Understanding others' actions and goals by mirror and mentalizing systems: A meta-analysis

This meta-analysis explores the role of the mirror and mentalizing systems in the understanding of other people's action goals. Based on over 200 fMRI studies, this analysis demonstrates that the mirror system - consisting of the anterior intraparietal sulcus and the premotor cortex - is engaged when one perceives articulated motions of body parts irrespective of their sensory (visual or auditory) or verbal format as well as when the perceiver executes them. This confirms the matching role of the mirror system in understanding biological action. Observation of whole-body motions and gaze engage the posterior superior temporal sulcus and most likely reflects an orientation response in line with the action or attention of the observed actor. In contrast, the mentalizing system - consisting of the temporo-parietal junction, the medial prefrontal cortex and the precuneus - is activated when behavior that enables inferences to be made about goals, beliefs or moral issues is presented in abstract terms (e.g., verbal stories or geometric shapes) and there is no perceivable biological motion of body parts. A striking overlap of brain activity at the temporo-parietal junction between social inferences and other, non-social observations (e.g., Posner's cuing task) suggests that this area computes the orientation or direction of the behavior in order to predict its likely end-state (or goal). No conclusions are drawn about the specific functionality of the precuneus and the medial prefrontal cortex. Because the mirror and mentalizing systems are rarely concurrently active, it appears that neither system subserves the other. Rather, they are complementary. There seems, however, to be a transition from the mirror to the mentalizing system even when body-part motions are observed by perceivers who are consciously deliberating about the goals of others and their behavioral executions, such as when perceived body motions are contextually inconsistent, implausible or pretended.

[1]  E. Naito,et al.  Enhanced neural activity in response to dynamic facial expressions of emotion: an fMRI study. , 2004, Brain research. Cognitive brain research.

[2]  Roel M. Willems,et al.  Complementary Systems for Understanding Action Intentions , 2008, Current Biology.

[3]  Susan T. Fiske,et al.  Dehumanizing the Lowest of the Low Responses to Extreme Out-Groups , 2006 .

[4]  Jordan Henry Grafman,et al.  Social contracts and precautions activate different neurological systems: An fMRI investigation of deontic reasoning , 2005, NeuroImage.

[5]  R. Passingham,et al.  Seeing or Doing? Influence of Visual and Motor Familiarity in Action Observation , 2006, Current Biology.

[6]  S. Pollmann,et al.  Covert Reorienting and Inhibition of Return: An Event-Related fMRI Study , 2002, Journal of Cognitive Neuroscience.

[7]  Andrea C. Pierno,et al.  Transfer of interfered motor patterns to self from others , 2006, The European journal of neuroscience.

[8]  Hanneke E. M. den Ouden,et al.  Thinking about intentions , 2005, NeuroImage.

[9]  G. Fink,et al.  Being with virtual others: Neural correlates of social interaction , 2006, Neuropsychologia.

[10]  T. Paus,et al.  Brain networks involved in viewing angry hands or faces. , 2006, Cerebral cortex.

[11]  Friedemann Pulvermüller,et al.  Brain mechanisms linking language and action , 2005, Nature Reviews Neuroscience.

[12]  Katja Fiehler,et al.  What activates the human mirror neuron system during observation of artificial movements: Bottom-up visual features or top-down intentions? , 2008, Neuropsychologia.

[13]  M. Jeannerod,et al.  The motor theory of social cognition: a critique , 2005, Trends in Cognitive Sciences.

[14]  Aina Puce,et al.  Viewing the motion of human body parts activates different regions of premotor, temporal, and parietal cortex , 2004, NeuroImage.

[15]  Parashkev Nachev,et al.  Space and the parietal cortex , 2007, Trends in Cognitive Sciences.

[16]  J. Mazziotta,et al.  Grasping the Intentions of Others with One's Own Mirror Neuron System , 2005, PLoS biology.

[17]  Armin Thron,et al.  An fMRI approach to particularize the frontoparietal network for visuomotor action monitoring: Detection of incongruence between test subjects’ actions and resulting perceptions , 2007, NeuroImage.

[18]  J. Grèzes,et al.  What is “mirror” in the premotor cortex? A review , 2008, Neurophysiologie Clinique/Clinical Neurophysiology.

[19]  Rebecca Saxe,et al.  The neural basis of belief encoding and integration in moral judgment , 2008, NeuroImage.

[20]  G. McCarthy,et al.  When Strangers Pass , 2004, Psychological science.

[21]  Volkmar Glauche,et al.  Functional properties and interaction of the anterior and posterior intraparietal areas in humans , 2003, The European journal of neuroscience.

