ABSTRACT Difficulties in reflecting on inner experience and identifying subjective feelings (alex... more ABSTRACT Difficulties in reflecting on inner experience and identifying subjective feelings (alexithymia) have been recognized as characteristic of autism spectrum disorders (ASD). To date, these difficulties have only been investigated with behavioural and self-report measures. This study is the first explicit attempt to investigate the neural activity associated with attention to subjective emotional responses in high-functioning adults with ASD. 15 individuals with ASD and 15 matched control (CTRL) participants completed two tasks ...
Affective touch Empathy Orbito-frontal cortex Anterior insula a b s t r a c t In spite of conside... more Affective touch Empathy Orbito-frontal cortex Anterior insula a b s t r a c t In spite of considerable progress in the understanding of the neural mechanisms underlying the experience of empathy, the majority of previous investigations have focused on how we share negative affective states (and in particular pain) of others, whereas only few studies have targeted empathy for positive emotions. This bias has precluded addressing one of the central tenets of the shared representations account of empathy, which is that different networks should be engaged when empathizing with emotions that are represented on different neural levels. The aim of the present study was to overcome this limitation and to test whether empathy for pleasant and unpleasant affective touch is underpinned by different neural networks. To this end we used functional magnetic resonance imaging (fMRI), with two independent replication experiments (N ¼ 18, N ¼ 32), and a novel paradigm enabling the joint investigation of first-hand and vicarious responses to pleasant and unpleasant affect induced via visuo-tactile stimulation. This revealed that empathy is subserved by distinct neural networks, with those regions recruited in the first-hand experience of positive or negative affective states also being specifically recruited when empathizing with these respective states in others. More specifically, the first-hand and vicarious experience of pleasant touch commonly recruited medial orbitofrontal cortex (OFC), while unpleasant touch was associated with shared activation in the right fronto-insular cortex. The observation that specifically tailored subsystems of the human brain are engaged to share positive versus negative touch of others brings fresh evidence to one of the major goals of the social neuroscience of empathy: to identify which specific aspects of the affective states of others are shared, and what role this plays in enabling the understanding of the emotions of others.
a b s t r a c t a r t i c l e i n f o Keywords: prosocial behavior virtual reality gICA salience ... more a b s t r a c t a r t i c l e i n f o Keywords: prosocial behavior virtual reality gICA salience network mPFC
Humans tend to use the self as a reference point to perceive the world and gain information about... more Humans tend to use the self as a reference point to perceive the world and gain information about other people's mental states. However, applying such a self-referential projection mechanism in situations where it is inappropriate can result in egocentrically biased judgments. To assess egocentricity bias in the emotional domain (EEB), we developed a novel visuo-tactile paradigm assessing the degree to which empathic judgments are biased by one's own emotions if they are incongruent to those of the person we empathize with. A first behavioral experiment confirmed the existence of such EEB, and two independent fMRI experiments revealed that overcoming biased empathic judgments is associated with increased activation in the right supramarginal gyrus (rSMG), in a location distinct from activations in right temporoparietal junction reported in previous social cognition studies. Using temporary disruption of rSMG with repetitive transcranial magnetic stimulation resulted in a substantial increase of EEB, and so did reducing visuo-tactile stimulation time as shown in an additional behavioral experiment. Our findings provide converging evidence from multiple methods and experiments that rSMG is crucial for overcoming emotional egocentricity. Effective connectivity analyses suggest that this may be achieved by early perceptual regulation processes disambiguating proprioceptive first-person information (touch) from exteroceptive third-person information (vision) during incongruency between self-and other-related affective states. Our study extends previous models of social cognition. It shows that although shared neural networks may underlie emotional understanding in some situations, an additional mechanism subserved by rSMG is needed to avoid biased social judgments in other situations.
