Jean Decety

Empathy accounts for the naturally occurring subjective experience of similarity between the feelings expressed by self and others without loosing sight of whose feelings belong to whom. Empathy involves not only the affective experience of the other person’s actual or inferred emotional state but also some minimal recognition and understanding of another’s emotional state. In light of multiple levels of analysis ranging from developmental psychology, social psychology, cognitive neuroscience, and clinical neuropsychology, this article proposes a model of empathy that involves parallel and distributed processing in a number of dissociable computational mechanisms. Shared neural representations, self-awareness, mental flexibility, and emotion regulation constitute the basic macrocomponents of empathy, which are underpinned by specific neural systems. This functional model may be used to make specific predictions about the various empathy deficits that can be encountered in different forms of social and neurological disorders.

A growing body of evidence suggests that empathy for pain is underpinned by neural structures that are also involved in the direct experience of pain. In order to assess the consistency of this finding, an image-based meta-analysis of nine independent functional magnetic resonance imaging (fMRI) investigations and a coordinate-based meta-analysis of 32 studies that had investigated empathy for pain using fMRI were conducted. The results indicate that a core network consisting of bilateral anterior insular cortex and medial/anterior cingulate cortex is associated with empathy for pain. Activation in these areas overlaps with activation during directly experienced pain, and we link their involvement to representing global feeling states and the guidance of adaptive behavior for both self- and other-related experiences. Moreover, the image-based analysis demonstrates that depending on the type of experimental paradigm this core network was co-activated with distinct brain regions: While viewing pictures of body parts in painful situations recruited areas underpinning action understanding (inferior parietal/ventral premotor cortices) to a stronger extent, eliciting empathy by means of abstract visual information about the other's affective state more strongly engaged areas associated with inferring and representing mental states of self and other (precuneus, ventral medial prefrontal cortex, superior temporal cortex, and temporo-parietal junction). In addition, only the picture-based paradigms activated somatosensory areas, indicating that previous discrepancies concerning somatosensory activity during empathy for pain might have resulted from differences in experimental paradigms. We conclude that social neuroscience paradigms provide reliable and accurate insights into complex social phenomena such as empathy and that meta-analyses of previous studies are a valuable tool in this endeavor.

There is a large body of psychological and neuroimaging experiments that have interpreted their findings in favor of a functional equivalence between action generation, action simulation, action verbalization, and perception of action. On the basis of these data, the concept of shared motor representations has been proposed. Indeed several authors have argued that our capacity to understand other people's behavior and to attribute intention or beliefs to others is rooted in a neural, most likely distributed, execution/observation mechanism. Recent neuroimaging studies have explored the neural network engaged during motor execution, simulation, verbalization, and observation. The focus of this metaanalysis is to evaluate in specific detail to what extent the activated foci elicited by these studies overlap.

Whether observation of distress in others leads to empathic concern and altruistic motivation, or to personal distress and egoistic motivation, seems to depend upon the capacity for self-other differentiation and cognitive appraisal. In this experiment, behavioral measures and event-related functional magnetic resonance imaging were used to investigate the effects of perspective-taking and cognitive appraisal while participants observed the facial expression of pain resulting from medical treatment. Video clips showing the faces of patients were presented either with the instruction to imagine the feelings of the patient ("imagine other") or to imagine oneself to be in the patient's situation ("imagine self"). Cognitive appraisal was manipulated by providing information that the medical treatment had or had not been successful. Behavioral measures demonstrated that perspective-taking and treatment effectiveness instructions affected participants' affective responses to the observed pain. Hemodynamic changes were detected in the insular cortices, anterior medial cingulate cortex (aMCC), amygdala, and in visual areas including the fusiform gyrus. Graded responses related to the perspective-taking instructions were observed in middle insula, aMCC, medial and lateral premotor areas, and selectively in left and right parietal cortices. Treatment effectiveness resulted in signal changes in the perigenual anterior cingulate cortex, in the ventromedial orbito-frontal cortex, in the right lateral middle frontal gyrus, and in the cerebellum. These findings support the view that humans' responses to the pain of others can be modulated by cognitive and motivational processes, which influence whether observing a conspecific in need of help will result in empathic concern, an important instigator for helping behavior.

To what extent do we share feelings with others? Neuroimaging investigations of the neural mechanisms involved in the perception of pain in others may cast light on one basic component of human empathy, the interpersonal sharing of affect. In this fMRI study, participants were shown a series of still photographs of hands and feet in situations that are likely to cause pain, and a matched set of control photographs without any painful events. They were asked to assess on-line the level of pain experienced by the person in the photographs. The results demonstrated that perceiving and assessing painful situations in others was associated with significant bilateral changes in activity in several regions notably, the anterior cingulate, the anterior insula, the cerebellum, and to a lesser extent the thalamus. These regions are known to play a significant role in pain processing. Finally, the activity in the anterior cingulate was strongly correlated with the participants' ratings of the others' pain, suggesting that the activity of this brain region is modulated according to subjects' reactivity to the pain of others. Our findings suggest that there is a partial cerebral commonality between perceiving pain in another individual and experiencing it oneself. This study adds to our understanding of the neurological mechanisms implicated in intersubjectivity and human empathy.

Accumulating evidence from cognitive neuroscience indicates that the right inferior parietal cortex, at the junction with the posterior temporal cortex, plays a critical role in various aspects of social cognition such as theory of mind and empathy. With a quantitative meta-analysis of 70 functional neuroimaging studies, the authors demonstrate that this area is also engaged in lower-level (bottom-up) computational processes associated with the sense of agency and reorienting attention to salient stimuli. It is argued that this domain-general computational mechanism is crucial for higher level social cognitive processing. NEUROSCIENTIST 13(6): 580—593, 2007. DOI: 10.1177/1073858407304654

PET was used to map brain regions that are associated with the observation of meaningful and meaningless hand actions. Subjects were scanned under four conditions which consisted of visually presented actions. In each of the four experimental conditions, they were instructed to watch the actions with one of two aims: to be able to recognize or to imitate them later. We found that differences in the meaning of the action, irrespective of the strategy used during observation, lead to different patterns of brain activity and clear left/right asymmetries. Meaningful actions strongly engaged the left hemisphere in frontal and temporal regions while meaningless actions involved mainly the right occipitoparietal pathway. Observing with the intent to recognize activated memory-encoding structures. In contrast, observation with the intent to imitate was associated with activation in the regions involved in the planning and in the generation of actions. Thus, the pattern of brain activation during observation of actions is dependent both on the nature of the required executive processing and the type of the extrinsic properties of the action presented.

The abilities to identify with others and to distinguish between self and other play a pivotal role in intersubjective transactions. Here, we marshall evidence from developmental science, social psychology and neuroscience (including clinical neuropsychology) that support the view of a common representation network (both at the computational and neural levels) between self and other. However, sharedness does not mean identicality, otherwise representations of self and others would completely overlap, and lead to confusion. We argue that self-awareness and agency are integral components for navigating within these shared representations. We suggest that within this shared neural network the inferior parietal cortex and the prefrontal cortex in the right hemisphere play a special role in interpersonal awareness.

Our ability to generate actions and to recognize actions performed by others is the bedrock of our social life. Behavioral evidence suggests that the processes underlying perception and action might share a common representational framework. That is, observers might understand the actions of another individual in terms of the same neural code that they use to produce the same actions themselves. What neurophysiological evidence, if any, supports such a hypothesis? In this article, brain imaging studies addressing this question are reviewed and examined in the light of the functional segregation of the perceptual mechanisms subtending visual recognition and those used for action. We suggest that there are not yet conclusive arguments for a clear neurophysiological substrate supporting a common coding between perception and action.

Empathy is the ability to experience and understand what others feel without confusion between oneself and others. Knowing what someone else is feeling plays a fundamental role in interpersonal interactions. In this paper, we articulate evidence from social psychology and cognitive neuroscience, and argue that empathy involves both emotion sharing (bottom-up information processing) and executive control to regulate and modulate this experience (top-down information processing), underpinned by specific and interacting neural systems. Furthermore, awareness of a distinction between the experiences of the self and others constitutes a crucial aspect of empathy. We discuss data from recent behavioral and functional neuroimaging studies with an emphasis on the perception of pain in others, and highlight the role of different neural mechanisms that underpin the experience of empathy, including emotion sharing, perspective taking, and emotion regulation.

Perspective-taking is a stepping stone to human empathy. When empathizing with another individual, one can imagine how the other perceives the situation and feels as a result. To what extent does imagining the other differs from imagining oneself in similar painful situations? In this functional magnetic resonance imaging experiment, participants were shown pictures of people with their hands or feet in painful or non-painful situations and instructed to imagine and rate the level of pain perceived from different perspectives. Both the Self's and the Other's perspectives were associated with activation in the neural network involved in pain processing, including the parietal operculum, anterior cingulate cortex (ACC; BA32) and anterior insula. However, the Self-perspective yielded higher pain ratings and involved the pain matrix more extensively in the secondary somatosensory cortex, the ACC (BA 24a′/24b′), and the insula proper. Adopting the perspective of the Other was associated with specific increase in the posterior cingulate/precuneus and the right temporo-parietal junction. These results show the similarities between Self- and Other-pain representation, but most interestingly they also highlight some distinctiveness between these two representations, which is a crucial aspect of human empathy. It may be what allows us to distinguish empathic responses to others versus our own personal distress. These findings are consistent with the view that empathy does not involve a complete Self–Other merging.

Whereas human pro-social behavior is often driven by empathic concern for another, it is unclear whether nonprimate mammals experience a similar motivational state. To test for empathically motivated pro-social behavior in rodents, we placed a free rat in an arena with a cagemate trapped in a restrainer. After several sessions, the free rat learned to intentionally and quickly open the restrainer and free the cagemate. Rats did not open empty or object-containing restrainers. They freed cagemates even when social contact was prevented. When liberating a cagemate was pitted against chocolate contained within a second restrainer, rats opened both restrainers and typically shared the chocolate. Thus, rats behave pro-socially in response to a conspecific's distress, providing strong evidence for biological roots of empathically motivated helping behavior.

The performance of subjects walking blindly to previously inspected visual targets (located at 5, 10 or 15 m from the subjects) was studied in 2 experiments. In Expt. 1, subjects selected as good visual imagers were instructed to build up a mental representation of the target. Then they had to either actually walk or imagine themselves walking to the target. Walking time was measured in both the actual and the mental performance. It was found that subjects took almost exactly the same time in the two conditions. Accuracy of these subjects was also measured in the actual walking task. They were found to make no direction errors and to slightly overshoot target location. Subjects from another, control, group, who received no instructions about visual imagery made much larger errors. In Expt. 2, actual and mental walking times were measured in the same subjects as in Expt. 1, while they carried a 25-kg weight on their shoulders. In this condition, actual walking time was the same as in Expt. 1, although mental walking time was found to increase systematically by about 30%. These results are discussed in terms of the neural parameters encoded in the motor program for actually executing or mentally performing an action.

