The organization and cognitive role of the mirror system

Observation of actions performed by others activates monkey ventral premotor cortex, where action meaning, but not object identity, is coded. In a functional MRI (fMRI) study, we investigated whether other monkey frontal areas respond to actions performed by others. Observation of a hand grasping objects activated four frontal areas: rostral F5 and areas 45B, 45A, and 46. Observation of an individual grasping an object also activated caudal F5, which indicates different degrees of action abstraction in F5. Observation of shapes activated area 45, but not premotor F5. Convergence of object and action information in area 45 may be important for full comprehension of actions. These data have been published on Science, 2005, 310: 332-336.

When we observe someone performing an action, do our brains simulate making that action? Acquired motor skills offer a unique way to test this question, since people differ widely in the actions they have learned to perform. We used functional magnetic resonance imaging to study differences in brain activity between watching an action that one has learned to do and an action that one has not, in order to assess whether the brain processes of action observation are modulated by the expertise and motor repertoire of the observer. Experts in classical ballet, experts in capoeira and inexpert control subjects viewed videos of ballet or capoeira actions. Comparing the brain activity when dancers watched their own dance style versus the other style therefore reveals the influence of motor expertise on action observation. We found greater bilateral activations in premotor cortex and intraparietal sulcus, right superior parietal lobe and left posterior superior temporal sulcus when expert dancers viewed movements that they had been trained to perform compared to movements they had not. Our results show that this 'mirror system' integrates observed actions of others with an individual's personal motor repertoire, and suggest that the human brain understands actions by motor simulation. These data have been published on Cerebral Cortex, 2005, 15: 1243-1249.

Processing of complex visual stimuli comprising facial movements, hand actions, and body movements is known to occur in the superior temporal sulcus (STS) of humans and nonhuman primates. The STS is also thought to play a role in the integration of multimodal sensory input. We investigated whether STS neurons coding the sight of actions also integrated the sound of those actions. For 23% of neurons responsive to the sight of an action, the sound of that action significantly modulated the visual response. The sound of the action increased or decreased the visually evoked response for an equal number of neurons. In the neurons whose visual response was increased by the addition of sound (but not those neurons whose responses were decreased), the audiovisual integration was dependent upon the sound of the action matching the sight of the action. These results suggest that neurons in the STS form multisensory representations of observed actions. These data have been published on Journal of Cognitive Neuroscience, 2005, 17: 377-391.

Inferior parietal lobule (IPL) neurons were studied when monkeys performed motor acts embedded in different actions and when they observed similar acts done by an experimenter. Most motor IPL neurons coding a specific act (e.g., grasping) showed markedly different activations when this act was part of different actions (e.g., for eating or for placing). Many motor IPL neurons also discharged during the observation of acts done by others. Most responded differentially when the same observed act was embedded in a specific action. These neurons fired during the observation of an act, before the beginning of the subsequent acts specifying the action. Thus, these neurons not only code the observed motor act but also allow the observer to understand the agent's intentions. The data have been published in Science, 2005, 308: 662-667.

Engagement of the primary motor cortex (MI) during the observation of actions has been debated for a long time. In the present study, we used the quantitative 14C-deoxyglucose method in monkeys that either grasped 3-D objects or observed the same movements executed by humans. We found that the forelimb regions of the MI and the primary somatosensory (SI) cortex were significantly activated in both cases. Our study resolves a debate in the literature, providing strong evidence for use of MI representations during the observation of actions. It demonstrates that the observation of an action is represented in the primary motor and somatosensory cortices as is its execution. It indicates that in terms of neural correlates, recognizing a motor behaviour is like executing the same behaviour, requiring the involvement of a distributed system encompassing not only the premotor but also the primary motor cortex. We suggest that movements and their proprioceptive components are stored as motor and somatosensory representations in motor and somatosensory cortices, respectively, and that these representations are recalled during observation of an action. These data have been published in Neuroimage, 2004, 23: 193-201.

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