particular, are an important source of information about their inner states [...] Desires, goals, and intentions can be read from eye gaze direction and body movements.’ Consequently, such stimuli have been often used in operationalizations of mentalizing (Baron-Cohen et al., 2001; Brunet et al., 2003; Gallagher and Frith, 2004; Völlm et al., 2006; Schulte-Rüther et al., 2007). Baron-Cohen et al. (2001) in their ‘Reading-the-mind-in-the-eyes’ test—used as a diagnostic tool for mentalizing deficits—showed pictures of eyes expressing different states of mind. With respect to activation of the STS, there is evidence that it is especially driven by social cues that convey mental states or communicative intent instead of the mere presence of faces or bodies alone. For example, it has been suggested that the STS especially encodes the intentional aspect occurring in the typical biological motion stimuli (Pelphrey et al., 2004b; Saxe et al., 2004b). In addition, the STS seems more active during the perception of expressive as opposed to instrumental gestures (Gallagher and Frith, 2004) and also responds to intentionally moving abstract—non-bodily—stimuli (Schultze et al., 2004, 2005). Accordingly, we did not find increased activation during 3P-MENT in other areas known to be modulated by attention to facial or bodily stimuli such as the fusiform face area (Kanwisher et al., 1997), extrastriate body area (Downing et al., 2001) or amygdala (Adolphs, 2003).
The TPJ has been crucially implicated in mentalizing (or ‘theory of mind’; Saxe and Wexler, 2005; Saxe et al., 2006), especially in the understanding of another person's false belief, but no significant TPJ activation was revealed in the present mentalizing task. Our stimulus material, however, did not consist of the typically used false belief tasks or theory-of-mind stories (Vogeley et al., 2001; Saxe and Wexler, 2005; Saxe et al., 2006) and did not tap into the assessment of false-belief reasoning (as only one aspect of the human mentalizing ability). Furthermore, the presently performed comparisons were different from typically computed comparisons such as ‘mentalizing vs no mentalizing’. Here, another important component of the mentalizing network, the MPFC, was also not activated. A possible explanation is provided by Saxe and colleagues (2006) who suggested that the MPFC is not only specifically engaged in mentalizing (Saxe and Wexler, 2005; Saxe et al., 2006) but also in self-reflection (Gusnard et al., 2001; Ochsner et al., 2004). Previously, authors have discussed a self-referential account to mentalizing—that is, the strategy to estimate others’ mental states by reflecting and using one's own thoughts, intentions or desires—especially in association with recruitment of the MPFC (as nicely demonstrated by Jenkins et al., 2008). Accordingly, one would not expect MPFC to be more activated for 3P-MENT vs, for example, 1P-MENT with both conditions involving self-reflection. Nonetheless, in accordance with previous evidence (Brunet et al., 2000; Gallagher et al. 2000; Völlm et al., 2006) we detected activation of the right precuneus and the middle frontal gyrus during mentalizing. Interestingly, these areas were not differentially activated for 3P-MENT in comparison to 3P-PERSP, and indeed partially also showed common activation in the conjunction analysis.
With respect to functional commonalities between mentalizing and perspective taking, both appear to rely on parietal and premotor cortices as indicated by the conjunction analysis. Does this commonly found activation reflect a mechanism within parietal and premotor areas underlying both processes? Interestingly, parietal as well as premotor cortices have been associated with the human mirror neuron system (Buccino et al., 2004; Keysers and Perrett, 2004). Mirror neurons, as originally found in the macaque, have been proposed as neural substrates of simulation theory (Gallese and Goldman, 1998; Keysers and Perrett, 2004). The underlying idea that we use our own experiences to simulate those of other people in order to understand or predict their behavior has indeed been suggested as one mechanism relevant for social cognition, including mentalizing and perspective taking (Gallese and Goldman, 1998; Langdon and Coltheart, 2001; Keysers and Perrett, 2004; cf. Saxe, 2005, who argues against simulation underlying mental state attributions). The posterior STS here, however, was selectively activated during mentalizing but not perspective taking. This finding is suggestive of a process within the STS that goes beyond simulation or other processes also involved in perspective taking. Such an additional process in support of mentalizing may be the analysis of higher complex, representational and socially relevant signals such as facial or bodily cues that convey a communicative intent or information on another person's mental states.
