Testosterone as a mediator of the tradeoff between cooperation and competition in the context of cooperative reproductive behaviors

Behavioral tradeoffs occur when the expression of one behavior detracts from the expression of another. Understanding the proximate mediators of behavioral tradeoffs is important as these tradeoffs can act as potential constraints on evolutionary responses to selection. Here, we describe the tradeoff between cooperation and competition faced by species that exhibit cooperative reproductive behaviors and propose that testosterone is a key hormonal mediator of the tradeoff. Cooperative reproductive behaviors occur when multiple individuals coordinate their efforts to gain a reproductive advantage over other individuals and/or those individuals attempting to reproduce in absence of cooperation. We propose that testosterone, a sex steroid known to mediate a number of physiological and behavioral actions associated with reproductive competition, is involved in mediating the tradeoff between cooperation and competition. To support this proposition, we first describe the importance of individual variation in behavior to the evolution of cooperative behaviors. We then describe how proximate mechanisms represent a prominent source of individual variation in social behaviors and highlight evidence suggesting testosterone mediates variation in cooperative behaviors. Two case studies in which the relationship between testosterone and cooperative behaviors have been investigated in detail are then summarized. Throughout we highlight the importance of studying individual variation to understand the mechanistic basis of behaviors, behavioral tradeoffs, and the evolution of cooperative reproductive behaviors more broadly.