[22]  Istvan Molnar-Szakacs,et al.  Observing complex action sequences: The role of the fronto-parietal mirror neuron system , 2006, NeuroImage.

[23]  Ran R. Hassin,et al.  The New Unconscious , 2006 .

[24]  J. Mazziotta,et al.  Cortical mechanisms of human imitation. , 1999, Science.

[25]  V. Bruce,et al.  Do the eyes have it? Cues to the direction of social attention , 2000, Trends in Cognitive Sciences.

[26]  Jody C. Culham,et al.  Does tool-related fMRI activity within the intraparietal sulcus reflect the plan to grasp? , 2007, NeuroImage.

[27]  R. Saxe Against simulation: the argument from error , 2005, Trends in Cognitive Sciences.

[28]  Justin L. Gardner,et al.  Executed and Observed Movements Have Different Distributed Representations in Human aIPS , 2008, The Journal of Neuroscience.

[29]  Kevin A. Pelphrey,et al.  Grasping the Intentions of Others: The Perceived Intentionality of an Action Influences Activity in the Superior Temporal Sulcus during Social Perception , 2004, Journal of Cognitive Neuroscience.

[30]  C. Frith,et al.  Experiencing Oneself vs Another Person as Being the Cause of an Action: The Neural Correlates of the Experience of Agency , 2002, NeuroImage.

[31]  Scott T. Grafton,et al.  Action outcomes are represented in human inferior frontoparietal cortex. , 2008, Cerebral cortex.

[32]  I. Johnsrude,et al.  Somatotopic Representation of Action Words in Human Motor and Premotor Cortex , 2004, Neuron.

[33]  D. Perrett,et al.  A region of right posterior superior temporal sulcus responds to observed intentional actions , 2004, Neuropsychologia.

[34]  G. Glover,et al.  Children's and adults’ neural bases of verbal and nonverbal ‘theory of mind’ , 2007, Neuropsychologia.

[35]  Diana C. Robertson,et al.  The neural processing of moral sensitivity to issues of justice and care , 2007, Neuropsychologia.

[36]  Maurizio Corbetta,et al.  Visuospatial reorienting signals in the human temporo‐parietal junction are independent of response selection , 2006, The European journal of neuroscience.

[37]  D. Dennett The Intentional Stance. , 1987 .

[38]  C. Frith,et al.  Adolescent development of the neural circuitry for thinking about intentions , 2007, Social cognitive and affective neuroscience.

[39]  R. E Passingham,et al.  Inferring false beliefs from the actions of oneself and others: an fMRI study , 2004, NeuroImage.

[40]  Luca Turella,et al.  Observing social interactions: The effect of gaze , 2008, Social neuroscience.

[41]  David J. Freedman,et al.  Experience-dependent representation of visual categories in parietal cortex , 2006, Nature.

[42]  Richard Henson,et al.  Forward inference using functional neuroimaging: dissociations versus associations , 2006, Trends in Cognitive Sciences.

[43]  E. Macaluso,et al.  Dissociation of stimulus relevance and saliency factors during shifts of visuospatial attention. , 2007, Cerebral cortex.

[44]  C. Frith,et al.  Meeting of minds: the medial frontal cortex and social cognition , 2006, Nature Reviews Neuroscience.

[45]  Jin Fan,et al.  Development of attentional networks: An fMRI study with children and adults , 2005, NeuroImage.

[46]  R. E Passingham,et al.  Activations related to “mirror” and “canonical” neurones in the human brain: an fMRI study , 2003, NeuroImage.

[47]  H. Aarts,et al.  The silence of the library: environment, situational norm, and social behavior. , 2003, Journal of personality and social psychology.

[48]  Sakiko Yoshikawa,et al.  Time course of superior temporal sulcus activity in response to eye gaze: a combined fMRI and MEG study. , 2008, Social cognitive and affective neuroscience.

[49]  Paula Tavares,et al.  Paying attention to social meaning: an FMRI study. , 2008, Cerebral cortex.

[50]  J. Downar,et al.  A multimodal cortical network for the detection of changes in the sensory environment , 2000, Nature Neuroscience.

[51]  G. Rizzolatti,et al.  Understanding motor events: a neurophysiological study , 2004, Experimental Brain Research.

[52]  J. Decety,et al.  The Role of the Right Temporoparietal Junction in Social Interaction: How Low-Level Computational Processes Contribute to Meta-Cognition , 2007, The Neuroscientist : a review journal bringing neurobiology, neurology and psychiatry.

[53]  Marcel Brass,et al.  The Inhibition of Imitative Response Tendencies , 2001, NeuroImage.

