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Corticospinal excitability (CSE) in humans measured with Transcranial Magnetic Stimulation (TMS) is generally increased by the perception of other people's actions. This perception can be unimodal (visual or auditory) or multimodal (visual and auditory). The increase in TMS-measured CSE is typically prominent for muscles involved in the perceived action (muscle specificity). There are two main classes of accounts for this phenomenon. One suggests that the motor system mirrors the actions that the observer perceives (the resonance account). The other suggests that the motor system predicts the actions that the observer perceives (the predictive account). To test these accounts (which need not be mutually exclusive), subjects were presented with four versions of three-note piano sequences: sound only, sight only, audiovisual, and audiovisual with sound lagging behind (the prediction violation condition). CSE was measured in two hand muscles used to play the notes. CSE increased reliably in one muscle only for the prediction violation condition, in line with the predictive account, while the other muscle demonstrated CSE increase for all conditions, in line with the resonance account. This finding supports both predictive coding accounts as well as resonance accounts of motor facilitation during action perception.

Stimulus-response compatibility (SRC)-the fact that some stimulus-response pairs are faster than others-is attributed in part to automatic activation of the stimulus-compatible response representation. Cognitive models of SRC propose that automatic response activation can be strategically suppressed if the automatic response is likely to interfere with behavior; in particular, suppression is thought to occur in preparation for incompatible responses and when the required stimulus-response mapping is unknown before stimulus presentation. We test this preparatory suppression hypothesis in the context of imitation, a special form of SRC particularly relevant to human social behavior. Using TMS, we measured muscle-specific corticospinal excitability during action observation (motor resonance) while human participants prepared to perform imitative and counterimitative responses to action videos. Motor resonance was suppressed during preparation to counterimitate and for unknown mappings, compared to preparation to imitate and a baseline measure of motor resonance. These results provide novel neurophysiological evidence that automatic activation of stimulus-compatible responses can be strategically suppressed when the automatic response is likely to interfere with task goals. Insofar as motor resonance measures mirror neuron system activity, these results also suggest that preparatory control of automatic imitative tendencies occurs through modulation of mirror neuron system activity.

Embodied metaphor theory suggests abstract concepts are metaphorically linked to more experientially basic ones and recruit sensorimotor cortex for their comprehension. To test whether words associated with spatial attributes reactivate traces in sensorimotor cortex, we recorded EEG from the scalp of healthy adults as they read words while performing a concurrent task involving either upward- or downward- directed arm movements. ERPs were time-locked to words associated with vertical space-either literally (ascend, descend) or metaphorically (inspire, defeat)-as participants made vertical movements that were either congruent or incongruent with the words. Congruency effects emerged 200-300 ms after word onset for literal words, but not until after 500 ms post-onset for metaphorically related words. Results argue against a strong version of embodied metaphor theory, but support a role for sensorimotor simulation in concrete language.