We used the quantitative 14 C-deoxyglucose method to map the activity pattern throughout the frontal cortex of rhesus monkeys, which either grasped a three-dimensional object or observed the same grasping movements executed by a human. We found that virtually the same frontal cortical networks were recruited for the generation and the perception of action, including the primary motor cortex (MI/F1), premotor cortical areas (F2, F5, and F6), the primary (SI) and supplementary (SSA) somatosensory cortex, medial cortical areas (8m and 9m), and the anterior cingulate. The overlapping networks for action execution and action observation support the notion that mental simulation of action could underlie the perception of others' actions. We suggest that the premotor and the somatotopic MI/F1 activations induced by action observation reflect motor grasp of the observed action, whereas the somatotopic SI and the SSA activations reflect recruitment of learned sensory-motor associations enabling perceptual understanding of the anticipated somatosensory feedback. We also found that the premotor activations were stronger for action observation, in contrast to the primary somatosensory-motor ones, which were stronger for action execution, and that activations induced by observation were bilateral, whereas those induced by execution were contralateral to the moving forelimb. We suggest that these differences in intensity and lateralization of activations between the executive and the perceptual networks help attribute the action to the correct agent, i.e., to the "self" during action execution and to the "other" during action observation. Accordingly, the "sense of agency" could be articulated within the core components of the circuitry supporting action execution/observation.
We used the (14)C-deoxyglucose method to map the functional activity in the cortex of the lateral and medial parietal convexity, the intraparietal and the parietoccipital sulci of monkeys which either reached and grasped a 3D-object or observed the same reaching-to-grasp movements executed by a human. Execution of reaching-to-grasp induced activations in the superior parietal areas SI-forelimb/convexity, PE, PE caudal (PEc); in the intraparietal areas PE intraparietal (PEip), medial intraparietal (MIP), 5 intraparietal posterior, ventral intraparietal (VIP), anterior intraparietal (AIP), lateral intraparietal dorsal; in the inferior parietal areas PF, PFG, PG; in the parietoccipital areas V6, V6A-dorsal; in the medial cortical areas PGm/7m and retrosplenial cortex. Observation of reaching-to-grasp activated areas SI-forelimb/convexity, PE lateral, PEc, PEip, MIP, VIP, AIP, PF, V6, PGm/7m, 31, and retrosplenial cortex. The common activations were stronger for execution than for observation and the interhemispheric differences were smaller for observation than for execution, contributing to the attribution of action to the correct agent. The extensive overlap of parietal networks activated for action execution and observation supports the "mental simulation theory" which assigns the role of understanding others' actions to the entire distributed neural network responsible for the execution of actions, and not the concept of "mirroring" which reflects the function of a certain class of cells in a couple of cortical areas.
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