[54]  Mirella Dapretto,et al.  Developmental changes in the neural basis of interpreting communicative intent. , 2006, Social cognitive and affective neuroscience.

[55]  D. Collins,et al.  Automatic 3D Intersubject Registration of MR Volumetric Data in Standardized Talairach Space , 1994, Journal of computer assisted tomography.

[56]  H. Walter,et al.  The intentional network: How the brain reads varieties of intentions , 2007, Neuropsychologia.

[57]  J. Mazziotta,et al.  Modulation of cortical activity during different imitative behaviors. , 2003, Journal of neurophysiology.

[58]  Luca Turella,et al.  Mirror neurons in humans: Consisting or confounding evidence? , 2009, Brain and Language.

[59]  Scott T. Grafton,et al.  Differential role of the orbital frontal lobe in emotional versus cognitive perspective-taking , 2006, Neuropsychologia.

[60]  David J. Turk,et al.  The angular gyrus computes action awareness representations. , 2008, Cerebral cortex.

[61]  M. Corbetta,et al.  An Event-Related Functional Magnetic Resonance Imaging Study of Voluntary and Stimulus-Driven Orienting of Attention , 2005, The Journal of Neuroscience.

[62]  Ravi S. Menon,et al.  Visually guided grasping produces fMRI activation in dorsal but not ventral stream brain areas , 2003, Experimental Brain Research.

[63]  G. R. Fink,et al.  The modulatory effects of nicotine on parietal cortex activity in a cued target detection task depend on cue reliability , 2006, Neuroscience.

[64]  Ewald Moser,et al.  The selection of intended actions and the observation of others' actions: A time-resolved fMRI study , 2006, NeuroImage.

[65]  E. Vatikiotis-Bateson,et al.  Perceiving Biological Motion: Dissociating Visible Speech from Walking , 2003, Journal of Cognitive Neuroscience.

[66]  R. Saxe,et al.  The neural basis of the interaction between theory of mind and moral judgment , 2007, Proceedings of the National Academy of Sciences.

[67]  H. Freund,et al.  The role of the inferior parietal cortex in linking the tactile perception and manual construction of object shapes. , 2001, Cerebral cortex.

[68]  F. Overwalle Social cognition and the brain: a meta-analysis. , 2009 .

[69]  Jana Schaich Borg,et al.  Consequences, Action, and Intention as Factors in Moral Judgments: An fMRI Investigation , 2006, Journal of Cognitive Neuroscience.

[70]  Chris I. Baker,et al.  Integration of Visual and Auditory Information by Superior Temporal Sulcus Neurons Responsive to the Sight of Actions , 2005, Journal of Cognitive Neuroscience.

[71]  K. Pelphrey,et al.  School-aged children exhibit domain-specific responses to biological motion , 2006, Social neuroscience.

[72]  C. Frith,et al.  Functional imaging of ‘theory of mind’ , 2003, Trends in Cognitive Sciences.

[73]  C. Keysers,et al.  Integrating simulation and theory of mind: from self to social cognition , 2007, Trends in Cognitive Sciences.

[74]  Tao Wu,et al.  The role of the dorsal stream for gesture production , 2006, NeuroImage.

[75]  Ruud G. J. Meulenbroek,et al.  Anatomical substrates of cooperative joint-action in a continuous motor task: Virtual lifting and balancing , 2008, NeuroImage.

[76]  Hiroshi Matsuda,et al.  Impaired self-awareness and theory of mind: An fMRI study of mentalizing in alexithymia , 2006, NeuroImage.

[77]  J. Haxby,et al.  Distinct representations of eye gaze and identity in the distributed human neural system for face perception , 2000, Nature Neuroscience.

[78]  Mitsuo Kawato,et al.  Activation of the Human Superior Temporal Gyrus during Observation of Goal Attribution by Intentional Objects , 2004, Journal of Cognitive Neuroscience.

[79]  Andrew N. Meltzoff,et al.  Neural circuits involved in imitation and perspective-taking , 2006, NeuroImage.

[80]  S. Baron-Cohen,et al.  Gaze Perception Triggers Reflexive Visuospatial Orienting , 1999 .

[81]  Gereon R Fink,et al.  Cerebral correlates of alerting, orienting and reorienting of visuospatial attention: an event-related fMRI study , 2004, NeuroImage.

[82]  J. Mazziotta,et al.  Lateralization of the Human Mirror Neuron System , 2006, The Journal of Neuroscience.

[83]  J. Haxby,et al.  fMRI Responses to Video and Point-Light Displays of Moving Humans and Manipulable Objects , 2003, Journal of Cognitive Neuroscience.