Autism is associated with an inability to identify and distinguish one's own feelings. We assesse... more Autism is associated with an inability to identify and distinguish one's own feelings. We assessed this inability using alexithymia and empathy questionnaires, and used fMRI to investigate brain activity while introspecting on emotion. Individuals with high functioning autism/Asperger syndrome (HFA/AS) were compared with matched controls. Participants rated stimuli from the International Affective Picture System twice, once according to the degree of un/pleasantness that the pictures induced, and once according to their color balance. The groups differed significantly on both alexithymia and empathy questionnaires. Alexithymia and lack of empathy were correlated, indicating a link between understanding one's own and others' emotions. For both groups a strong relationship between questionnaire scores and brain activity was found in the anterior insula (AI), when participants were required to assess their feelings to unpleasant pictures. Regardless of selfreported degree of emotional awareness, individuals with HFA/AS differed from controls when required to introspect on their feelings by showing reduced activation in self-reflection/mentalizing regions. Thus, we conclude that difficulties in emotional awareness are related to hypoactivity in AI in both individuals with HFA/AS and controls, and that the particular difficulties in emotional awareness in individuals with HFA/AS are not related to their impairments in selfreflection/mentalizing.
Recent research has shown that experiencing events that represent a significant threat to social ... more Recent research has shown that experiencing events that represent a significant threat to social bonds activates a network of brain areas associated with the sensory-discriminative aspects of pain. In the present study, we investigated whether the same brain areas are involved when witnessing social exclusion threats experienced by others. Using a within-subject design, we show that an ecologically valid experience of social exclusion recruits areas coding the somatosensory components of physical pain (posterior insular cortex and secondary somatosensory cortex). Furthermore, we show that this pattern of activation not only holds for directly experienced social pain, but also during empathy for social pain. Finally, we report that subgenual cingulate cortex is the only brain area conjointly active during empathy for physical and social pain. This supports recent theories that affective processing and homeostatic regulation are at the core of empathic responses.
Please cite this article as: L.TomovaB. von DawansM. HeinrichsG. SilaniC. Lamm Is stress affectin... more Please cite this article as: L.TomovaB. von DawansM. HeinrichsG. SilaniC. Lamm Is stress affecting our ability to tune into others? Evidence for gender differences in the effects of stress on self-other distinction (2014), http://dx.
Please cite this article in press as: Seidel, E.M., et al., The impact of social exclusion vs. in... more Please cite this article in press as: Seidel, E.M., et al., The impact of social exclusion vs. inclusion on subjective and hormonal reactions in females and males. Psychoneuroendocrinology (2013), http://dx.
Little is known about the neurobiological mechanisms underlying prosocial decisions and how they ... more Little is known about the neurobiological mechanisms underlying prosocial decisions and how they are modulated by social factors such as perceived group membership. The present study investigates the neural processes preceding the willingness to engage in costly helping toward ingroup and outgroup members. Soccer fans witnessed a fan of their favorite team (ingroup member) or of a rival team (outgroup member) experience pain. They were subsequently able to choose to help the other by enduring physical pain themselves to reduce the other's pain. Helping the ingroup member was best predicted by anterior insula activation when seeing him suffer and by associated self-reports of empathic concern. In contrast, not helping the outgroup member was best predicted by nucleus accumbens activation and the degree of negative evaluation of the other. We conclude that empathy-related insula activation can motivate costly helping, whereas an antagonistic signal in nucleus accumbens reduces the propensity to help.