This study investigated agency, the feeling of being causally involved in an action. This is the feeling that leads us to attribute an action to ourselves rather than to another person. We were interested in the effects of experimentally modulating this experience on brain areas known to be involved in action recognition and self-recognition. We used a device that allowed us to modify the subject's degree of control of the movements of a virtual hand presented on a screen. Four main conditions were used: (1) a condition where the subject had a full control of the movements of the virtual hand, (2) a condition where the movements of the virtual hand appeared rotated by 25 degrees with respect to the movements made by the subject, (3) a condition where the movements of the virtual hand appeared rotated by 50 degrees, and (4) a condition where the movements of the virtual hand were produced by another person and did not correspond to the subject's movements. The activity of two main brain areas appeared to be modulated by the degree of discrepancy between the movement executed and the movement seen on the screen. In the inferior part of the parietal lobe, specifically on the right side, the less the subject felt in control of the movements of the virtual hand, the higher the level of activation. A reverse covariation was observed in the insula. These results demonstrate that the level of activity of specific brain areas maps onto the experience of causing or controlling an action. The implication of these results for understanding pathological conditions is discussed.

In recent years, abundant evidence from behavioral and cognitive studies and functional-imaging experiments has indicated that individuals come to understand the emotional and affective states expressed by others with the help of the neural architecture that produces such states in themselves. Such a mechanism gives rise to shared representations, which constitutes one important aspect of empathy, although not the sole one. We suggest that other components, including people's ability to monitor and regulate cognitive and emotional processes to prevent confusion between self and other, are equally necessary parts of a functional model of empathy. We discuss data from recent functional-imaging studies in support of such a model and highlight the role of specific brain regions, notably the insula, the anterior cingulate cortex, and the right temporo-parietal region. Because this model assumes that empathy relies on dissociable information-processing mechanisms, it predicts a variety of structural or functional dysfunctions, depending on which mechanism is disrupted.

Both developmental and neurophysiological research suggest a common coding between perceived and generated actions. This shared representational network is innately wired in humans. We review psychological evidence concerning the imitative behaviour of newborn human infants. We suggest that the mechanisms involved in infant imitation provide the foundation for understanding that others are 'like me' and underlie the development of theory of mind and empathy for others. We also analyse functional neuroimaging studies that explore the neurophysiological substrate of imitation in adults. We marshal evidence that imitation recruits not only shared neural representations between the self and the other but also cortical regions in the parietal cortex that are crucial for distinguishing between the perspective of self and other. Imitation is doubly revealing: it is used by infants to learn about adults, and by scientists to understand the organization and functioning of the brain.

A large number of cognitive neuroscience studies point to the similarities in the neural circuits activated during the generation, imagination, as well as observation of one's own and other's behavior. Such findings support the shared representations account of social cognition, which is suggested to provide the basic mechanism for social interaction. Mental simulation may also be a representational tool to understand the self and others. However, successfully navigating these shared representations--both within oneself and between individuals--constitutes an essential functional property of any autonomous agent. It will be argued that self-awareness and agency, mediated by the temporoparietal (TPJ) area and the prefrontal cortex, are critical aspects of the social mind. Thus, differences as well as similarities between self and other representations at the neural level may be related to the degrees of self-awareness and agency. Overall, these data support the view that social cognition draws on both domain-general mechanisms and domain-specific embodied representations.

Motor imagery may be defined as a dynamic state during which representations of a given motor act are internally rehearsed in working memory without any overt motor output. What neural processes underlie the generation of motor imagery? This paper reviews physiological evidence from measurements of regional brain activity and from measurements of autonomic responses in normal subjects and behavioral observations from brain damaged patients. It is proposed that motor imagery shares neural mechanisms with processes used in motor control. This review emphasizes the importance of the prefrontal cortex and its connections to the basal ganglia in maintaining dynamic motor representations in working memory. This view fits with the general idea that the prefrontal cortex is responsible for the creation and maintenance of explicit representations that guide thought and action.

Perspective-taking is a complex cognitive process involved in social cognition. This positron emission tomography (PET) study investigated by means of a factorial design the interaction between the emotional and the perspective factors. Participants were asked to adopt either their own (first person) perspective or the (third person) perspective of their mothers in response to situations involving social emotions or to neutral situations. The main effect of third-person versus first-person perspective resulted in hemodynamic increase in the medial part of the superior frontal gyrus, the left superior temporal sulcus, the left temporal pole, the posterior cingulate gyrus, and the right inferior parietal lobe. A cluster in the postcentral gyrus was detected in the reverse comparison. The amygdala was selectively activated when subjects were processing social emotions, both related to self and other. Interaction effects were identified in the left temporal pole and in the right postcentral gyrus. These results support our prediction that the frontopolar, the somatosensory cortex, and the right inferior parietal lobe are crucial in the process of self/ other distinction. In addition, this study provides important building blocks in our understanding of social emotion processing and human empathy.

Several authors have demonstrated that theory of mind is associated with a cerebral pattern of activity involving the medial prefrontal cortex. This study was designed to determine the cerebral regions activated during attribution of intention to others, a task which requires theory-of-mind skills. Eight healthy subjects performed three nonverbal tasks using comic strips while PET scanning was performed. One condition required subjects to attribute intentions to the characters of the comic strips. The other two conditions involved only physical logic and knowledge about objects' properties: one condition involved characters, whereas the other only represented objects. The comparison of the attribution of intention condition with the physical logic with characters condition was associated with rCBF increases in the right middle and medial prefrontal cortex including Brodmann's area (BA) 9, the right inferior prefrontal cortex (BA 47), the right inferior temporal gyrus (BA 20), the left superior temporal gyrus (BA 38), the left cerebellum, the bilateral anterior cingulate, and the middle temporal gyri (BA 21). The comparison of the physical logic with characters condition and the physical logic without characters condition showed the activation of the lingual gyri (BA 17, 18, 19), the fusiform gyri (BA 37), the middle (BA 21) and superior (BA 22, 38) temporal gyri on both sides, and the posterior cingulate. These data suggest that attribution of intentions to others is associated with a complex cerebral activity involving the right medial prefrontal cortex when a nonverbal task is used. The laterality of this function is discussed.

This paper addresses the issue of the functional correlates of motor imagery, using mental chronometry, monitoring the autonomic responses and measuring cerebral blood flow in humans. The timing of mentally simulated actions closely mimic actual movement times. Autonomic responses during motor imagery parallel the autonomic responses to actual exercise. Cerebral blood flow increases are observed in the motor cortices involved in the programming of actual movement (i.e. premotor cortex, anterior cingulate, inferior parietal lobule and cerebellum). These three sources of data provide converging support for the hypothesis that imagined and executed actions share, to some extent, the same central structures.

Cooperation and competition are two basic modes of social cognition that necessitate monitoring of both one's own and others' actions, as well as adopting a specific mental set. In this fMRI, study individuals played a specially designed computer game, according to a set of predefined rules, either in cooperation with or in competition against another person. The hemodynamic response during these conditions was contrasted to that of the same subjects playing the game independently. Both cooperation and competition stances resulted in activation of a common frontoparietal network subserving executive functions, as well as the anterior insula, involved in autonomic arousal. Moreover, distinct regions were found to be selectively associated with cooperation and competition, notably the orbitofrontal cortex in the former and the inferior parietal and medial prefrontal cortices in the latter. This pattern reflects the different mental frameworks implicated in being cooperative versus competitive with another person. In accordance with evidence from evolutionary psychology as well as from developmental psychology, we argue that cooperation is a socially rewarding process and is associated with specific left medial orbitofrontal cortex involvement.

Empathy, which implies a shared interpersonal experience, is implicated in many aspects of social cognition, notably prosocial behavior, morality and the regulation of aggression. The purpose of this paper is to critically examine the current knowledge in developmental and affective neuroscience with an emphasis on the perception of pain in others. It will be argued that human empathy involves several components: affective arousal, emotion understanding and emotion regulation, each with different developmental trajectories. These components are implemented by a complex network of distributed, often recursively connected, interacting neural regions including the superior temporal sulcus, insula, medial and orbitofrontal cortices, amygdala and anterior cingulate cortex, as well as autonomic and neuroendocrine processes implicated in social behaviors and emotional states. Decomposing the construct of empathy into subcomponents that operate in conjunction in the healthy brain and examining their developmental trajectory provides added value to our current approaches to understanding human development. It can also benefit our understanding of both typical and atypical development.

The role of the primary motor cortex (M1) during mental simulation of movement is open to debate. In the present study, functional magnetic resonance imaging (fMRI) signals were measured in normal right-handed subjects during actual and mental execution of a finger-to-thumb opposition task with either the right or the left hand. There were no significant differences between the two hands with either execution or simulation. A significant involvement of contralateral M1 (30% of the activity found during execution) was detected in four of six subjects. Premotor cortex (PM) and the rostral part of the posterior SMA were activated bilaterally during motor imagery. These findings support the hypothesis that motor imagery involves virtually all stages of motor control.

The psychological construct of empathy refers to an intersubjective induction process by which positive and negative emotions are shared, without losing sight of whose feelings belong to whom. Empathy can lead to personal distress or to empathic concern (sympathy). The goal of this paper is to address the underlying cognitive processes and their neural underpinnings that constitute empathy within a developmental neuroscience perspective. In addition, we focus on how these processes go awry in developmental disorders marked by impairments in social cognition, such as autism spectrum disorder, and conduct disorder. We argue that empathy involves both bottom-up and top-down information processing, underpinned by specific and interacting neural systems. We discuss data from developmental psychology as well as cognitive neuroscience in support of such a model, and highlight the impact of neural dysfunctions on social cognitive developmental behavior. Altogether, bridging developmental science and cognitive neuroscience helps approach a more complete understanding of social cognition. Synthesizing these two domains also contributes to a better characterization of developmental psychopathologies that impacts the development of effective treatment strategies.

Positron emission tomography (PET) was used to explore the neural correlates of a potential involvement of motor representation during the perception of visually presented objects with different tasks. The main result of this study was that the perception of objects, irrespective of the task (judgement of the vertical orientation, motor imagery, and silent generation of the noun or of the corresponding action verb), versus perception of non-objects, was associated with rCBF increases in a common set of cortical regions. The occipito-temporal junction, the inferior parietal lobule, the SMA-proper, the pars triangularis in the inferior frontal gyrus, the dorsal and ventral precentral gyrus were engaged in the left hemisphere. The ipsilateral cerebellum was also involved. These activations are congruent with the idea of an involvement of motor representation already during the perception of object and thus provide neurophysiological evidence that the perception of objects automatically affords actions that can be made toward them. Besides this common set of cortical areas, each task engaged specific regions.

