Integrating Tinbergen's inquiries: Mimicry and play in humans and other social mammals.

Visual signals convey emotions and intentions between individuals. Darwin underlined that human facial expressions represent a shared heritage between our species and many other social mammals. Social play is a fertile field to examine the role and the potential communicative function of facial expressions. The relaxed open-mouth (or play face) is a context-specific playful expression, which is widespread in human and non-human mammals. Here, we focus on playful communication by applying Tinbergen's four areas of inquiry: proximate causation, ontogeny, function, and evolution. First of all we explore mimicry by focusing on its neural substrates and factors of modulation within playful and non-playful context (proximate causation). Play face is one of the earliest facial expressions to appear and be mimicked in neonates. The motor resonance between infants and their caregivers is essential later in life when individuals begin to engage in increasingly complex social interactions, including play (ontogeny). The success of a playful session can be evaluated by its duration in time. Mirroring facial expressions prolongs the session by favoring individuals to fine-tune their own motor sequences accordingly (function). Finally, through a comparative approach we also demonstrate that the elements constituting play communication and mimicry are sensitive to the quality of interindividual relationships of a species, thus reflecting the nature of its social network and style (evolution). In conclusion, our goal is to integrate Tinbergen's four areas of ethological inquiry to provide a broader framework regarding the importance of communication and mimicry in the play domain of humans and other social mammals.