Faces in the mirror, from the neuroscience of mimicry to the emergence of mentalizing.

In the current opinion paper, we provide a comparative perspective on specific aspects of primate empathic abilities, with particular emphasis on the mirror neuron system associated with mouth/face actions and expression. Mouth and faces can be very salient communicative classes of stimuli that allow an observer access to the emotional and physiological content of other individuals. We thus describe patterns of activations of neural populations related to observation and execution of specific mouth actions and emotional facial expressions in some species of monkeys and in humans. Particular attention is given to dynamics of face-to-face interactions in the early phases of development and to the differences in the anatomy of facial muscles among different species of primates. We hypothesize that increased complexity in social environments and patterns of social development have promoted specializations of facial musculature, behavioral repertoires related to production and recognition of facial emotional expression, and their neural correlates. In several primates, mirror circuits involving parietal-frontal regions, insular regions, cingulate cortices, and amygdala seem to support automatic forms of embodied empathy, which probably contribute to facial mimicry and behavioural synchrony. In humans these circuits interact with specific prefrontal and temporo-parietal cortical regions, which facilitates higher order cognitive functions such as cognitive empathy and mental state attribution. Our analysis thus suggests that the evolution of higher forms of empathy, such as mentalizing, is also linked to the coupling between the perceptual and motor system related to face processing, which may have undergone a process of exaptation during primate phylogeny.