The physiological basis of mental states can be effectively studied by combining cognitive psychology with human neuroscience. Recent research has employed mental motor imagery in normal and brain-damaged subjects to decipher the content and the structure of covert processes preceding the execution of action. The mapping of brain activity during motor imagery discloses a pattern of activation similar to that of an executed action.

Watching or imagining other people experiencing pain activates the central nervous system's pain matrix in the observer. Without emotion regulation skills, repeated exposure to the suffering of others in healthcare professionals may be associated with the adverse consequences of personal distress, burnout and compassion fatigue, which are detrimental to their wellbeing. Here, we recorded event-related potentials (ERP) from physicians and matched controls as they were presented with visual stimuli depicting body parts pricked by a needle (pain) or touched by a Q-tip (no-pain). The results showed early N110 differentiation between pain and no-pain over the frontal area as well as late P3 over the centro-parietal regions were observed in the control participants. In contrast, no such early and late ERP responses were detected in the physicians. Our results indicate that emotion regulation in physicians has very early effects, inhibiting the bottom-up processing of the perception of pain in others. It is suggested that physicians' down-regulation of the pain response dampens their negative arousal in response to the pain of others and thus may have many beneficial consequences including freeing up cognitive resources necessary for being of assistance.

Empathy reflects the natural ability to perceive and be sensitive to the emotional states of others, coupled with a motivation to care for their well-being. It has evolved in the context of parental care for offspring, as well as within kinship bonds, to help facilitate group living. In this paper, we integrate the perspectives of evolution, animal behaviour, developmental psychology, and social and clinical neuroscience to elucidate our understanding of the proximate mechanisms underlying empathy. We focus, in particular, on processing of signals of distress and need, and their relation to prosocial behaviour. The ability to empathize, both in animals and humans, mediates prosocial behaviour when sensitivity to others' distress is paired with a drive towards their welfare. Disruption or atypical development of the neural circuits that process distress cues and integrate them with decision value leads to callous disregard for others, as is the case in psychopathy. The realization that basic forms of empathy exist in non-human animals is crucial for gaining new insights into the underlying neurobiological and genetic mechanisms of empathy, enabling translation towards therapeutic and pharmacological interventions.

Perceiving the pain of others activates a large part of the pain matrix in the observer [1]. Because this shared neural representation can lead to empathy or personal distress [2, 3], regulatory mechanisms must operate in people who inflict painful procedures in their practice with patient populations in order to prevent their distress from impairing their ability to be of assistance. In this functional magnetic resonance imaging MRI study, physicians who practice acupuncture were compared to naive participants while observing animated visual stimuli depicting needles being inserted into different body parts, including the mouth region, hands, and feet. Results indicate that the anterior insula somatosensory cortex, periaqueducal gray, and anterior cingulate cortex were significantly activated in the control group, but not in the expert group, who instead showed activation of the medial and superior prefrontal cortices and the temporoparietal junction, involved in emotion regulation and theory of mind.

This study was designed to investigate mentally simulated actions in a virtual reality environment. Naive human subjects (n = 15) were instructed to imagine themselves walking in a three-dimensional virtual environment toward gates of different apparent widths placed at three different apparent distances. Each subject performed nine blocks of six trials in a randomised order. The response time (reaction time and mental walking time) was measured as the duration between an acoustic go signal and a motor signal produced by the subject. There was a combined effect on response time of both gate width and distance. Response time increased for decreasing apparent gate widths when the gate was placed at different distances. These results support the notion that mentally simulated actions are governed by central motor rules.

The observation of a patient (A.T.) with a bilateral posterior parietal lesion of vascular origin is reported. A.T. presented a bilateral (more marked on the right) deficit in grasping simple objects (neutral cylindrical dowels) without deficit in reaching toward the location of these objects. The major symptom was an exaggerated anticipatory opening of the fingers with poor correlation with object size, resulting in awkward grasps. It was present both when the hand was visible to the subject and when it was not. This deficit was much less marked if the neutral objects were replaced by usual objects of the same sizes. Finally, in tasks where she had to indicate with her fingers the size of visual objects presented as virtual images through a mirror, or the size of imagined usual objects, A.T. performed normally. These results are discussed within the framework of a dual representation of objects. Only the "pragmatic" representation for steering object-oriented actions would be impaired in this patient as a result of posterior parietal damage. By contrast the semantic representation for object identification would be intact.

Imitation is a natural mechanism involving perception-action coupling which plays a central role in the development of understanding that other people, like the self, are mental agents. PET was used to examine the hemodynamic changes occurring in a reciprocal imitation paradigm. Eighteen subjects (a) imitated the actions of the experimenter, (b) had their actions imitated by the experimenter, (c) freely produced actions, or (d) freely produced actions while watching different actions made by the experimenter. In a baseline condition, subjects simply watched the experimenter's actions. Specific increases were detected in the left STS and in the inferior parietal cortex in conditions involving imitation. The left inferior parietal is specifically involved in producing imitation, whereas the right homologous region is more activated when one's own actions are imitated by another person. This pattern of results suggests that these regions play a specific role in distinguishing internally produced actions from those generated by others.

Empathy is thought to play a key role in motivating prosocial behavior, guiding our preferences and behavioral responses, and providing the affective and motivational base for moral development. While these abilities have traditionally been examined using behavioral methods, recent work in evolutionary biology, developmental and cognitive neuroscience has begun to shed light on the neural circuitry that instantiate them. The purpose of this article is to critically examine the current knowledge in the field of affective neuroscience and provide an integrative and comprehensive view of the computational mechanisms that underlie empathy. This framework is of general interest and relevance for theory as well as for assisting future research in the domains of affective developmental neuroscience and psychopathology.

Empathy is a concept central to psychiatry, psychotherapy and clinical psychology. The construct of empathy involves not only the affective experience of the other person's actual or inferred emotional state but also some minimal recognition and understanding of another's emotional state. It is proposed, in the light of multiple levels of analysis including social psychology, cognitive neuroscience and clinical neuropsychology, a model of empathy that involves both bottom-up and top-down information processing underpinned by parallel and distributed computational mechanisms. The predictive validity of this model is explored with reference to clinical conditions. As many psychiatric conditions are associated with deficits or even lack of empathy, we discuss a limited number of these disorders including psychopathy/antisocial personality disorders, borderline and narcissistic personality disorders, autistic spectrum disorders, and alexithymia. We argue that future clinical investigations of empathy disorders can only be informative if behavioral, dispositional and biological factors are combined.

Whether emotion is a source of moral judgments remains controversial. This study combined neurophysiological measures, including functional magnetic resonance imaging, eye-tracking, and pupillary response with behavioral measures assessing affective and moral judgments across age. One hundred and twenty-six participants aged between 4 and 37 years viewed scenarios depicting intentional versus accidental actions that caused harm/damage to people and objects. Morally, salient scenarios evoked stronger empathic sadness in young participants and were associated with enhanced activity in the amygdala, insula, and temporal poles. While intentional harm was evaluated as equally wrong across all participants, ratings of deserved punishments and malevolent intent gradually became more differentiated with age. Furthermore, age-related increase in activity was detected in the ventromedial prefrontal cortex in response to intentional harm to people, as well as increased functional connectivity between this region and the amygdala. Our study provides evidence that moral reasoning involves a complex integration between affective and cognitive processes that gradually changes with age and can be viewed in dynamic transaction across the course of ontogenesis. The findings support the view that negative emotion alerts the individual to the moral salience of a situation by bringing discomfort and thus can serve as an antecedent to moral judgment.

Recent neuroscientific evidence suggests that empathy for pain activates similar neural representations as the first-hand experience of pain. However, empathy is not an all-or-none phenomenon but it is strongly malleable by interpersonal, intrapersonal and situational factors. This study investigated how two different top-down mechanisms - attention and cognitive appraisal - affect the perception of pain in others and its neural underpinnings.We performed one behavioral (N = 23) and two functional magnetic resonance imaging (fMRI) experiments (N = 18). In the first fMRI experiment, participants watched photographs displaying painful needle injections, and were asked to evaluate either the sensory or the affective consequences of these injections. The role of cognitive appraisal was examined in a second fMRI experiment in which participants watched injections that only appeared to be painful as they were performed on an anesthetized hand. Perceiving pain in others activated the affective-motivational and sensory-discriminative aspects of the pain matrix. Activity in the somatosensory areas was specifically enhanced when participants evaluated the sensory consequences of pain. Perceiving non-painful injections into the anesthetized hand also led to signal increase in large parts of the pain matrix, suggesting an automatic affective response to the putatively harmful stimulus. This automatic response was modulated by areas involved in self/other distinction and valence attribution - including the temporo-parietal junction and medial orbitofrontal cortex.Our findings elucidate how top-down control mechanisms and automatic bottom-up processes interact to generate and modulate other-oriented responses. They stress the role of cognitive processing in empathy, and shed light on how emotional and bodily awareness enable us to evaluate the sensory and affective states of others.

To better understand clinical empathy and what factors can undermine its experience and outcome in care-giving settings, a large-scale study was conducted with 7,584 board certified practicing physicians. Online validated instruments assessing different aspects of empathy, distress, burnout, altruistic behavior, emotional awareness, and well-being were used. Compassion satisfaction was strongly associated with empathic concern, perspective taking and altruism, while compassion fatigue (burnout and secondary traumatic stress) was more closely related to personal distress and alexithymia. Gender had a highly selective effect on empathic concern, with women displaying higher values, which led to a wide array of negative and devalued feelings. Years of experience did not influence dispositional measures per se after controlling for the effect of age and gender. Participants who experienced compassion fatigue with little to no compassion satisfaction showed the highest scores on personal distress and alexithymia as well as the strongest indicators of compassion fatigue. Physicians who have difficulty regulating their negative arousal and describing and identifying emotions seem to be more prone to emotional exhaustion, detachment, and a low sense of accomplishment. On the contrary, the ability to engage in self-other awareness and regulate one's emotions and the tendency to help others, seem to contribute to the sense of compassion that comes from assisting patients in clinical practice.

Is it important to adopt the perspective of the model when learning a new skill? Is the “mirror system” equally involved when the teacher is facing or side-by-side with students? In this functional MRI study, we measured the cerebral hemodynamic changes in participants who watched video-clips depicting simple hand or foot actions. The participants either watched passively or imitated these actions. Half the video-clips depicted actions filmed from the perspective of the participant (1st-person perspective) and half from a frontal view as if watching someone else (3rd-person perspective). Behavioral results showed that latency to imitate was significantly shorter for the 1st-person perspective than the 3rd-person perspective. Functional imaging results demonstrate that the observation of intransitive actions engaged primary visual and extrastriate visual areas, but not the premotor cortex. Imitation vs. observation of actions yielded enhanced signal in the contralateral somatosensory and motor cortices, cerebellum, left inferior parietal lobule and superior parietal cortex, and left ventral premotor cortex. Activity in the lateral occipital cortex around the extrastriate body area was significantly enhanced during imitation, as compared to observation of actions confirming that this region involvement reaches beyond the perception of body parts. Moreover, comparisons of the two visual perspectives showed more activity in the left sensory–motor cortex for 1st-person, even during observation alone, and in the lingual gyrus for 3rd-person perspective. These findings suggest that the 1st-person perspective is more tightly coupled to the sensory-motor system than the 3rd-person perspective, which requires additional visuospatial transformation.