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TOPABSTRACTMATERIALS AND METHODSRESULTSDISCUSSIONSUPPLEMENTARY DATAREFERENCES |
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Supplementary data are available at SCAN online.
Received January 6, 2008. Accepted July 11, 2008.
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Adolphs R. Cognitive neuroscience of human social behaviour. Nature Reviews. Neuroscience (2003) 4(3):165–78.[Web of Science][Medline]
Aichhorn M, Perner J, Kronbichler M, Staffen W, Ladurner G. Do visual perspective tasks need theory of mind? NeuroImage (2006) 30(3):1059–68.[CrossRef][Web of Science][Medline]
Allison T, Puce A, McCarthy G. Social perception from visual cues: role of the STS region. Trends in Cognitive Science (2000) 4(7):267–78.[CrossRef][Web of Science][Medline]
Arzy S, Thut G, Mohr C, Michel CM, Blanke O. Neural basis of embodiment: distinct contributions of temporoparietal junction and extrastriate body area. The Journal of Neuroscience (2006) 26(31):8074–81.
Ashburner J, Friston KJ. Rigid body registration. In: Human Brain Function.—Frackowiak RS, Friston KJ, Frith CD, et al, eds. (2003) 2nd. London, UK: Academic Press. 635–55.
Ashburner J, Friston KJ. Unified segmentation. Neuroimage (2005) 26(3):839–51.[CrossRef][Web of Science][Medline]
Baron-Cohen S. Mindblindness: An Essay on Autism and Theory of Mind. (1997) Cambridge: MIT Press.
Baron-Cohen S, Wheelwright S, Hill J. The ‘reading the mind in the eyes’ test revised version: a study with normal adults, and adults with Asperger syndrome or high-functioning autism. Journal of Child Psychology and Psychiatry (2001) 42:241–52.[CrossRef][Web of Science][Medline]
Baron-Cohen S, Knickmeyer RC, Belmonte MK. Sex differences in the brain: implications for explaining autism. Science (2005) 310(5749):819–23.
Bente G, Krämer NC, Petersen A, de Ruiter JP. Computer animated movement and person perception: methodological advances in nonverbal behavior research. Journal of Nonverbal Behavior (2001) 25(3):151–66.[CrossRef][Web of Science]
Bewernick BH, David N, Vogeley K. Certainty of mental attribution in a nonverbal theory of mind task. Journal of Cognitive Neuroscience (2005) 69–70.
Blakemore SJ, Boyer P, Pachot-Clouard M, Meltzoff A, Segebarth C, Decety J. The detection of contingency and animacy from simple animations in the human brain. Cereb Cortex (2003) 13(8):837–44.
Blanke O, Mohr C, Michel CM, et al. Linking out-of-body experience and self processing to mental own-body imagery at the temporoparietal junction. The Journal of Neuroscience (2005) 25(3):550–7.
Bonda E, Petrides M, Ostry D, Evans A. Specific involvement of human parietal systems and the amygdala in the perception of biological motion. The Journal of Neuroscience (1996) 16(11):3737–44.
Brunet E, Sarfati Y, Hardy-Bayle MC, Decety J. Abnormalities of brain function during a nonverbal theory of mind task in schizophrenia. Neuropsychologia (2003) 41(12):1574–82.[CrossRef][Web of Science][Medline]
Buccino G, Lui F, Canessa N, et al. Neural circuits involved in the recognition of actions performed by nonconspecifics: an fMRI study. Journal of Cognitive Neuroscience (2004) 16(1):114–26.[CrossRef][Web of Science][Medline]
Castelli F, Happe F, Frith U, Frith C. Movement and mind: a functional imaging study of perception and interpretation of complex intentional movement patterns. NeuroImage (2000) 12(3):314–25.[CrossRef][Web of Science][Medline]
Cohen MX, David N, Vogeley K, Elger CE. Gamma-band activity in the human superior temporal sulcus during mentalizing from nonverbal social cues. In: Psychophysiology. (in press).
Collins DL, Neelin P, Peters TM, Evans AC. Automatic 3d intersubject registration of mr volumetric data in standardized Talairach space. Journal of Computer Assisted Tomography (1994) 18(2):192–205.[Web of Science][Medline]
David N, Bewernick BH, Cohen MX, et al. Neural representations of self versus other: visual-perspective taking and agency in a virtual ball-tossing game. Journal of Cognitive Neuroscience (2006) 18(6):898–910.[CrossRef][Web of Science][Medline]
Downing PE, Jiang Y, Shuman M, Kanwisher N. A cortical area selective for visual processing of the human body. Science (2001) 293(5539):2470–3.
