Are there brain regions that are specialized for the execution of imitative actions? We compared two hypotheses of imitation: the mirror neuron system (MNS) hypothesis predicts frontal and parietal engagement which is specific to imitation, while the Grist-Mills hypothesis predicts no difference in brain activation between imitative and matched non-imitative actions. Our delayed imitation fMRI paradigm included two tasks, one where correct performance was defined by a spatial rule and another where it was defined by an item-based rule. For each task, participants could learn a sequence from a video of a human hand performing the task, from a matched "Ghost" condition, or from text instructions. When participants executed actions after seeing the Hand demonstration (compared to Ghost and Text demonstrations), no activation differences occurred in frontal or parietal regions; rather, activation was localized primarily to occipital cortex. This adds to a growing body of evidence which indicates that imitation-specific responses during action execution do not occur in canonical mirror regions, contradicting the mirror neuron system hypothesis. However, activation differences did occur between action execution in the Hand and Ghost conditions outside MNS regions, which runs counter to the Grist-Mills hypothesis. We conclude that researchers should look beyond these hypotheses as well as classical MNS regions to describe the ways in which imitative actions are implemented by the brain.
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