Measurement of cardiac and respiratory activity during mental simulation of locomotion at increasing speed revealed a covariation of heart rate and pulmonary ventilation with the degree of imagined effort. The degree of vegetative activation of a subject mentally running at 12 km/h was comparable to that of a subject actually walking at 5 km/h. This effect cannot be explained by an increase in peripheral (e.g. muscular) metabolic demands. Indeed, oxygen uptake decreased during motor imagery. This finding is suggestive of a commonality of neural structures responsible for mental imagery of movement and those responsible for programming actual movement. In addition, it provides an quantifiable way of testing mental imagery in relation to movement by using easily accessible biological markers.

Social communication requires shared representations as well as a cognitive flexibility for successful interactions between self and other. What neural mechanisms underlie the ability to distinguish between our own perspective vs. the perspective of others at a conceptual level? In this PET study subjects who were medical students were asked according to the experimental conditions to respond to a list of health-related questions, taking either their own perspective or the perspective of a 'lay person'. Third-person perspective as compared to first-person perspective was associated with activation in the medial part of the superior frontal gyrus, in the left superior temporal sulcus, in the left temporal pole and in the right inferior parietal lobe. The reverse comparison revealed a specific activation in the postcentral gyrus for the first-person conceptual perspective. This study provides congruent results at the conceptual level with previous studies investigating the neural correlates of self/other distinction at the motor level, and opens a new area of research in which conceptual cognition can be viewed in the continuity of motor cognition.

The ontogeny of human empathy is better understood with reference to the evolutionary history of the social brain. Empathy has deep evolutionary, biochemical, and neurological underpinnings. Even the most advanced forms of empathy in humans are built on more basic forms and remain connected to core mechanisms associated with affective communication, social attachment, and parental care. In this paper, we argue that it is essential to consider empathy within a neurodevelopmental framework that recognizes both the continuities and changes in socioemotional understanding from infancy to adulthood. We bring together neuroevolutionary and developmental perspectives on the information processing and neural mechanisms underlying empathy and caring, and show that they are grounded in multiple interacting systems and processes. Moreover, empathy in humans is assisted by other abstract and domain-general high-level cognitive abilities such as executive functions, mentalizing and language, as well as the ability to differentiate another's mental states from one's own, which expand the range of behaviors that can be driven by empathy.

Abstract This experiment was designed to investigate the neural network engaged by the perception of human movements using positron emission tomography. Perception of meaningful and of meaningless hand actions without any purpose was contrasted with the perception of the same kind of stimuli with the goal to imitate them later. A condition that consisted of the perception of stationary hands served as a baseline level. Perception of meaningful actions and meaningless actions without any aim was associated with activation of a common set of cortical regions. In both hemispheres, the occipito-temporal junction (Ba 37/19) and the superior occipital gyrus (Ba 19) were involved. In the left hemisphere, the middle temporal gyrus (Ba 21) and the inferior parietal lobe (Ba 40) were found to be activated. These regions are interpreted as related to the analysis of hand movements. The precentral gyrus, within the area of hand representation (Ba 4), was activated in the left hemisphere. In addition to this common network, meaningful and meaningless movements engaged specific networks, respectively: meaningful actions were associated with activations mainly located in the left hemisphere in the inferior frontal gyrus (Ba 44/45) and the fusiform gyrus (Ba 38/20), whereas meaningless actions involved the dorsal pathway (inferior parietal lobe, Ba 40 and superior parietal lobule, Ba 7) bilaterally and the right cerebellum. In contrast, meaningful and meaningless actions shared almost the same network when the aim of the perception was to im itate. Activations were located in the right cerebellum and bilaterally in the dorsal pathway reaching the prem otor cortex. Additional bilateral activations were located in the SMA and in the orbitofrontal cortex during observation of meaningful actions.

Morality and empathy are fundamental components of human nature across cultures. However, the wealth of empirical findings from developmental, behavioral, and social neuroscience demonstrates a complex relation between morality and empathy. At times, empathy guides moral judgment, yet other times empathy can interfere with it. To better understand such relations, we propose abandoning the catchall term of empathy in favor of more precise concepts, such as emotional sharing, empathic concern, and affective perspective-taking.

Because awareness of emotional states in the self is a prerequisite to recognizing such states in others, alexithymia (ALEX), difficulty in identifying and expressing one's own emotional states, should involve impairment in empathy. Using functional magnetic resonance imaging (fMRI), we compared an ALEX group (n = 16) and a non-alexithymia (non-ALEX) group (n = 14) for their regional hemodynamic responses to the visual perception of pictures depicting human hands and feet in painful situations. Subjective pain ratings of the pictures and empathy-related psychological scores were also compared between the 2 groups. The ALEX group showed less cerebral activation in the left dorsolateral prefrontal cortex (DLPFC), the dorsal pons, the cerebellum, and the left caudal anterior cingulate cortex (ACC) within the pain matrix. The ALEX group showed greater activation in the right insula and inferior frontal gyrus. Furthermore, alexithymic participants scored lower on the pain ratings and on the scores related to mature empathy. In conclusion, the hypofunction in the DLPFC, brain stem, cerebellum, and ACC and the lower pain-rating and empathy-related scores in ALEX are related to cognitive impairments, particularly executive and regulatory aspects, of emotional processing and support the importance of self-awareness in empathy.

Positron emission tomography (PET) was used to investigate the neural correlates of feeling sympathy for someone else (i.e. the affinity, association, or relationship between persons wherein whatever affects one similarly affects the other). While undergoing PET scans, subjects were presented with a series of video-clips showing individuals (who were semi-professional stage actors) telling sad and neutral stories, as if they had personally experienced them. These stories were told with either congruent or incongruent motor expression of emotion (MEE). At the end of each movie, subjects were asked to rate the mood of the communicator and also how likable they found that person. Watching sad stories versus neutral stories was associated with increased activity in emotion processing-related structures, as well as in a set of cortical areas that belong to a "shared representation" network, including the right inferior parietal cortex. Motor expression of emotion, regardless of the narrative content of the stories, resulted in a specific regional cerebral blood flow (rCBF) increase in the left inferior frontal gyrus. The condition of mismatch between the narrative content of the stories and the motor expression of emotion elicited a significant skin conductance response and strong rCBF increase in the ventromedial prefrontal cortex and superior frontal gyrus which are involved in dealing with social conflict. Taken together, these results are consistent with a model of feeling sympathy that relies on both the shared representation and the affective networks. Interestingly, this network was not activated when subjects watched inappropriate social behavior.

Measurements in man of regional cerebral blood flow (rCBF) have demonstrated a number of cortical and subcortical events coupled to sensory stimulation or motor performance. It has also been shown that local activity changes take place in the cortex during 'pure' mental activity such as motor imagery (unaccompanied by sensory input or motor output). Thus, our group has previously shown that imagination of hand movements gives predominantly a frontal cortical rCBF activation while the corresponding hand movement activates the rolandic hand area mainly. In this paper we report tomographic rCBF measurements with a 133-Xenon SPECT technique during imagined tennis movements and silent counting. Both procedures gave rise to a significant cerebellar activation in addition to cortical rCBF changes. Apparently, the cerebellum may participate in pure mental activity. It possibly plays a role for the temporal organization of neuronal events related to cognition.

There is strong evidence that empathy has deep evolutionary, biochemical, and neurological underpinnings. Even the most advanced forms of empathy in humans are built on more basic forms and remain connected to core mechanisms associated with affective communication, social attachment, and parental care. Social neuroscience has begun to examine the neurobiological mechanisms that instantiate empathy, especially in response to signals of distress and pain, and how certain dispositional and contextual moderators modulate its experience. Functional neuroimaging studies document a circuit that responds to the perception of others' distress. Activation of this circuit reflects an aversive response in the observer, and this information may act as a trigger to inhibit aggression or prompt motivation to help. Moreover, empathy in humans is assisted by other domain-general high-level cognitive abilities, such as executive functions, mentalizing, and language, which expand the range of behaviors that can be driven by empathy.

This study compared the temporal organization of graphic movements executed either actually or mentally. Six subjects had to perform two tasks, writing a sentence and drawing a cube, either as a real performance or as an imagined one, with either the right or the left hand, and with either a small or a large tracing amplitude. In the same subject, for the same hand, mental and actual movement times were both very stable and very close from trial to trial regardless of the tracing amplitude. Thus, mental movements mimic closely real movements in their temporal organization and are likely to involve the same planning program.

aÉcole de psychologie and CIRRIS and CRULRG, Université Laval, Québec, Que., Canada bDépartement de Stomatologie, Université de Montréal, Montréal, Que., Canada cDepartment of Psychology and Center for Cognitive and Social Neuroscience, The University of Chicago, Chicago, IL, USA *Corresponding author. Tel.: +1 418 656 2131 ext.5151; fax: +1 418 529 3548. E-mail address:[email protected] Submitted July 17, 2006; revised September 6, 2006; accepted September 8, 2006.

ORIGINAL RESEARCH article Front. Hum. Neurosci., 24 September 2013Sec. Cognitive Neuroscience https://doi.org/10.3389/fnhum.2013.00489

Abstract Empathy and sympathy play crucial roles in much of human social interaction and are necessary components for healthy coexistence. Sympathy is thought to be a proxy for motivating prosocial behavior and providing the affective and motivational base for moral development. The purpose of the present study was to use functional MRI to characterize developmental changes in brain activation in the neural circuits underpinning empathy and sympathy. Fifty‐seven individuals, whose age ranged from 7 to 40 years old, were presented with short animated visual stimuli depicting painful and non‐painful situations. These situations involved either a person whose pain was accidentally caused or a person whose pain was intentionally inflicted by another individual to elicit empathic (feeling as the other) or sympathetic (feeling concern for the other) emotions, respectively. Results demonstrate monotonic age‐related changes in the amygdala, supplementary motor area, and posterior insula when participants were exposed to painful situations that were accidentally caused. When participants observed painful situations intentionally inflicted by another individual, age‐related changes were detected in the dorsolateral prefrontal and ventromedial prefrontal cortex, with a gradual shift in that latter region from its medial to its lateral portion. This pattern of activation reflects a change from a visceral emotional response critical for the analysis of the affective significance of stimuli to a more evaluative function. Further, these data provide evidence for partially distinct neural mechanisms subserving empathy and sympathy, and demonstrate the usefulness of a developmental neurobiological approach to the new emerging area of moral neuroscience.