Duvernoy HM. The Human Brain. (1999) 2nd. Wien, New York: Springer.
Eickhoff SB, Heim S, Zilles K, Amunts K. Testing anatomically specified hypotheses in functional imaging using cytoachtectonic maps. NeuroImage (2006) 32(2):570–82.[CrossRef][Web of Science][Medline]
Evans AC, Marrett S, Neelin P, et al. Anatomical mapping of functional activation in stereotactic coordinate space. Neuroimage (1992) 1(1):43–53.[Medline]
Flavell JH, Everett BA, Croft K, Flavell ER. Young children's knowledge about visual perception: further evidence for the level 1 - level 2 distinction. Developmental Psychology (1981) 17:99–103.[CrossRef][Web of Science]
Flavell JH, Shipstead SG, Croft K. Young children's knowledge about visual perception: hiding objects from others. Child Development (1978) 49(4):1208–11.[CrossRef][Web of Science][Medline]
Friston KJ, Penny WD, Glaser DE. Conjunction revisited. NeuroImage (2005) 25(3):661–7.[CrossRef][Web of Science][Medline]
Frith CD, Frith U. The neural basis of mentalizing. Neuron (2006) 50(4):531–4.[CrossRef][Web of Science][Medline]
Frith U, Frith CD. Interacting minds - a biological basis. Science (1999) 286(5445):1692–5.
Frith U, Frith CD. Development and neurophysiology of mentalizing. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences (2003) 358(1431):459–73.[CrossRef][Web of Science]
Gallagher HL, Frith CD. Functional imaging of ‘theory of mind’. Trends in Cognitive Science (2003) 7(2):77–83.[CrossRef][Web of Science][Medline]
Gallagher HL, Frith CD. Dissociable neural pathways for the perception and recognition of expressive and instrumental gestures. Neuropsychologia (2004) 42(13):1725–36.[Web of Science][Medline]
Gallagher HL, Happe F, Brunswick N, Fletcher PC, Frith U, Frith CD. Reading the mind in cartoons and stories: an fMRI study of ‘theory of mind’ in verbal and nonverbal tasks. Neuropsychologia (2000) 38(1):11–21.[CrossRef][Web of Science][Medline]
Gallagher HL, Jack AI, Roepstorff A, Frith CD. Imaging the intentional stance in a competitive game. NeuroImage (2002) 16(3 Pt 1):814–21.[CrossRef][Web of Science][Medline]
Gallese V, Goldman A. Mirror neurons and the simulation theory of mind-reading. Trends in Cognitive Science (1998) 2(2):493–501.[CrossRef][Web of Science]
Grossman ED, Blake R. Brain areas active during visual perception of biological motion. Neuron (2002) 35(6):1167–75.[CrossRef][Web of Science][Medline]
Gusnard DA, Akbudak E, Shulman GL, Raichle ME. Medial prefrontal cortex and self-referential mental activity: relation to a default mode of brain function. Proceedings of the National Academy of Sciences of the United States of America (2001) 98(7):4259–64.
Hare B, Call J, Agnetta B, Tomasello M. Chimpanzees know what conspecifics do and do not see. Animal Behaviour (2000) 59(4):771–85.[CrossRef][Web of Science][Medline]
Hare B, Call J, Tomasello M. Do chimpanzees know what conspecifics know? Animal Behaviour (2001) 61(1):139–51.[CrossRef][Web of Science][Medline]
Jackson PL, Meltzoff AN, Decety J. Neural circuits involved in imitation and perspective-taking. NeuroImage (2006) 31(1):429–39.[CrossRef][Web of Science][Medline]
Jenkins AC, Macrae CN, Mitchell JP. Repetition suppression of ventromedial prefrontal activity during judgments of self and others. Proceedings of the National Academy of Sciences of the United States of America (2008) 105(11):4507–12.
Kanwisher N, McDermott J, Chun MM. The fusiform face area: a module in human extrastriate cortex specialized for face perception. Journal of Neuroscience (1997) 17(11):4302–11.