[84]  G. Rizzolatti,et al.  Neurophysiological mechanisms underlying the understanding and imitation of action , 2001, Nature Reviews Neuroscience.

[85]  Hannes Rakoczy,et al.  Minds, persons, and space: An fMRI investigation into the relational complexity of higher-order intentionality , 2008, Consciousness and Cognition.

[86]  D. Na,et al.  Functional magnetic resonance imaging during pantomiming tool-use gestures , 2001, Experimental Brain Research.

[87]  Anjan Chatterjee,et al.  Specificity of Action Representations in the Lateral Occipitotemporal Cortex , 2006, Journal of Cognitive Neuroscience.

[88]  Josef Perner,et al.  Do visual perspective tasks need theory of mind? , 2006, NeuroImage.

[89]  E. Macaluso,et al.  Supramodal Effects of Covert Spatial Orienting Triggered by Visual or Tactile Events , 2002, Journal of Cognitive Neuroscience.

[90]  Aaron C. Koralek,et al.  Two Takes on the Social Brain: A Comparison of Theory of Mind Tasks , 2007, Journal of Cognitive Neuroscience.

[91]  R. J. Seitz,et al.  A fronto‐parietal circuit for object manipulation in man: evidence from an fMRI‐study , 1999, The European journal of neuroscience.

[92]  G. Rizzolatti,et al.  A unifying view of the basis of social cognition , 2004, Trends in Cognitive Sciences.

[93]  Pierre Fonlupt,et al.  Listening to a walking human activates the temporal biological motion area , 2005, NeuroImage.

[94]  Eleanor A. Maguire,et al.  Spontaneous mentalizing during an interactive real world task: An fMRI study , 2006, Neuropsychologia.

[95]  Ori Friedman,et al.  Core mechanisms in ‘theory of mind’ , 2004, Trends in Cognitive Sciences.

[96]  D. Perrett,et al.  Opinion TRENDS in Cognitive Sciences Vol.8 No.11 November 2004 Demystifying social cognition: a Hebbian perspective , 2022 .

[97]  M Corbetta,et al.  Multiple neural correlates of detection in the human brain. , 2001, Proceedings of the National Academy of Sciences of the United States of America.

[98]  Jeffrey M. Zacks,et al.  Human Brain Activity Time-Locked to Narrative Event Boundaries , 2007, Psychological science.

[99]  K. Paller,et al.  Brain networks for analyzing eye gaze. , 2003, Brain research. Cognitive brain research.

[100]  Kiyotaka Nemoto,et al.  The neural network for the mirror system and mentalizing in normally developed children: an fMRI study , 2004, Neuroreport.

[101]  R. Blake,et al.  Brain Areas Active during Visual Perception of Biological Motion , 2002, Neuron.

[102]  Jean Decety,et al.  Motivation modulates the activity of the human mirror-neuron system. , 2007, Cerebral cortex.

[103]  Ran R. Hassin,et al.  Automatic goal inferences. , 2005 .

[104]  R. Saxe Uniquely human social cognition , 2006, Current Opinion in Neurobiology.

[105]  M. Corbetta,et al.  Neural Systems for Visual Orienting and Their Relationships to Spatial Working Memory , 2002, Journal of Cognitive Neuroscience.

[106]  R. Marois,et al.  Visual Short-Term Memory Load Suppresses Temporo-Parietal Junction Activity and Induces Inattentional Blindness , 2005, Psychological science.

[107]  F. Van Overwalle,et al.  Electrophysiological time course and brain areas of spontaneous and intentional trait inferences. , 2007, Social cognitive and affective neuroscience.

[108]  C Dohle,et al.  Human anterior intraparietal area subserves prehension , 1998, Neurology.

[109]  Ivanei E. Bramati,et al.  The Neural Correlates of Moral Sensitivity: A Functional Magnetic Resonance Imaging Investigation of Basic and Moral Emotions , 2002, The Journal of Neuroscience.

[110]  M. Iacoboni Neural mechanisms of imitation , 2005, Current Opinion in Neurobiology.

[111]  Beate Sodian,et al.  Neural correlates of true and false belief reasoning , 2007, NeuroImage.

[112]  Rebecca Saxe,et al.  Reading minds versus following rules: Dissociating theory of mind and executive control in the brain , 2006, Social neuroscience.

[113]  J. Decety,et al.  Functional anatomy of execution, mental simulation, observation, and verb generation of actions: A meta‐analysis , 2001, Human brain mapping.