Because youth with aggressive conduct disorder (CD) often inflict pain on others, it is important to determine if they exhibit atypical empathic responses to viewing others in pain. In this initial functional magnetic resonance imaging (fMRI) study, eight adolescents with aggressive CD and eight matched controls with no CD symptoms were scanned while watching animated visual stimuli depicting other people experiencing pain or not experiencing pain. Furthermore, these situations involved either an individual whose pain was caused by accident or an individual whose pain was inflicted on purpose by another person. After scanning, participants rated how painful the situations were. In both groups the perception of others in pain was associated with activation of the pain matrix, including the ACC, insula, somatosensory cortex, supplementary motor area and periaqueductal gray. The pain matrix was activated to a specific extent in participants with CD, who also showed significantly greater amygdala, striatal, and temporal pole activation. When watching situations in which pain was intentionally inflicted, control youth exhibited signal increase in the medial prefrontal cortex, lateral orbitofrontal cortex, and right temporo-parietal junction, whereas youth with CD only exhibited activation in the insula and precentral gyrus. Furthermore, connectivity analyses demonstrated that youth with CD exhibited less amygdala/prefrontal coupling when watching pain inflicted by another than did control youth. These preliminary findings suggest that youth with aggressive CD exhibit an atypical pattern of neural response to viewing others in pain that should be explored in further studies.

Abstract Prior research has shown that perceived social isolation (loneliness) motivates people to attend to and connect with others but to do so in a self-protective and paradoxically self-defeating fashion. Although recent research has shed light on the neural correlates of social perception, cooperation, empathy, rejection, and love, little is known about how individual differences in loneliness relate to neural responses to social and emotional stimuli. Using functional magnetic resonance imaging, we show that there are at least two neural mechanisms differentiating social perception in lonely and nonlonely young adults. For pleasant depictions, lonely individuals appear to be less rewarded by social stimuli, as evidenced by weaker activation of the ventral striatum to pictures of people than of objects, whereas nonlonely individuals showed stronger activation of the ventral striatum to pictures of people than of objects. For unpleasant depictions, lonely individuals were characterized by greater activation of the visual cortex to pictures of people than of objects, suggesting that their attention is drawn more to the distress of others, whereas nonlonely individuals showed greater activation of the right and left temporo-parietal junction to pictures of people than of objects, consistent with the notion that they are more likely to reflect spontaneously on the perspective of distressed others.

The complexities of the brain and nervous system make neuroscience an inherently interdisciplinary pursuit, one that comprises disparate basic, clinical, and applied disciplines. Behavioral neuroscientists approach the brain and nervous system as instruments of sensation and response; cognitive neuroscientists view the same systems as a solitary computer with a focus on representations and processes. The Oxford Handbook of Social Neuroscience marks the emergence of a third broad perspective in this field. Social neuroscience emphasizes the functions that emerge through the coaction and interaction of conspecifics, the neural mechanisms that underlie these functions, and the commonality and differences across social species and superorganismal structures. With an emphasis on the neural, hormonal, cellular, and genetic mechanisms underlying social behavior, social neuroscience places emphasis on the associations and influences between social and biological levels of organization. This complex interdisciplinary perspective demands theoretical, methodological, statistical, and inferential rigor to effectively integrate basic, clinical, and applied perspectives on the nervous system and brain.

This investigation combined behavioral and functional neuroimaging measures to explore whether perception of pain is modulated by the target's stigmatized status and whether the target bore responsibility for that stigma. During fMRI scanning, participants were exposed to a series of short video clips featuring age-matched individuals experiencing pain who were (a) similar to the participant (healthy), (b) stigmatized but not responsible for their stigmatized condition (infected with AIDS as a result of an infected blood transfusion), or (c) stigmatized and responsible for their stigmatized condition (infected with AIDS as a result of intravenous drug use). Explicit pain and empathy ratings for the targets were obtained outside of the MRI environment, along with a variety of implicit and explicit measures of AIDS bias. Results showed that participants were significantly more sensitive to the pain of AIDS transfusion targets as compared with healthy and AIDS drug targets, as evidenced by significantly higher pain and empathy ratings during video evaluation and significantly greater hemodynamic activity in areas associated with pain processing (i.e., right anterior insula, anterior midcingulate cortex, periaqueductal gray). In contrast, significantly less activity was observed in the anterior midcingulate cortex for AIDS drug targets as compared with healthy controls. Further, behavioral differences between healthy and AIDS drug targets were moderated by the extent to which participants blamed AIDS drug individuals for their condition. Controlling for both explicit and implicit AIDS bias, the more participants blamed these targets, the less pain they attributed to them as compared with healthy controls. The present study reveals that empathic resonance is moderated early in information processing by a priori attitudes toward the target group.

Abstract Previous research on the neural underpinnings of empathy has been limited to affective situations experienced in a similar way by an observer and a target individual. In daily life we also interact with people whose responses to affective stimuli can be very different from our own. How do we understand the affective states of these individuals? We used functional magnetic resonance imaging to assess how participants empathize with the feelings of patients who reacted with no pain to surgical procedures but with pain to a soft touch. Empathy for pain of these patients activated the same areas (insula, medial/anterior cingulate cortex) as empathy for persons who responded to painful stimuli in the same way as the observer. Empathy in a situation that was aversive only for the observer but neutral for the patient recruited areas involved in self–other distinction (dorsomedial prefrontal cortex) and cognitive control (right inferior frontal cortex). In addition, effective connectivity between the latter and areas implicated in affective processing was enhanced. This suggests that inferring the affective state of someone who is not like us can rely upon the same neural structures as empathy for someone who is similar to us. When strong emotional response tendencies exist though, these tendencies have to be overcome by executive functions. Our results demonstrate that the fronto-cortical attention network is crucially involved in this process, corroborating that empathy is a flexible phenomenon which involves both automatic and controlled cognitive mechanisms. Our findings have important implications for the understanding and promotion of empathy, demonstrating that regulation of one's egocentric perspective is crucial for understanding others.

Background Psychopathic traits are associated with increases in antisocial behaviors such as aggression and are characterized by reduced empathy for others' distress. This suggests that psychopathic traits may also impair empathic pain sensitivity. However, whether psychopathic traits affect responses to the pain of others versus the self has not been previously assessed. Method We used whole‐brain functional magnetic resonance imaging to measure neural activation in 14 adolescents with oppositional defiant disorder or conduct disorder and psychopathic traits, as well as 21 healthy controls matched on age, gender, and intelligence. Activation in structures associated with empathic pain perception was assessed as adolescents viewed photographs of pain‐inducing injuries. Adolescents imagined either that the body in each photograph was their own or that it belonged to another person. Behavioral and neuroimaging data were analyzed using random‐effects analysis of variance. Results Youths with psychopathic traits showed reduced activity within regions associated with empathic pain as the depicted pain increased. These regions included rostral anterior cingulate cortex, ventral striatum (putamen), and amygdala. Reductions in amygdala activity particularly occurred when the injury was perceived as occurring to another. Empathic pain responses within both amygdala and rostral anterior cingulate cortex were negatively correlated with the severity of psychopathic traits as indexed by PCL : YV scores. Conclusions Youths with psychopathic traits show less responsiveness in regions implicated in the affective response to another's pain as the perceived intensity of this pain increases. Moreover, this reduced responsiveness appears to predict symptom severity.

In mammals, helping is preferentially provided to members of one’s own group. Yet, it remains unclear how social experience shapes pro-social motivation. We found that rats helped trapped strangers by releasing them from a restrainer, just as they did cagemates. However, rats did not help strangers of a different strain, unless previously housed with the trapped rat. Moreover, pair-housing with one rat of a different strain prompted rats to help strangers of that strain, evidence that rats expand pro-social motivation from one individual to phenotypically similar others. To test if genetic relatedness alone can motivate helping, rats were fostered from birth with another strain and were not exposed to their own strain. As adults, fostered rats helped strangers of the fostering strain but not rats of their own strain. Thus, strain familiarity, even to one’s own strain, is required for the expression of pro-social behavior.

Why do people tend to care for upholding principles of justice? This study examined the association between individual differences in the affective, motivational and cognitive components of empathy, sensitivity to justice, and psychopathy in participants (N 265) who were also asked to rate the permissibility of everyday moral situations that pit personal benefit against moral standards of justice. Counter to common sense, emotional empathy was not associated with sensitivity to injustice for others. Rather, individual differences in cognitive empathy and empathic concern predicted sensitivity to justice for others, as well as the endorsement of moral rules. Psychopathy coldheartedness scores were inversely associated with motivation for justice. Moreover, hierarchical multiple linear regression analysis revealed that self-focused and other-focused orientations toward justice had opposing influences on the permissibility of moral judgments. High scores on psychopathy were associated with less moral condemnation of immoral behavior. Together, these results contribute to a better understanding of the information processing mechanisms underlying justice motivation, and may guide interventions designed to foster justice and moral behavior. In order to promote justice motivation, it may be more effective to encourage perspective taking and reasoning than emphasizing emotional sharing with the misfortune of others.

In mammals, empathy is crucial for living in social groups and caring for others. In this paper, we consider the structural and functional organization of empathy. We propose that empathy subsumes a variety of neurobiological processes and partially dissociable information processing subsystems, each of which has a unique evolutionary history. Even the most advanced and flexible forms of empathy in humans are built on more basic forms and remain connected to core subcortical and neurohormonal mechanisms associated with affective communication, parental care and social attachment processes. Considering empathy within a framework that recognizes both the continuities and the changes within a phylogenetic perspective provides a richer understanding of empathy and related neurobehavioral processes.

<h3>Importance</h3>A marked lack of empathy is a hallmark characteristic of individuals with psychopathy. However, neural processes associated with empathic processing have not yet been directly examined in psychopathy, especially in response to the perception of other people in pain and distress.<h3>Objective</h3>To identify potential differences in patterns of neural activity in incarcerated individuals with psychopathy and incarcerated persons serving as controls during the perception of empathy-eliciting stimuli depicting other people experiencing pain.<h3>Design</h3>In a case-control study, brain activation patterns elicited by dynamic stimuli depicting individuals being harmed and facial expressions of pain were compared between incarcerated individuals with psychopathy and incarcerated controls.<h3>Setting</h3>Participants were scanned on the grounds of a correctional facility using the Mind Research Network's mobile 1.5-T magnetic resonance imaging system.<h3>Participants</h3>Eighty incarcerated men were classified according to scores on the Hare Psychopathy Checklist–Revised (PCL-R) as high (27 men; PCL-R, ≥30), intermediate (28 men; PCL-R, 21-29), or low (25 men; PCL-R, ≤20) levels of psychopathy.<h3>Main Outcome Measure</h3>Neurohemodynamic response to empathy-eliciting dynamic scenarios revealed by functional magnetic resonance imaging.<h3>Results</h3>Participants in the psychopathy group exhibited significantly less activation in the ventromedial prefrontal cortex, lateral orbitofrontal cortex, and periaqueductal gray relative to controls but showed greater activation in the insula, which was positively correlated with scores on both PCL-R factors 1 and 2.<h3>Conclusions and Relevance</h3>In response to pain and distress cues expressed by others, individuals with psychopathy exhibit deficits in the ventromedial prefrontal cortex and orbitofrontal cortex regardless of stimulus type and display selective impairment in processing facial cues of distress in regions associated with cognitive mentalizing. A better understanding of the neural responses to empathy-eliciting stimuli in psychopathy is necessary to inform intervention programs.