We investigated whether individuals with autism spectrum disorders (ASD) would show attentional m... more We investigated whether individuals with autism spectrum disorders (ASD) would show attentional modulation for social (face) and non-social (house) stimuli. Sixteen individuals with ASD and 16 matched control participants completed a task in which pairs of face and house stimuli were present on every trial, with one of the pairs randomly assigned to attended locations and the other to unattended locations. Both massunivariate (SPM) and region of interest analyses suggested that responses to houses were modulated by attention in both groups, but that only the control participants demonstrated attentional modulation of faceselective regions. Thus, the participants with ASD demonstrated a lack of attentional modulation which was particularly evident for the social stimulus. Analyses of effective connectivity indicated that these results were due to a failure of attention to modulate connectivity between extrastriate areas and V1. We discuss how these results may suggest a mechanism to explain the reduced salience of social stimuli in ASD. D
rCBF PET scans were used to study the physiological bases of lip-reading, a natural skill of extr... more rCBF PET scans were used to study the physiological bases of lip-reading, a natural skill of extracting language from mouth movements, which contributes to speech perception in everyday life. Viewing connected mouth movements that could not be lexically identified and that evoke perception of isolated speech sounds (non-lexical lip-reading) was associated with bilateral activation of auditory association cortex around Wernicke's area, of left dorsal premotor cortex and left opercular-premotor-division of the left inferior frontal gyrus (Broca's area). The supplementary motor area was active as well. These areas have all been implicated in phonogical processing, speech and mouth motor planning and execution. In addition, non-lexical lip-reading also differentially activated visual motion areas. Lexical access through lip-reading was associated with a similar pattern of activation and with additional foci in ventral-and dorsolateral prefrontal cortex bilaterally and in left inferior parietal cortex. Linear regression analysis of cerebral blood flow and proficiency for lexical lip-reading further clarified the role of these areas in gaining access to language through lipreading. The results suggest cortical activation circuits for lip-reading from action representations which may differentiate lexical access from nonlexical processes.
■ Prior lesion and functional imaging studies have highlighted the importance of the left ventral... more ■ Prior lesion and functional imaging studies have highlighted the importance of the left ventral occipito-temporal (LvOT) cortex for visual word recognition. Within this area, there is a posterioranterior hierarchy of subregions that are specialized for different stages of orthographic processing. The aim of the present fMRI study was to dissociate the effects of subword orthographic typicality (e.g., cider [high] vs. cynic [low]) from the effect of lexicality (e.g., pollen [word] vs. pillen [pseudoword]). We therefore orthogonally manipulated the orthographic typicality of written words and pseudowords (nonwords and pseudohomophones) in a visual lexical decision task. Consistent with previous studies, we identified greater activation for pseudowords than words (i.e., an effect of lexicality) in posterior LvOT cortex. In addition, we revealed higher activation for atypical than typical strings, irrespective of lexicality, in a left inferior occipital region that is posterior to LvOT cortex. When lexical decisions were made more difficult in the context of pseudohomophone foils, left anterior temporal activation also increased for atypical relative to typical strings. The latter finding agrees with the behavior of patients with progressive anterior temporal lobe degeneration, who have particular difficulty recognizing words with atypical orthography. The most novel outcome of this study is that, within a distributed network of regions supporting orthographic processing, we have identified a left inferior occipital region that is particularly sensitive to the typicality of subword orthographic patterns. ■
In this study, we tested the validity of 2 popular assumptions about empathy: (a) empathy can be ... more In this study, we tested the validity of 2 popular assumptions about empathy: (a) empathy can be enhanced by oxytocin, a neuropeptide known to be crucial in affiliative behavior, and (b) individual differences in prosocial behavior are positively associated with empathic brain responses. To do so, we measured brain activity in a double-blind placebo-controlled study of 20 male participants either receiving painful stimulation to their own hand (self condition) or observing their female partner receiving painful stimulation to her hand (other condition). Prosocial behavior was measured using a monetary economic interaction game with which participants classified as prosocial (N ϭ 12) or selfish (N ϭ 6), depending on whether they cooperated with another player. Empathy-relevant brain activation (anterior insula) was neither enhanced by oxytocin nor positively associated with prosocial behavior. However, oxytocin reduced amygdala activation when participants received painful stimulation themselves (in the nonsocial condition). Surprisingly, this effect was driven by "selfish" participants. The results suggest that selfish individuals may not be as rational and unemotional as usually suggested, their actions being determined by their feeling anxious rather than by reason.
Thirteen right brain-damaged patients who were found to neglect pictures presented on the left of... more Thirteen right brain-damaged patients who were found to neglect pictures presented on the left of a display were presented the same stimuli, intermixed with foils, in a yes-no recognition test. Fifty per cent of patients claimed to have already seen one or more of the previously neglected pictures. This demonstrates that visual information that fails to access consciousness in neglect patients does retain the ability to surface as explicit memory at a later stage.