[114]  Scott T. Grafton,et al.  Actions or Hand-Object Interactions? Human Inferior Frontal Cortex and Action Observation , 2003, Neuron.

[115]  Alex Martin,et al.  Representation of Manipulable Man-Made Objects in the Dorsal Stream , 2000, NeuroImage.

[116]  Christian Keysers,et al.  Aplasics Born without Hands Mirror the Goal of Hand Actions with Their Feet , 2007, Current Biology.

[117]  Annette Baumgaertner,et al.  Polymodal conceptual processing of human biological actions in the left inferior frontal lobe , 2007, The European journal of neuroscience.

[118]  Diederik A. Stapel,et al.  Silence and Table Manners: When Environments Activate Norms , 2008, Personality & social psychology bulletin.

[119]  井関 一海,et al.  Neural mechanisms involved in mental imagery and observation of gait , 2008 .

[120]  Gereon R Fink,et al.  Differential involvement of the posterior temporal cortex in mentalizing but not perspective taking. , 2008, Social cognitive and affective neuroscience.

[121]  R. Campbell,et al.  Reading Speech from Still and Moving Faces: The Neural Substrates of Visible Speech , 2003, Journal of Cognitive Neuroscience.

[122]  Jordan Grafman,et al.  Functional Networks in Emotional Moral and Nonmoral Social Judgments , 2002, NeuroImage.

[123]  Jaap Ham,et al.  Smart and easy: Co-occurring activation of spontaneous trait inferences and spontaneous situational inferences , 2003 .

[124]  Henrik Walter,et al.  Understanding Intentions in Social Interaction: The Role of the Anterior Paracingulate Cortex , 2004, Journal of Cognitive Neuroscience.

[125]  T. Allison,et al.  Social perception from visual cues: role of the STS region , 2000, Trends in Cognitive Sciences.

[126]  J. Mazziotta,et al.  Functional segregation within pars opercularis of the inferior frontal gyrus: evidence from fMRI studies of imitation and action observation. , 2005, Cerebral cortex.

[127]  D Yves von Cramon,et al.  Premotor cortex in observing erroneous action: an fMRI study. , 2003, Brain research. Cognitive brain research.

[128]  C. Keysers,et al.  Facial expressions: What the mirror neuron system can and cannot tell us , 2007, Social neuroscience.

[129]  G. Rizzolatti,et al.  Action observation activates premotor and parietal areas in a somatotopic manner: an fMRI study , 2001, The European journal of neuroscience.

[130]  Gian Luca Romani,et al.  Neural systems underlying observation of humanly impossible movements: an FMRI study. , 2005, Cerebral cortex.

[131]  William Gaetz,et al.  Activation of area MT/V5 and the right inferior parietal cortex during the discrimination of transient direction changes in translational motion. , 2007, Cerebral cortex.

[132]  Emily S. Cross,et al.  Building a motor simulation de novo: Observation of dance by dancers , 2006, NeuroImage.

[133]  David I. Perrett,et al.  An fMRI study of joint attention experience , 2005, NeuroImage.

[134]  M. Brass,et al.  Investigating Action Understanding: Inferential Processes versus Action Simulation , 2007, Current Biology.

[135]  D Yves von Cramon,et al.  Sequences of Abstract Nonbiological Stimuli Share Ventral Premotor Cortex with Action Observation and Imagery , 2004, The Journal of Neuroscience.

[136]  Alex Martin,et al.  NEURAL FOUNDATIONS FOR UNDERSTANDING SOCIAL AND MECHANICAL CONCEPTS , 2003, Cognitive neuropsychology.

[137]  Rebecca Elliott,et al.  Neuronal correlates of theory of mind and empathy: A functional magnetic resonance imaging study in a nonverbal task , 2006, NeuroImage.

[138]  Klaas E. Stephan,et al.  Visuospatial attention: how to measure effects of infrequent, unattended events in a blocked stimulus design , 2004, NeuroImage.

[139]  R. Hari,et al.  Viewing Lip Forms Cortical Dynamics , 2002, Neuron.

[140]  R. Leiguarda,et al.  The neural substrate of gesture recognition , 2008, Neuropsychologia.

[141]  Karl J. Friston,et al.  Action selectivity in parietal and temporal cortex. , 2005, Brain research. Cognitive brain research.

[142]  Shigeru Sato,et al.  Comprehension of implicit meanings in social situations involving irony: A functional MRI study , 2007, NeuroImage.

[143]  M. Brass,et al.  The inhibition of imitative and overlearned responses: a functional double dissociation , 2005, Neuropsychologia.