In the past decade, a flurry of empirical and theoretical research on morality and empathy has taken place, and interest and usage in the media and the public arena have increased. At times, in both popular culture and academia, morality and empathy are used interchangeably, and quite often the latter is considered to play a foundational role for the former. In this article, we argue that although there is a relationship between morality and empathy, it is not as straightforward as apparent at first glance. Moreover, it is critical to distinguish among the different facets of empathy (emotional sharing, empathic concern, and perspective taking), as each uniquely influences moral cognition and predicts differential outcomes in moral behavior. Empirical evidence and theories from evolutionary biology as well as developmental, behavioral, and affective and social neuroscience are comprehensively integrated in support of this argument. The wealth of findings illustrates a complex and equivocal relationship between morality and empathy. The key to understanding such relations is to be more precise on the concepts being used and, perhaps, abandoning the muddy concept of empathy.

Empathy reflects an innate ability to perceive and be sensitive to the emotional states of others coupled with a motivation to care for their wellbeing. It has evolved in the context of parental care for offspring as well as within kinship. Current work demonstrates that empathy is underpinned by circuits connecting the brainstem, amygdala, basal ganglia, anterior cingulate cortex, insula and orbitofrontal cortex, which are conserved across many species. Empirical studies document that empathetic reactions emerge early in life, and that they are not automatic. Rather they are heavily influenced and modulated by interpersonal and contextual factors, which impact behavior and cognitions. However, the mechanisms supporting empathy are also flexible and amenable to behavioral interventions that can promote caring beyond kin and kith.

Guided by indirect evidence, recent approaches propose a tripartite crosstalk among interoceptive signaling, emotional regulation, and low-level social cognition. Here we examined the neurocognitive convergence of such domains. First, we performed three meta-analyses of functional magnetic resonance imaging studies to identify which areas are consistently coactivated by these three systems. Multi-level Kernel Density Analysis (MKDA) revealed major overlaps in the right anterior insular and frontotemporal regions (viz., the orbitofrontal and inferior frontal gyri, the amygdala, and mid temporal lobe/subcortical structures). Second, we explored such domains in patients with fronto-insulo-temporal damage. Relative to controls, the patients showed behavioral impairments of interoception, emotional processing, and social cognition, with preservation of other cognitive functions. Convergent results from both studies offer direct support for a model of insular-frontotemporal regions integrating interoception, emotion, and social cognition.

It is acknowledged that empathy plays a critical role in the physician-patient relationship and has a positive impact on health outcomes. However, as the field of empathy expands, the lack of conceptual coherence challenges advances in medicine. In fact, in some medical settings, there is little added theoretical or clinical value in applying the all-encompassing term of empathy, which is by nature multidimensional, interpersonal, and modulated by context. Functional neuroimaging studies of health professionals, designed to examine patterns of brain activation in response to empathy-eliciting situations, bring theoretical clarity to the neurocognitive mechanisms that underlie interpersonal sensitivity, emotional empathy, cognitive empathy, and caring. These components are relatively independent but often interact and are deeply interwoven in the fabric of the brain. Nevertheless, it seems clear from this review that cultivating empathic concern or compassion in today's medicine is more important than other aspects of empathy, like vicariously experiencing and introspecting about patients' emotions. Specific neurobiological mechanisms explain the benefit to patients of their physicians' perceived empathy.

Perception of biological motions plays a major adaptive role in identifying, interpreting, and predicting the actions of others. It may therefore be hypothesized that the perception of biological motions is subserved by a specific neural network. Here we used fMRI to verify this hypothesis. In a group of 10 healthy volunteers, we explored the hemodynamic responses to seven types of visual motion displays: drifting random dots, random dot cube, random dot cube with masking elements, upright point-light walker, inverted point-light walker, upright point-light walker display with masking elements, and inverted point-light walker display with masking elements. A gradient in activation was observed in the occipitotemporal junction. The responses to rigid motion were localized posteriorly to those responses elicited by nonrigid motions. Our results demonstrate that in addition to the posterior portion of superior temporal sulcus, the left intraparietal cortex is involved in the perception of nonrigid biological motions.

1. Healthy subjects actually performed and mentally simulated a leg exercise at two levels of work (15 and 19 kg loads). Heart rate, respiration rate and end‐tidal PCO2 were measured in both conditions. In addition, muscular metabolism was simultaneously measured using 31P nuclear magnetic resonance (NMR) spectroscopy. 2. During actual exercise, heart and respiration rates increased, first abruptly and then gradually in relation to the level of work. End‐tidal PCO2 was unaltered. NMR spectra showed a drop in phosphocreatine (PCr) and an increase in inorganic phosphate (Pi) concentrations. Intracellular pH fell to 6.65 at maximal effort with a 19 kg load. 3. During mental simulation, both heart and ventilatory rate increased immediately after mental exercise was begun. This increase was proportional to the amount of simulated exercise. Heart rate remained about 25% below the level observed during actual exercise. The increase in respiration rate, by contrast, was more marked than during actual exercise. Finally, end‐tidal PCO2 decreased progressively to about 18% of the resting value. 4. During mental simulation, NMR spectra were unchanged with respect to the resting values. 5. Subjects rated their sensation of fatigue using an analog rating scale, during both actual exercise and mental simulation. During mental exercise, the sensation of fatigue was greater with the 19 kg load than with the 15 kg load. 6. These results demonstrate that mental simulation of action can activate heart and respiration control mechanisms. They suggest that autonomic activation during imagined action pertains to the more general phenomenon of preparation for action.

Several studies have indicated that schizophrenic patients show impaired performance in various aspects of social cognition, including theory of mind, emotion processing, and agency judgments. Neuroimaging studies that have compared patients and healthy subjects during such mental activity indicate an abnormal hemodynamic response in the medial prefrontal cortex, the prefrontal cortex, the amygdala, the inferior parietal lobe, i.e., a set of regions known to be critically involved in social cognition. This paper addresses a number of issues raised by schizophrenia research into theory of mind, emotion perception and self-agency with regards to the neural systems that mediate social cognition. In healthy subjects, typical brain patterns are associated with theory of mind, emotion perception and self-agency; some activated clusters overlap, while others are distinct. For instance, activations in the paracingulate gyrus are almost systematically associated with theory of mind tasks, while the amygdala is mainly involved in emotion perception tasks. Additional foci are frequently found activated during those tasks: superior temporal sulcus, inferior frontal area. Moreover, the inferior parietal lobe is thought to contribute to agency judgments. In the light of the data on brain abnormalities and neurochemical dysfunctions in schizophrenia, we discuss the interaction of social cognitive dysfunction with the supposed information processing abnormalities caused by dopamine dysregulation.

This paper reviews findings from cognitive and sport psychology, as well as from neurophysiology, concerning mental simulation of movement. A neuropsychological hypothesis is advanced to explain why mental practice can improve motor skill learning. Mental practice activates certain brain structures selectively as shown by measurements of regional cerebral blood flow. It appears likely that this activation improves the subsequent control of execution of movements. It is pointed out that the study of simulation of movements may not only be of value for sport training but also have importance for the rehabilitation of patients with motor disturbances following lesions of the central nervous system.

Mutual gaze may be described as a psychological process during which two persons have the feeling of a brief link between their two minds. In the monkey, specific cell assemblies in the superior temporal cortex of the brain are responsive to gaze. This suggests that the brain may have evolved mechanisms for interpreting direct eye contact. These mechanisms could depend on the activation of specific brain regions. Positron emission tomography was used to measure activity in brain regions in healthy volunteers while they were looking at faces featuring, respectively, eye contact, averted gaze, or no gaze. As expected a region known to be involved in face processing was found to be activated in the ventral occipito-temporal region, especially in the right hemisphere. Averted gaze and mutual gaze triggered blood flow responses in similar areas which were different from those involved in face processing. These areas included the occipital part of the fusiform gyrus, the right parietal lobule, the right inferior temporal gyrus, and the middle temporal gyrus in both hemispheres. These results are consistent with the hypothesis that perception of eyes regardless of the direction of the gaze is subserved by a distributed network. However, no conclusive evidence was found for specific area(s) devoted to mutual gaze processing.

We investigated whether observation of actions reproduced in three-dimensional virtual reality would engage perceptual and visuomotor brain processes different from those induced by the observation of real hand actions. Participants were asked to passively observe grasping actions of geometrical objects made by a real hand or by hand reconstructions of different quality in 3D virtual reality as well as on a 2D TV screen. We found that only real actions in natural environment activated a visuospatial network including the right posterior parietal cortex. Observation of virtual-reality hand actions engaged prevalent visual perceptual processes within lateral and mesial occipital regions. Thus, only perception of actions in reality maps onto existing action representations, whereas virtual-reality conditions do not access the full motor knowledge available to the central nervous system.

Contingencies between objects and people can be mechanical or intentional–social in nature. In this fMRI study we used simplified stimuli to investigate brain regions involved in the detection of mechanical and intentional contingencies. Using a factorial design we manipulated the ‘animacy’ and ‘contingency’ of stimulus movement, and the subject’s attention to the contingencies. The detection of mechanical contingency between shapes whose movement was inanimate engaged the middle temporal gyrus and right intraparietal sulcus. The detection of intentional contingency between shapes whose movement was animate activated superior parietal networks bilaterally. These activations were unaffected by attention to contingency. Additional regions, the right middle frontal gyrus and left superior temporal sulcus, became activated by the animate–contingent stimuli when subjects specifically attended to the contingent nature of the stimuli. Our results help to clarify neural networks previously associated with ‘theory of mind’ and agency detection. In particular, the results suggest that low-level perception of agency in terms of objects reacting to other objects at a distance is processed by parietal networks. In contrast, the activation of brain regions traditionally associated with theory of mind tasks appears to require attention to be directed towards agency and contingency.