ABSTRACT Difficulties in reflecting on inner experience and identifying subjective feelings (alex... more ABSTRACT Difficulties in reflecting on inner experience and identifying subjective feelings (alexithymia) have been recognized as characteristic of autism spectrum disorders (ASD). To date, these difficulties have only been investigated with behavioural and self-report measures. This study is the first explicit attempt to investigate the neural activity associated with attention to subjective emotional responses in high-functioning adults with ASD. 15 individuals with ASD and 15 matched control (CTRL) participants completed two tasks ...
Affective touch Empathy Orbito-frontal cortex Anterior insula a b s t r a c t In spite of conside... more Affective touch Empathy Orbito-frontal cortex Anterior insula a b s t r a c t In spite of considerable progress in the understanding of the neural mechanisms underlying the experience of empathy, the majority of previous investigations have focused on how we share negative affective states (and in particular pain) of others, whereas only few studies have targeted empathy for positive emotions. This bias has precluded addressing one of the central tenets of the shared representations account of empathy, which is that different networks should be engaged when empathizing with emotions that are represented on different neural levels. The aim of the present study was to overcome this limitation and to test whether empathy for pleasant and unpleasant affective touch is underpinned by different neural networks. To this end we used functional magnetic resonance imaging (fMRI), with two independent replication experiments (N ¼ 18, N ¼ 32), and a novel paradigm enabling the joint investigation of first-hand and vicarious responses to pleasant and unpleasant affect induced via visuo-tactile stimulation. This revealed that empathy is subserved by distinct neural networks, with those regions recruited in the first-hand experience of positive or negative affective states also being specifically recruited when empathizing with these respective states in others. More specifically, the first-hand and vicarious experience of pleasant touch commonly recruited medial orbitofrontal cortex (OFC), while unpleasant touch was associated with shared activation in the right fronto-insular cortex. The observation that specifically tailored subsystems of the human brain are engaged to share positive versus negative touch of others brings fresh evidence to one of the major goals of the social neuroscience of empathy: to identify which specific aspects of the affective states of others are shared, and what role this plays in enabling the understanding of the emotions of others.
a b s t r a c t a r t i c l e i n f o Keywords: prosocial behavior virtual reality gICA salience ... more a b s t r a c t a r t i c l e i n f o Keywords: prosocial behavior virtual reality gICA salience network mPFC
Humans tend to use the self as a reference point to perceive the world and gain information about... more Humans tend to use the self as a reference point to perceive the world and gain information about other people's mental states. However, applying such a self-referential projection mechanism in situations where it is inappropriate can result in egocentrically biased judgments. To assess egocentricity bias in the emotional domain (EEB), we developed a novel visuo-tactile paradigm assessing the degree to which empathic judgments are biased by one's own emotions if they are incongruent to those of the person we empathize with. A first behavioral experiment confirmed the existence of such EEB, and two independent fMRI experiments revealed that overcoming biased empathic judgments is associated with increased activation in the right supramarginal gyrus (rSMG), in a location distinct from activations in right temporoparietal junction reported in previous social cognition studies. Using temporary disruption of rSMG with repetitive transcranial magnetic stimulation resulted in a substantial increase of EEB, and so did reducing visuo-tactile stimulation time as shown in an additional behavioral experiment. Our findings provide converging evidence from multiple methods and experiments that rSMG is crucial for overcoming emotional egocentricity. Effective connectivity analyses suggest that this may be achieved by early perceptual regulation processes disambiguating proprioceptive first-person information (touch) from exteroceptive third-person information (vision) during incongruency between self-and other-related affective states. Our study extends previous models of social cognition. It shows that although shared neural networks may underlie emotional understanding in some situations, an additional mechanism subserved by rSMG is needed to avoid biased social judgments in other situations.