[144]  C. Keysers,et al.  Towards a unifying neural theory of social cognition. , 2006, Progress in brain research.

[145]  Peter Thier,et al.  The posterior superior temporal sulcus is involved in social communication not specific for the eyes , 2008, Neuropsychologia.

[146]  Jason P. Mitchell Mentalizing and Marr: An information processing approach to the study of social cognition , 2006, Brain Research.

[147]  Alison J. Wiggett,et al.  Patterns of fMRI Activity Dissociate Overlapping Functional Brain Areas that Respond to Biological Motion , 2006, Neuron.

[148]  A. Cavanna,et al.  The precuneus: a review of its functional anatomy and behavioural correlates. , 2006, Brain : a journal of neurology.

[149]  F. Nielsen,et al.  Right temporoparietal cortex activation during visuo-proprioceptive conflict. , 2004, Cerebral Cortex.

[150]  C. Frith,et al.  Interacting minds--a biological basis. , 1999, Science.

[151]  R. Dolan,et al.  An fMRI study of intentional and unintentional (embarrassing) violations of social norms. , 2002, Brain : a journal of neurology.

[152]  Evelyn C. Ferstl,et al.  What Does the Frontomedian Cortex Contribute to Language Processing: Coherence or Theory of Mind? , 2002, NeuroImage.

[153]  Gregory McCarthy,et al.  Regional Brain Activation Evoked When Approaching a Virtual Human on a Virtual Walk , 2005, Journal of Cognitive Neuroscience.

[154]  J. Mazziotta,et al.  Neural mechanisms of empathy in humans: A relay from neural systems for imitation to limbic areas , 2003, Proceedings of the National Academy of Sciences of the United States of America.

[155]  A. Villringer,et al.  An fMRI study of simple ethical decision-making , 2003, Neuroreport.

[156]  Frank Van Overwalle,et al.  Please Scroll down for Article Social Neuroscience Erp Time Course and Brain Areas of Spontaneous and Intentional Goal Inferences Erp Time Course and Brain Areas of Spontaneous and Intentional Goal Inferences , 2022 .

[157]  Kent A. Kiehl,et al.  Neural correlates of the processing of another’s mistakes: A possible underpinning for social and observational learning , 2008, NeuroImage.

[158]  Lisa A. Parr,et al.  The evolution of social cognition , 2011 .

[159]  R. J. Seitz,et al.  A parieto-premotor network for object manipulation: evidence from neuroimaging , 1999, Experimental Brain Research.

[160]  C. Frith,et al.  Reading the mind in cartoons and stories: an fMRI study of ‘theory of mind’ in verbal and nonverbal tasks , 2000, Neuropsychologia.

[161]  M. Corbetta,et al.  Control of goal-directed and stimulus-driven attention in the brain , 2002, Nature Reviews Neuroscience.

[163]  Michael B. Miller,et al.  Neural Correlates of Detecting Pretense: Automatic Engagement of the Intentional Stance under Covert Conditions , 2004, Journal of Cognitive Neuroscience.

[164]  Christian Keysers,et al.  The anthropomorphic brain: The mirror neuron system responds to human and robotic actions , 2007, NeuroImage.

[165]  R. Miall,et al.  A system in the human brain for predicting the actions of others , 2004, Nature Neuroscience.

[166]  H. Forssberg,et al.  Differential fronto-parietal activation depending on force used in a precision grip task: an fMRI study. , 2001, Journal of neurophysiology.

[167]  D. Perrett,et al.  Cortical and subcortical mechanisms at the core of imitation , 2007, Social neuroscience.

[168]  Livia Colle,et al.  The neural basis for understanding non-intended actions , 2007, NeuroImage.

[169]  S. Carey,et al.  Understanding other minds: linking developmental psychology and functional neuroimaging. , 2004, Annual review of psychology.

[170]  Scott T. Grafton,et al.  Goal Representation in Human Anterior Intraparietal Sulcus , 2006, The Journal of Neuroscience.

[171]  Scott T. Grafton,et al.  Beyond grasping: Representation of action in human anterior intraparietal sulcus , 2007, NeuroImage.

[172]  Scott T. Grafton,et al.  Cortical topography of human anterior intraparietal cortex active during visually guided grasping. , 2005, Brain research. Cognitive brain research.

[173]  H. Wimmer,et al.  Beliefs about beliefs: Representation and constraining function of wrong beliefs in young children's understanding of deception , 1983, Cognition.

[174]  E. Redcay The superior temporal sulcus performs a common function for social and speech perception: Implications for the emergence of autism , 2008, Neuroscience & Biobehavioral Reviews.