There is converging evidence from developmental and cognitive psychology, as well as from neuroscience, to suggest that the self is both special and social, and that self–other interaction is the driving force behind self-development. We review experimental findings which demonstrate that human infants are motivated for social interactions and suggest that the development of an awareness of other minds is rooted in the implicit notion that others are like the self. We then marshal evidence from functional neuroimaging explorations of the neurophysiological substrate of shared representations between the self and others, using various ecological paradigms such as mentally representing one’s own actions versus others’ actions, watching the actions executed by others, imitating the others’ actions versus being imitated by others. We suggest that within this shared neural network the inferior parietal cortex and the prefrontal cortex in the right hemisphere play a special role in the essential ability to distinguish the self from others, and in the way the self represents the other. Interestingly, the right hemisphere develops its functions earlier than the left.

Perception of apparent motion operates somewhat differently for objects and human figures. Depending on the interstimulus interval, the latter d may give rise to either perception of a direct path (i.e. biologically impossible) or indirect path (i.e. biologically possible). Here, PET was used to investigate whether a change in brain activity accompanies this perceptual shift. We found neural encoding of apparent motion to be a function of the intrinsic properties of the stimulus presented (object vs human) as well as the kind of human movement path perceived (biomechanically possible vs impossible). Motor and parietal cortex were only involved for possible motion which suggests that these regions are selectively activated to process actions which conform to the capabilities of the observer.

Accumulating evidence demonstrates that similar neural circuits are activated during the first-hand experience of pain and the observation of pain in others. However, most functional MRI studies did not detect signal change in the primary somatosensory cortex during pain empathy. To test if the perception of pain in others involves the primary somatosensory cortex, neuromagnetic oscillatory activity was recorded from the primary somatosensory cortex in 16 participants while they observed static pictures depicting body parts in painful and non-painful situations. The left median nerve was stimulated at the wrist, and the poststimulus rebounds of the approximately 10-Hz somatosensory cortical oscillations were quantified. Compared to the baseline condition, the level of the approximately 10-Hz oscillations was suppressed during both of the observational situations, indicating the activation of the primary somatosensory cortex. Importantly, watching painful compared to non-painful situations suppressed somatosensory oscillations to a significant stronger degree. In addition, the suppression caused by perceiving others in the painful relative to the non-painful situations correlated with the perspective taking subscale of the interpersonal reaction index. These results, consistent with the mirror-neuron system, demonstrate that the perception of pain in others modulates neural activity in primary somatosensory cortex and supports the idea that the perception of pain in others elicits subtle somatosensory activity that may be difficult to detect by fMRI techniques.

The question of whether mental simulation of an action has an effect on the spinal reflex circuits was examined in normal humans. Subjects were instructed either to exert or to mentally simulate a strong or a weak pressure on a pedal with the left or the right foot. Changes in the H- and T-reflexes activated by electrical and mechanical stimuli were measured on both legs during motor performance as well as during mental simulation of the same task. Asynchronous EMG activity of the soleus muscles was simultaneously recorded. Reflex excitability increased during performance of the pressure. It was larger when the H-reflex was triggered in the muscle involved in the task as compared to the contralateral side. Because actual performance modified the tension of the tendon and the location of the stimulus, ipsilateral changes of T-reflex amplitude could not be evaluated. Mental simulation of foot pressure in this condition resulted in a large increase of spinal reflex excitability, which was only slightly weaker than the reflex facilitation associated with the actual performance. Changes in T-reflex amplitude, but not in H-reflex amplitude, depended upon the lateralization and force of the simulated pressure, being larger in the leg involved in the simulation than in the contralateral leg, and larger for a strong than for a weak simulated movement. EMG activity was found to be weakly increased during mental imagery. This increase was significantly, although slightly, modulated by the lateralization and intensity of the imagined movement. However, no correlation was found across subjects between reflex amplitude and the amplitude of EMG activity.

The purpose of this study was to identify the functional anatomy of the mechanisms involved in visually guided prehension and in object recognition in humans. The cerebral blood flow of seven subjects was investigated by positron emission tomography. Three conditions were performed using the same set of stimuli. In the 'grasping' condition, subjects were instructed to accurately grasp the objects. In the 'matching' condition, subjects were requested to compare the shape of the presented object with that of the previous one. In the 'pointing' condition (control), subjects pointed towards the objects. The comparison between grasping and pointing showed a regional cerebral blood flow (rCBF) increase in the anterior part of the inferior parietal cortex and part of the posterior parietal cortex. The comparison between grasping and matching showed an rCBF increase in the cerebellum, the left frontal cortex around the central sulcus, the mesial frontal cortex and the left inferior parietal cortex. Finally, the comparison between matching and pointing showed an rCBF increase in the right temporal cortex and the right posterior parietal cortex. Thus object-oriented action and object recognition activate a common posterior parietal area, suggesting that some kind of within-object spatial analysis was processed by this area whatever the goal of the task.

Diffusion tensor imaging (DTI) studies in adolescents with autism spectrum disorders (ASD) indicate aberrant neurodevelopment of frontal white matter (WM), potentially underlying abnormal social cognition and communication in ASD. Here, we further use tract-based spatial statistics (TBSS) to examine the developmental change of WM skeleton (i.e., the most compact whole-brain WM) during adolescence in ASD. This whole-brain DTI used TBSS measures fractional anisotropy (FA) and longitudinal and radial diffusivities in fifty adolescents, 25 ASD and 25 controls. Results show that adolescents with ASD versus controls had significantly reduced FA in the right posterior limb of internal capsule (increased radial diffusivity distally and reduced longitudinal diffusivity centrally). Adolescents with ASD versus controls (covarying for age and IQ) had significantly greater FA in the frontal lobe (reduced radial diffusivity), right cingulate gyrus (reduced radial diffusivity), bilateral insula (reduced radial diffusivity and increased longitudinal diffusivity), right superior temporal gyrus (reduced radial diffusivity), and bilateral middle cerebellar peduncle (reduced radial diffusivity). Notably, a significant interaction with age by group was found in the right paracentral lobule and bilateral superior frontal gyrus as indicated by an age-related FA gain in the controls whilst an age-related FA loss in the ASD. To our knowledge, this is the first study to use TBSS to examine WM in individuals with ASD. Our findings indicate that the frontal lobe exhibits abnormal WM microstructure as well as an aberrant neurodevelopment during adolescence in ASD, which support the frontal disconnectivity theory of autism.

When we attend to other people in pain, the neural circuits underpinning the processing of first-hand experience of pain are activated in the observer. This basic somatic sensorimotor resonance plays a critical role in the primitive building block of empathy and moral reasoning that relies on the sharing of others’ distress. However, the full-blown capacity of human empathy is more sophisticated than the mere simulation of the target's affective state. Indeed, empathy is about both sharing and understanding the emotional state of others in relation to oneself. In this functional magnetic resonance imaging (fMRI) study, 17 typically developing children (range 7–12 yr) were scanned while presented with short animated visual stimuli depicting painful and non-painful situations. These situations involved either a person whose pain was accidentally caused or a person whose pain was intentionally inflicted by another individual. After scanning, children rated how painful these situations appeared. Consistent with previous fMRI studies of pain empathy with adults, the perception of other people in pain in children was associated with increased hemodynamic activity in the neural circuits involved in the processing of first-hand experience of pain, including the insula, somatosensory cortex, anterior midcingulate cortex, periaqueductal gray, and supplementary motor area. Interestingly, when watching another person inflicting pain onto another, regions that are consistently engaged in representing social interaction and moral behavior (the temporo-parietal junction, the paracingulate, orbital medial frontal cortices, amygdala) were additionally recruited, and increased their connectivity with the fronto-parietal attention network. These results are important to set the standard for future studies with children who exhibit social cognitive disorders (e.g., antisocial personality disorder, conduct disorder) and are often deficient in experiencing empathy or guilt.

Being in a close relationship is essential to human existence. Such closeness can be described as including other in the self and be underpinned on social attachment system, which evolved from a redirection of nociceptive mechanisms. To what extent does imagining a loved-one differs from imagining an unfamiliar individual being in painful situations? In this functional MRI study, participants were exposed to animated stimuli depicting hands or feet in painful and non-painful situations, and instructed to imagine these scenarios from three different perspectives: self, loved-one and stranger after being primed with their respective photographs. In line with previous studies, the three perspectives were associated with activation of the neural network involved in pain processing. Specifically, adopting the perspective of a loved-one increased activity in the anterior cingulate cortex and insula, whereas imagining a stranger induced a signal increase in the right temporo-parietal junction (TPJ) and superior frontal gyrus. The closer the participants' relationships were with their partner, the greater the deactivation in the right TPJ. A negative effective connectivity between the right TPJ and the insula, and a positive one with the superior frontal gyrus were found when participants imagined the perspective of a stranger. These results demonstrate that intimacy affects the bottom-up information processing involved in empathy, as indicated by greater overlap between neural representations of the self and the other.

We examined seven right-handed, asymmetrical (right side affected) Parkinson's disease patients and seven age-matched controls in a manual finger sequencing test using left and right hands in vision, no vision, and motor imagery conditions. All patients displayed motor asymmetry, favoring the left hand. They also displayed motor imagery asymmetry, mentally simulating movement more slowly with their right affected hand than with their left hand. Additionally, impairment in mental hand rotation correlated significantly with the imagery asymmetry. These data support two related hypotheses: (a) Motor sequence imagery and execution share common neural structures. (b) The frontostriatal system is among these shared structures.

PET was used to explore the neural network involved in the perception of meaningless action. In two conditions, subjects observed learned and unknown meaningless actions without any purpose. In two other conditions, subjects observed the same type of stimuli for later imitation. The control condition, which consisted of the presention of stationary hands, served as a baseline. Unsurprisingly, a common network that forms part of the dorsal pathway was engaged in all conditions when compared with stationary hands, and this was interpreted as being devoted to the analysis of hand movements. One of the most striking results of the present study was that some brain areas were strongly modulated by the learning level, independent of the subject's intention. Two different effects were observed: a reduced activity in posterior regions within the common network, which correlated with specific increases in the frontopolar area 10 and in the angular gyrus during the perception of learned meaningless actions compared with the perception of unknown actions. Finally, the major effect of the subject's intention to imitate was a strong increase in the dorsal pathway extending to the lateral premotor cortex and to the dorsolateral prefrontal cortex, which reflects the information processing needed for prospective action. Overall, our results provide evidence for both an effect of the visuomotor learning level and of the subject's intention on the neural network involved during the perception of human meaningless actions.

The eye region and gaze behaviour are known to play a major role in conveying information about direction of attention and emotional dispositions. Positron emission tomography scanning was used to explore the cerebral structures involved while subjects were asked to attribute hostile or friendly intentions to video-taped actors who directed attention towards or away from the subjects. As expected, a number of brain regions known to be involved in emotion processing was found activated when subjects had to attribute an emotion regardless of gaze direction. In addition, results indicate that gaze direction has an impact on the brain regions recruited to interpret emotions. The anterior region of the superior temporal gyrus (STG) was selectively activated during analysis of emotions through eye contact. This result provides neurophysiological evidence for privileged processing when an individual becomes personally involved as the object of another's emotions.