Keysers C, Perrett DI. Demystifying social cognition: a Hebbian perspective. Trends in Cognitive Science (2004) 8(11):501–7.[CrossRef][Web of Science][Medline]
Kiebel S, Holmes AP. The general linear model. In: Human Brain Function.—Frackowiak RS, Friston KJ, Frith CD, et al, eds. (2003) 2nd. London, UK: Academic Press. 725–60.
Langdon R, Coltheart M. Visual perspective-taking and schizotypy: evidence for a simulation-based account of mentalizing in normal adults. Cognition (2001) 82(1):1–26.[CrossRef][Web of Science][Medline]
Maldjian JA, Laurienti PJ, Kraft RA, Burdette JH. An automated method for neuroanatomic and cytoarchitectonic atlas-based interrogation of fMRI data sets. NeuroImage (2003) 19:1233–9.[CrossRef][Web of Science][Medline]
Meltzoff AN. Origins of theory of mind, cognition and communication. Journal of Communications Disorders (1999) 32(4):251–69.[CrossRef]
Narumoto J, Yamada H, Iidaka T, et al. Brain regions involved in verbal or non-verbal aspects of facial emotion recognition. Neuroreport (2000) 11(11):2571–6.[Web of Science][Medline]
Newcombe N. The development of spatial perspective taking. Advances in Child Development and Behavior (1989) 22:203–47.[Web of Science][Medline]
Ochsner KN, Knierim K, Ludlow DH, et al. Reflecting upon feelings: an fMRI study of neural systems supporting the attribution of emotion to self and other. Journal of Cognitive Neuroscience (2004) 16(10):1746–72.[CrossRef][Web of Science][Medline]
Oldfield RC. The assessment and analysis of handedness: the Edinburgh Inventory. Neuropsychologia, 9 (1971) 97–113.
Parsons TD, Larson P, Kratz K, et al. Sex differences in mental rotation and spatial rotation in a virtual environment. Neuropsychologia (2004) 42(4):555–62.[CrossRef][Web of Science][Medline]
Parsons LM. Imagined transformation of one's body. Journal of Experimantal Psychology. General (1987) 116:172–91.[CrossRef]
Pelphrey K, Adolphs R, Morris JP. Neuroanatomical substrates of social cognition dysfunction in autism. Mental Retardation and Developmental Disabilities Research Reviews (2004a) 10(4):259–71.[CrossRef][Web of Science][Medline]
Pelphrey KA, Morris JP, McCarthy G. Grasping the intentions of others: the perceived intentionality of an action influences activity in the superior temporal sulcus during social perception. Journal of Cognitive Neuroscience (2004b) 16(10):1706–16.[CrossRef][Web of Science][Medline]
Penny WD, Holmes AP. Random effects analysis. In: Human Brain Function.—Frackowiak RS, Friston KJ, Frith CD, et al, eds. (2003) 2nd. London, UK: Academic Press. 843–50.
Piaget J, Inhelder B, eds. The coordination of perspectives. In: The Child's; Conception of Space. (1967) New York: Norton & Co. 209–46.
Povinelli DJ, Eddy TJ. What young chimpanzees know about seeing. Monographs of the Society for Research in Child Development (1996) 61(3). i–vi, 1–152; discussion 153–91.
Premack D, Woodruff G. Does the chimpanzee have a theory of mind? Behavioural and Brain Sciences (1978) 4:515–26.