[175]  Keiji Tanaka,et al.  The role of the medial prefrontal cortex in achieving goals , 2004, Current Opinion in Neurobiology.

[176]  Mahzarin R. Banaji,et al.  Neural Correlates of Stereotype Application , 2009, Journal of Cognitive Neuroscience.

[177]  Rieko Osu,et al.  The Neural Substrates of Biological Motion Perception: an fMRI Study , 2022 .

[178]  T. Heatherton,et al.  Detecting agency from the biological motion of veridical vs animated agents. , 2007, Social cognitive and affective neuroscience.

[179]  N. George,et al.  Facing the gaze of others , 2008, Neurophysiologie Clinique/Clinical Neurophysiology.

[180]  Jonathan D. Nelson,et al.  Human cortical representations for reaching: Mirror neurons for execution, observation, and imagery , 2007, NeuroImage.

[181]  C. Keysers,et al.  Empathy and the Somatotopic Auditory Mirror System in Humans , 2006, Current Biology.

[182]  J. Haxby,et al.  Parallel Visual Motion Processing Streams for Manipulable Objects and Human Movements , 2002, Neuron.

[183]  P. Fonlupt Perception and judgement of physical causality involve different brain structures. , 2003, Brain research. Cognitive brain research.

[184]  R. Passingham,et al.  Brain Mechanisms for Inferring Deceit in the Actions of Others , 2004, The Journal of Neuroscience.

[185]  R Saxe,et al.  People thinking about thinking people The role of the temporo-parietal junction in “theory of mind” , 2003, NeuroImage.

[186]  T. Allison,et al.  Temporal Cortex Activation in Humans Viewing Eye and Mouth Movements , 1998, The Journal of Neuroscience.

[187]  S. Blakemore,et al.  Social cognitive neuroscience: where are we heading? , 2004, Trends in Cognitive Sciences.

[188]  Jonathan D. Cohen,et al.  An fMRI Investigation of Emotional Engagement in Moral Judgment , 2001, Science.

[189]  G. Rizzolatti,et al.  Neural Circuits Underlying Imitation Learning of Hand Actions An Event-Related fMRI Study , 2004, Neuron.

[190]  M. Makuuchi Is Broca's area crucial for imitation? , 2005, Cerebral cortex.

[191]  Nava Rubin,et al.  Brain areas selective for both observed and executed movements. , 2007, Journal of neurophysiology.

[192]  Volkmar Glauche,et al.  The human action recognition system and its relationship to Broca’s area: an fMRI study , 2003, NeuroImage.

[193]  J. Gottlieb From Thought to Action: The Parietal Cortex as a Bridge between Perception, Action, and Cognition , 2007, Neuron.

[194]  K. Zilles,et al.  Mind Reading: Neural Mechanisms of Theory of Mind and Self-Perspective , 2001, NeuroImage.

[195]  G. Rizzolatti,et al.  Neural Circuits Involved in the Recognition of Actions Performed by Nonconspecifics: An fMRI Study , 2004, Journal of Cognitive Neuroscience.

[196]  F. Heider,et al.  An experimental study of apparent behavior , 1944 .

[197]  Scott T. Grafton,et al.  A distributed left hemisphere network active during planning of everyday tool use skills. , 2004, Cerebral cortex.

[198]  A. Schnitzler,et al.  Do simple intransitive finger movements consistently activate frontoparietal mirror neuron areas in humans? , 2007, NeuroImage.

[199]  Andrew D. Engell,et al.  The Neural Bases of Cognitive Conflict and Control in Moral Judgment , 2004, Neuron.

[200]  Christian Keysers,et al.  Testing Simulation Theory with Cross-Modal Multivariate Classification of fMRI Data , 2008, PloS one.

[201]  Gregory McCarthy,et al.  Taking an “intentional stance” on eye-gaze shifts: A functional neuroimaging study of social perception in children , 2005, NeuroImage.

[202]  Olaf B. Paulson,et al.  Similar brain networks for detecting visuo-motor and visuo-proprioceptive synchrony , 2006, NeuroImage.

[203]  Jean Decety,et al.  Who caused the pain? An fMRI investigation of empathy and intentionality in children , 2008, Neuropsychologia.

[204]  Robert Oostenveld,et al.  Population activity in the human dorsal pathway predicts the accuracy of visual motion detection. , 2007, Journal of neurophysiology.

[205]  B. Puri,et al.  The human mirror system: A motor resonance theory of mind-reading , 2007, Brain Research Reviews.

[206]  Jason P. Mitchell Activity in right temporo-parietal junction is not selective for theory-of-mind. , 2008, Cerebral cortex.

