Neural foundations of overt and covert actions.

We used fMRI to assess the human brain areas activated for execution, observation and 1st person motor imagery of a visually guided tracing task with the index finger. Voxel-level conjunction analysis revealed several cortical areas activated in common across all three motor conditions, namely, the upper limb representation of the primary motor and somatosensory cortices, the dorsal and ventral premotor, the superior and inferior parietal cortices as well as the posterior part of the superior and middle temporal gyrus including the temporo-parietal junction (TPj) and the extrastriate body area (EBA). Functional connectivity analyses corroborated the notion that a common sensory-motor fronto-parieto-temporal cortical network is engaged for execution, observation, and imagination of the very same action. Taken together these findings are consistent with the more parsimonious account of motor cognition provided by the mental simulation theory rather than the recently revised mirror neuron view Action imagination and observation were each associated with several additional functional connections, which may serve the distinction between overt action and its covert counterparts, and the attribution of action to the correct agent. For example, the central position of the right middle and inferior frontal gyrus in functional connectivity during motor imagery may reflect the suppression of movements during mere imagination of action, and may contribute to the distinction between 'imagined' and 'real' action. Also, the central role of the right EBA in observation, assessed by functional connectivity analysis, may be related to the attribution of action to the 'external agent' as opposed to the 'self'.