Price CJ, Friston KJ. Cognitive conjunction: a new approach to brain activation experiments. NeuroImage (1997) 5(4 Pt 1):261–70.[CrossRef][Web of Science][Medline]
Reed T, Peterson C. A comparative study of autistic subjects’ performance at two levels of visual and cognitive perspective taking. Journal of Autism and Developmental Disorders (1990) 20(4):555–67.[CrossRef][Web of Science][Medline]
Rilling JK, Sanfey AG, Aronson JA, Nystrom LE, Cohen JD. The neural correlates of theory of mind within interpersonal interactions. NeuroImage (2004) 22(4):1694–703.[CrossRef][Web of Science][Medline]
Ruby P, Decety J. Effect of subjective perspective taking during simulation of action: a PET investigation of agency. Nature Neuroscience (2001) 4(5):546–50.[Web of Science][Medline]
Ruby P, Decety J. What you believe versus what you think they believe: a neuroimaging study of conceptual perspective-taking. The European Journal of Neuroscience (2003) 17(11):2475–80.[CrossRef][Web of Science][Medline]
Ruby P, Decety J. How would you feel versus how do you think she would feel? A neuroimaging study of perspective-taking with social emotions. Journal of Cognitive Neuroscience (2004) 16(6):988–99.[CrossRef][Web of Science][Medline]
Saxe R. Against simulation: the argument from error. Trends in Cognitive Science (2005) 9(4):174–9.[CrossRef][Web of Science][Medline]
Saxe R. Uniquely human social cognition. Current Opinion in Neurobiology (2006) 16(2):235–9.[CrossRef][Web of Science][Medline]
Saxe R, Kanwisher N. People thinking about thinking people. The role of the temporo-parietal junction in ‘theory of mind’. Neuroimage (2003) 19(4):1835–42.[CrossRef][Web of Science][Medline]
Saxe R, Wexler A. Making sense of another mind: the role of the right temporo-parietal junction. Neuropsychologia (2005) 43(10):1391–9.[CrossRef][Web of Science][Medline]
Saxe R, Carey S, Kanwisher N. Understanding other minds: linking developmental psychology and functional neuroimaging. Annual Review of Psychology (2004a) 55:87–124.[CrossRef][Web of Science][Medline]
Saxe R, Xiao DK, Kovacs G, Perrett DI, Kanwisher N. A region of right posterior superior temporal sulcus responds to observed intentional actions. Neuropsychologia (2004b) 42(11):1435–46.[CrossRef][Web of Science][Medline]
Saxe R, Moran J, Scholz J, Gabrieli J. Overlapping and non-overlapping brain regions for theory of mind and self reflection in individual subjects. In: Social Cognitive and Affective Neuroscience, 1. (2006) 229–234.
Schilbach L, Wohlschlaeger AM, Kraemer NC, et al. Being with virtual others: neural correlates of social interaction. Neuropsychologia (2006) 44(5):718–30.[CrossRef][Web of Science][Medline]
Schultz J, Friston KJ, O’Doherty J, Wolpert DM, Frith CD. Activation in posterior superior temporal sulcus parallels parameter inducing the percept of animacy. Neuron (2005) 45(4):625–35.[CrossRef][Web of Science][Medline]
Schultz J, Imamizu H, Kawato M, Frith CD. Activation of the human superior temporal gyrus during observation of goal attribution by intentional objects. Journal of Cognitive Neuroscience (2004) 16(10):1695–705.[CrossRef][Web of Science][Medline]
Talairach J, Tournoux P. Co-planar Stereotaxic Atlas of the Human Brain. (1988) Stuttgart: Georg Thieme Verlag.
Vogeley K, Bussfeld P, Newen A, et al. Mind reading: neural mechanisms of theory of mind and self-perspective. NeuroImage (2001) 14(1 Pt 1):170–81.[CrossRef][Web of Science][Medline]
Vogeley K, Fink GR. Neural correlates of the first-person-perspective. Trends in Cognitive Science (2003) 7(1):38–42.[CrossRef][Web of Science][Medline]
Vogeley K, May M, Ritzl A, Falkai P, Zilles K, Fink GR. Neural correlates of first-person perspective as one constituent of human self-consciousness. Journal of Cognitive Neuroscience (2004) 16(5):817–27.[CrossRef][Web of Science][Medline]
Völlm BA, Taylor AN, Richardson P, et al. Neuronal correlates of theory od mind and empathy: a functional magentic resonance study in a nonverbal task. NeuroImage (2006) 29(1):90–8.[Web of Science][Medline]
Yirmiya N, Sigman M, Zacks D. Perceptual perspective-taking and seriation abilities in high-functioning children with autism. Development and Psychopathology (1994) 6:263–72.[Web of Science]
Zacks J, Rypma B, Gabrieli JD, Tversky B, Glover GH. Imagined transformations of bodies: an fMRI investigation. Neuropsychologia (1999) 37(9):1029–40.[CrossRef][Web of Science][Medline]
Zacks JM, Vettel JM, Michelon P. Imagined viewer and object rotations dissociated with event-related fMRI. Journal of Cognitive Neuroscience (2003) 15(7):1002–18.[CrossRef][Web of Science][Medline]
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