[207]  M. Iacoboni,et al.  The self and social cognition: the role of cortical midline structures and mirror neurons , 2007, Trends in Cognitive Sciences.

[208]  G. Rizzolatti,et al.  Parietal Lobe: From Action Organization to Intention Understanding , 2005, Science.

[209]  P. Christiaan Klink,et al.  General Validity of Levelt's Propositions Reveals Common Computational Mechanisms for Visual Rivalry , 2008, PloS one.

[210]  M. Kronbichler,et al.  Thinking of mental and other representations: The roles of left and right temporo-parietal junction , 2006, Social neuroscience.

[211]  P. Sinha,et al.  Functional neuroanatomy of biological motion perception in humans , 2001, Proceedings of the National Academy of Sciences of the United States of America.

[212]  Mark Mühlau,et al.  Left inferior parietal dominance in gesture imitation: an fMRI study , 2005, Neuropsychologia.

[213]  Ricarda Ines Schubotz,et al.  I see you moving - Motion class dependency in observers' non-primary motor areas revealed by functional Magnetic Resonance Imaging (fMRI) , 2005 .

[214]  A. Meltzoff,et al.  The detection of contingency and animacy from simple animations in the human brain. , 2003, Cerebral cortex.

[215]  Gereon R. Fink,et al.  Cue validity modulates the neural correlates of covert endogenous orienting of attention in parietal and frontal cortex , 2006, NeuroImage.

[216]  C. Frith,et al.  Social interaction modifies neural response to gaze shifts. , 2007, Social cognitive and affective neuroscience.

[217]  T. Ito,et al.  The influence of processing objectives on the perception of faces: An ERP study of race and gender perception , 2005, Cognitive, affective & behavioral neuroscience.

[218]  Luca Turella,et al.  When Gaze Turns into Grasp , 2006, Journal of Cognitive Neuroscience.

[219]  E. Vakil,et al.  Motor and non-motor sequence learning in patients with basal ganglia lesions: the case of serial reaction time (SRT) , 2000, Neuropsychologia.

[220]  Lindsey J. Powell,et al.  It's the Thought That Counts , 2006, Psychological science.

[221]  J. Downar,et al.  The Effect of Task Relevance on the Cortical Response to Changes in Visual and Auditory Stimuli: An Event-Related fMRI Study , 2001, NeuroImage.

[222]  Vittorio Gallese,et al.  Listening to Action-related Sentences Activates Fronto-parietal Motor Circuits , 2005, Journal of Cognitive Neuroscience.

[223]  G. Orban,et al.  Specificity of regions processing biological motion , 2005, The European journal of neuroscience.

[224]  T. Allison,et al.  Functional anatomy of biological motion perception in posterior temporal cortex: an FMRI study of eye, mouth and hand movements. , 2005, Cerebral cortex.

[225]  R. Saxe,et al.  Making sense of another mind: The role of the right temporo-parietal junction , 2005, Neuropsychologia.

[226]  M. Sereno,et al.  Point-Light Biological Motion Perception Activates Human Premotor Cortex , 2004, The Journal of Neuroscience.

[227]  Cheryl L. Dickter,et al.  Racial ingroup and outgroup attention biases revealed by event-related brain potentials. , 2007, Social cognitive and affective neuroscience.

[228]  Isabell Wartenburger,et al.  Individual differences in moral judgment competence influence neural correlates of socio-normative judgments. , 2008, Social cognitive and affective neuroscience.

[229]  Gaspare Galati,et al.  A selective representation of the meaning of actions in the auditory mirror system , 2008, NeuroImage.

[230]  Nadim Joni Shah,et al.  Neural Representations of Self versus Other: Visual-Spatial Perspective Taking and Agency in a Virtual Ball-tossing Game , 2006, Journal of Cognitive Neuroscience.

[231]  R. Passingham,et al.  Action observation and acquired motor skills: an FMRI study with expert dancers. , 2005, Cerebral cortex.

[232]  P. Eslinger,et al.  Frontopolar and anterior temporal cortex activation in a moral judgment task: preliminary functional MRI results in normal subjects. , 2001, Arquivos de neuro-psiquiatria.

[233]  Isabell Wartenburger,et al.  Influence of bodily harm on neural correlates of semantic and moral decision-making , 2005, NeuroImage.

[234]  J. Decety,et al.  Neural correlates of feeling sympathy , 2003, Neuropsychologia.

[235]  J. Downar,et al.  A cortical network sensitive to stimulus salience in a neutral behavioral context across multiple sensory modalities. , 2002, Journal of neurophysiology.