Studying of the impact of social context on the perception of pain in others is important for understanding the role of intentionality in interpersonal sensitivity, empathy, and implicit moral reasoning. Here we used an event-related fMRI with pain and social context (i.e., the number of individuals in the stimuli) as the two factors to investigate how different social contexts and resulting perceived agency modulate the neural response to the perception of pain in others. Twenty-six healthy participants were scanned while presented with short dynamic visual stimuli depicting painful situations accidentally caused by or intentionally caused by another individual. The main effect of perception of pain was associated with signal increase in the aMCC, insula, somatosensory cortex, SMA and PAG. Importantly, perceiving the presence of another individual led to specific hemodynamic increase in regions involved in representing social interaction and emotion regulation including the temporoparietal junction, medial prefrontal cortex, inferior frontal gyrus, and orbitofrontal cortex. Furthermore, the functional connectivity pattern between the left amygdala and other brain areas was modulated by the perceived agency. Our study demonstrates that the social context in which pain occurs modulate the brain response to other's pain. This modulation may reflect successful adaptation to potential danger present in a social interaction. Our results contribute to a better understanding of the neural mechanisms underpinning implicit moral reasoning that concern actions that can harm other people.

Agency is the sense that I am the one generating an action. In this neuroimaging experiment, subjects controlled a circle with a mouse while requested either to lead another circle (i.e., being the agent) or to follow it (i.e., being acted upon). Clusters within the right intraparietal sulcus were associated with following for the most rostral and leading for the most caudal ones. Bilateral activity in the inferior parietal lobule in conditions involving confusion about the origin of the action confirmed its role in agency. A lateralization effect was also found in these conditions, the response being stronger in the left inferior parietal lobule when subjects were not the agent of the performed action, and in the right when they were.

Accumulative empirical evidence has been reviewed in support of the notion that the production and perception of action as well as the interpretation of others’ actions are functionally connected, and indeed, rely on common distributed neural systems in the premotor and parietal cortices. We suggest that these neural systems sustain shared representations between self and other that are crucial in social interactions. The inferior parietal cortex plays a special role in the sense of agency, which is a fundamental aspect to navigate within this neural network. The role of other brain areas that implement and regulate these shared representations remains to be specified.

Theory of mind (ToM), the specific ability to attribute thoughts and feelings to oneself and others is generally impaired in schizophrenia. Previous studies demonstrated a deficit of the attribution of intentions to others among patients having formal thought disorder. During nonverbal tasks, such a function requires both the visual perception of human figures and the understanding of their intentions. These processes are considered to involve the superior temporal sulcus and the medial prefrontal cortex, respectively. Are the functional patterns of activation associated with those processes abnormal in schizophrenia? Seven schizophrenic patients on medication performed a nonverbal attribution of intentions task as well as two matched physical logic tasks, with and without human figures, while H2O15 PET-scanning was performed. Data from the patients were compared to those of eight healthy controls matched for verbal IQ and sex. The experimental design allowed dissociating the effect of the perception of human figures from that of the attribution of intentions. During attribution of intentions, significant activations in the right prefrontal cortex were detected in the control subjects. Those activations were not found in the schizophrenic group. However, in both groups, the perception of human figure elicited bilateral activation of the occipitotemporal regions and of the posterior part of the superior temporal sulcus. Schizophrenic patients performing a nonverbal attribution of intentions task have an abnormal cerebral activity.

Recent neuropsychological investigations of apraxia have led to new hypotheses about the representational defects associated with imitation impairments in neurological patients. This fMRI experiment investigated the relation between imitation and the body schema in healthy subjects. Experimental conditions were derived from a factorial plan, and participants were asked to watch a human model performing bodily gestures and then to execute either an identical or a different action, with the same or different limbs. Brain areas activated when subjects imitated the model were traced to the representation of the action (main effect of performing an identical action regardless of limb), to the body schema (using the same limb regardless of action), or to both. The first set of analyses yielded a network associated with visual perception, indicating that action representation is primarily visuospatial not motor, while the second analysis highlighted regions involved in body schema including the inferior parietal cortex and the insula. It is suggested that imitation of simple body gestures requires both a visuospatial description of the observed model, sustained by visual perception areas in the right occipitotemporal and superior parietal cortices and a visuospatial description of one’s own body, supported by the left inferior parietal lobule. These results favor a model of praxis proposing that imitation deficits in left inferior parietal lobe patients with apraxia reflect primarily an impairment of the body schema, while deficits of praxis in right parietal patients are limited to gestures demanding in terms of visuospatial analysis.

Females frequently perform better in empathy, interpersonal sensitivity, and emotional recognition than do males. The mirror-neuron system has been proposed to play an important role in social cognition. It remains to be clarified, however, whether the neuroanatomy underlying the human mirror neuron system exhibits sex differences. With the use of voxel-based morphometry analysis, a whole-brain unbiased technique to characterize regional cerebral volume differences in structural magnetic resonance images, concurrent with the dispositional empathy measures, we demonstrate that young adult females (n=25) had significantly larger gray matter volume in the pars opercularis and inferior parietal lobule than matched males (n=25) participants. Moreover, higher self-report scores in the emotional empathic disposition was tightly coupled with larger gray matter volume of the pars opercularis across all female and male participants (P=0.002). These results indicate that the existence of neuroanatomical sex differences in the human mirror-neuron system. They also suggest that the network of the human mirror-neuron system is strongly linked to empathy competence.

The 'broken mirror' theory of autism, which proposes that a dysfunction of the human mirror neuron system (MNS) is responsible for the core social and cognitive deficits in individuals with autism spectrum disorders (ASD), has received considerable attention despite weak empirical evidence.In this electroencephalographic study, we examined mu suppression, as an indicator of sensorimotor resonance, concurrent with oculomotor performance while individuals (n = 20) with ASD and control participants (n = 20) either executed hand actions or observed hand actions or a moving dot. No difference in visual attention between groups was found as indicated by fixation duration and normalized fixation number on the presented stimuli.The mu suppression over the sensorimotor cortex was significantly affected by experimental conditions, but not by group membership, nor by the interaction between groups and conditions. Individuals with ASD, similar to the controls, exhibited stronger mu suppression when watching hand actions relative to a moving dot. Notably, participants with ASD failed to imitate the observed actions while their mu suppression indicating the MNS activity was intact. In addition, the mu suppression during the observation of hand actions was positively associated with the communication competence of individuals with ASD.Our study clearly challenges the broken mirror theory of autism. The functioning of the mirror neuron system might be preserved in individuals with ASD to a certain degree. Less mu suppression to action observation coupled with more communicational severity can reflect the symptom heterogeneity of ASD. Additional research needs to be done, and more caution should be used when reaching out to the media.

Results from recent functional neuroimaging studies suggest that facial expressions of pain trigger empathic mimicry responses in the observer, in the sense of an activation in the pain matrix. However, pain itself also signals a potential threat in the environment and urges individuals to escape or avoid its source. This evolutionarily primitive aspect of pain processing, i.e., avoidance from the threat value of pain, seems to conflict with the emergence of empathic concern, i.e., a motivation to approach toward the other. The present study explored whether the affective values of targets influence the detection of pain at the unconscious level. We found that the detection of pain was facilitated by unconscious negative affective processing rather than by positive affective processing. This suggests that detection of pain is primarily influenced by its inherent threat value, and that empathy and empathic concern may not rely on a simple reflexive resonance as generally thought. The results of this study provide a deeper understanding of how fundamental the unconscious detection of pain is to the processes involved in the experience of empathy and sympathy.

Patients with first-rank symptoms (FRS) of schizophrenia do not experience all of their actions and personal states as their own. FRS may be associated with an impaired ability to correctly attribute an action to its origin. In the present study, we examined regional cerebral blood flow (rCBF) with positron emission tomography during an action-attribution task in a group of patients with FRS. We used a device previously used with healthy subjects that allows the experimenter to modulate the subject's degree of movement control (and thus action attribution) of a virtual hand presented on a screen. In healthy subjects, the activity of the right angular gyrus and the insula cortex appeared to be modulated by the subject's degree of movement control of the virtual hand. In the present study, the schizophrenic patients did not show this pattern. We found an aberrant relationship between the subject's degree of control of the movements and rCBF in the right angular gyrus and no modulation in the insular cortex. The implications of these results for understanding pathological conditions such as schizophrenia are discussed.

Lack of empathy is a hallmark of social impairments in individuals with autism spectrum disorder (ASD). However, the concept empathy encompasses several socio-emotional and behavioral components underpinned by interacting brain circuits. This study examined empathic arousal and social understanding in individuals with ASD and matched controls by combining pressure pain thresholds (PPT) with functional magnetic resonance imaging (study 1) and electroencephalography/event-related potentials and eye-tracking responses (study 2) to empathy-eliciting stimuli depicting physical bodily injuries. Results indicate that participants with ASD had lower PPT than controls. When viewing body parts being accidentally injured, increased hemodynamic responses in the somatosensory cortex (SI/SII) but decreased responses in the anterior mid-cingulate and anterior insula as well as heightened N2 but preserved late-positive potentials (LPP) were detected in ASD participants. When viewing a person intentionally hurting another, decreased hemodynamic responses in the medial prefrontal cortex and reduced LPP were observed in the ASD group. PPT was a mediator for the SI/SII response in predicting subjective unpleasantness ratings to others' pain. Both ASD and control groups had comparable mu suppression, indicative of typical sensorimotor resonance. The findings demonstrate that, in addition to reduced pain thresholds, individuals with ASD exhibit heightened empathic arousal but impaired social understanding when perceiving others' distress.

Neuroscience research indicates that moral reasoning is underpinned by distinct neural networks including the posterior superior temporal sulcus (pSTS), amygdala, and ventromedial prefrontal cortex, which support communication between computational systems underlying affective states, cognitions, and motivational processes. To characterize real-time neural processing underpinning moral computations, high-density event-related potentials were measured in participants while they viewed short, morally laden visual scenarios depicting intentional and accidental harmful actions. Current source density maxima in the right pSTS as fast as 62 ms poststimulus first distinguished intentional vs. accidental actions. Responses in the amygdala/temporal pole (122 ms) and ventromedial prefrontal cortex (182 ms) were then evoked by the perception of harmful actions, indicative of fast information processing associated with early stages of moral cognition. Our data strongly support the notion that intentionality is the first input to moral computations. They also demonstrate that emotion acts as a gain antecedent to moral judgment by alerting the individual to the moral salience of a situation and provide evidence for the pervasive role of affect in moral sensitivity and reasoning.