Autism is associated with an inability to identify and distinguish one's own feelings. We assesse... more Autism is associated with an inability to identify and distinguish one's own feelings. We assessed this inability using alexithymia and empathy questionnaires, and used fMRI to investigate brain activity while introspecting on emotion. Individuals with high functioning autism/Asperger syndrome (HFA/AS) were compared with matched controls. Participants rated stimuli from the International Affective Picture System twice, once according to the degree of un/pleasantness that the pictures induced, and once according to their color balance. The groups differed significantly on both alexithymia and empathy questionnaires. Alexithymia and lack of empathy were correlated, indicating a link between understanding one's own and others' emotions. For both groups a strong relationship between questionnaire scores and brain activity was found in the anterior insula (AI), when participants were required to assess their feelings to unpleasant pictures. Regardless of selfreported degree of emotional awareness, individuals with HFA/AS differed from controls when required to introspect on their feelings by showing reduced activation in self-reflection/mentalizing regions. Thus, we conclude that difficulties in emotional awareness are related to hypoactivity in AI in both individuals with HFA/AS and controls, and that the particular difficulties in emotional awareness in individuals with HFA/AS are not related to their impairments in selfreflection/mentalizing.
Recent research has shown that experiencing events that represent a significant threat to social ... more Recent research has shown that experiencing events that represent a significant threat to social bonds activates a network of brain areas associated with the sensory-discriminative aspects of pain. In the present study, we investigated whether the same brain areas are involved when witnessing social exclusion threats experienced by others. Using a within-subject design, we show that an ecologically valid experience of social exclusion recruits areas coding the somatosensory components of physical pain (posterior insular cortex and secondary somatosensory cortex). Furthermore, we show that this pattern of activation not only holds for directly experienced social pain, but also during empathy for social pain. Finally, we report that subgenual cingulate cortex is the only brain area conjointly active during empathy for physical and social pain. This supports recent theories that affective processing and homeostatic regulation are at the core of empathic responses.
Please cite this article as: L.TomovaB. von DawansM. HeinrichsG. SilaniC. Lamm Is stress affectin... more Please cite this article as: L.TomovaB. von DawansM. HeinrichsG. SilaniC. Lamm Is stress affecting our ability to tune into others? Evidence for gender differences in the effects of stress on self-other distinction (2014), http://dx.
Please cite this article in press as: Seidel, E.M., et al., The impact of social exclusion vs. in... more Please cite this article in press as: Seidel, E.M., et al., The impact of social exclusion vs. inclusion on subjective and hormonal reactions in females and males. Psychoneuroendocrinology (2013), http://dx.
Little is known about the neurobiological mechanisms underlying prosocial decisions and how they ... more Little is known about the neurobiological mechanisms underlying prosocial decisions and how they are modulated by social factors such as perceived group membership. The present study investigates the neural processes preceding the willingness to engage in costly helping toward ingroup and outgroup members. Soccer fans witnessed a fan of their favorite team (ingroup member) or of a rival team (outgroup member) experience pain. They were subsequently able to choose to help the other by enduring physical pain themselves to reduce the other's pain. Helping the ingroup member was best predicted by anterior insula activation when seeing him suffer and by associated self-reports of empathic concern. In contrast, not helping the outgroup member was best predicted by nucleus accumbens activation and the degree of negative evaluation of the other. We conclude that empathy-related insula activation can motivate costly helping, whereas an antagonistic signal in nucleus accumbens reduces the propensity to help.
We investigated whether individuals with autism spectrum disorders (ASD) would show attentional m... more We investigated whether individuals with autism spectrum disorders (ASD) would show attentional modulation for social (face) and non-social (house) stimuli. Sixteen individuals with ASD and 16 matched control participants completed a task in which pairs of face and house stimuli were present on every trial, with one of the pairs randomly assigned to attended locations and the other to unattended locations. Both massunivariate (SPM) and region of interest analyses suggested that responses to houses were modulated by attention in both groups, but that only the control participants demonstrated attentional modulation of faceselective regions. Thus, the participants with ASD demonstrated a lack of attentional modulation which was particularly evident for the social stimulus. Analyses of effective connectivity indicated that these results were due to a failure of attention to modulate connectivity between extrastriate areas and V1. We discuss how these results may suggest a mechanism to explain the reduced salience of social stimuli in ASD. D
rCBF PET scans were used to study the physiological bases of lip-reading, a natural skill of extr... more rCBF PET scans were used to study the physiological bases of lip-reading, a natural skill of extracting language from mouth movements, which contributes to speech perception in everyday life. Viewing connected mouth movements that could not be lexically identified and that evoke perception of isolated speech sounds (non-lexical lip-reading) was associated with bilateral activation of auditory association cortex around Wernicke's area, of left dorsal premotor cortex and left opercular-premotor-division of the left inferior frontal gyrus (Broca's area). The supplementary motor area was active as well. These areas have all been implicated in phonogical processing, speech and mouth motor planning and execution. In addition, non-lexical lip-reading also differentially activated visual motion areas. Lexical access through lip-reading was associated with a similar pattern of activation and with additional foci in ventral-and dorsolateral prefrontal cortex bilaterally and in left inferior parietal cortex. Linear regression analysis of cerebral blood flow and proficiency for lexical lip-reading further clarified the role of these areas in gaining access to language through lipreading. The results suggest cortical activation circuits for lip-reading from action representations which may differentiate lexical access from nonlexical processes.
■ Prior lesion and functional imaging studies have highlighted the importance of the left ventral... more ■ Prior lesion and functional imaging studies have highlighted the importance of the left ventral occipito-temporal (LvOT) cortex for visual word recognition. Within this area, there is a posterioranterior hierarchy of subregions that are specialized for different stages of orthographic processing. The aim of the present fMRI study was to dissociate the effects of subword orthographic typicality (e.g., cider [high] vs. cynic [low]) from the effect of lexicality (e.g., pollen [word] vs. pillen [pseudoword]). We therefore orthogonally manipulated the orthographic typicality of written words and pseudowords (nonwords and pseudohomophones) in a visual lexical decision task. Consistent with previous studies, we identified greater activation for pseudowords than words (i.e., an effect of lexicality) in posterior LvOT cortex. In addition, we revealed higher activation for atypical than typical strings, irrespective of lexicality, in a left inferior occipital region that is posterior to LvOT cortex. When lexical decisions were made more difficult in the context of pseudohomophone foils, left anterior temporal activation also increased for atypical relative to typical strings. The latter finding agrees with the behavior of patients with progressive anterior temporal lobe degeneration, who have particular difficulty recognizing words with atypical orthography. The most novel outcome of this study is that, within a distributed network of regions supporting orthographic processing, we have identified a left inferior occipital region that is particularly sensitive to the typicality of subword orthographic patterns. ■
In this study, we tested the validity of 2 popular assumptions about empathy: (a) empathy can be ... more In this study, we tested the validity of 2 popular assumptions about empathy: (a) empathy can be enhanced by oxytocin, a neuropeptide known to be crucial in affiliative behavior, and (b) individual differences in prosocial behavior are positively associated with empathic brain responses. To do so, we measured brain activity in a double-blind placebo-controlled study of 20 male participants either receiving painful stimulation to their own hand (self condition) or observing their female partner receiving painful stimulation to her hand (other condition). Prosocial behavior was measured using a monetary economic interaction game with which participants classified as prosocial (N ϭ 12) or selfish (N ϭ 6), depending on whether they cooperated with another player. Empathy-relevant brain activation (anterior insula) was neither enhanced by oxytocin nor positively associated with prosocial behavior. However, oxytocin reduced amygdala activation when participants received painful stimulation themselves (in the nonsocial condition). Surprisingly, this effect was driven by "selfish" participants. The results suggest that selfish individuals may not be as rational and unemotional as usually suggested, their actions being determined by their feeling anxious rather than by reason.
Thirteen right brain-damaged patients who were found to neglect pictures presented on the left of... more Thirteen right brain-damaged patients who were found to neglect pictures presented on the left of a display were presented the same stimuli, intermixed with foils, in a yes-no recognition test. Fifty per cent of patients claimed to have already seen one or more of the previously neglected pictures. This demonstrates that visual information that fails to access consciousness in neglect patients does retain the ability to surface as explicit memory at a later stage.
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