How feedback and feed-forward mechanisms link determinants of social dominance
In many animal societies, individuals differ consistently in their ability to win agonistic interactions, resulting in dominance hierarchies. These differences arise due to a range of factors that can influence individuals’ abilities to win agonistic interactions, spanning from genetically driven traits through to individuals’ recent interaction history. Yet, despite a century of study since Schjelderup-Ebbe’s seminal paper on social dominance, we still lack a general understanding of how these different factors work together to determine individuals’ positions in hierarchies. Here, we first outline five widely studied factors that can influence interaction outcomes: intrinsic attributes, resource value asymmetry, winner-loser effects, dyadic interaction-outcome history and third-party support. A review of the evidence shows that whilst different factors have been shown to be important in specific systems, there are few empirical cases where one factor has a definitive effect. We then propose that mixed empirical support for a single factor is likely to arise due to feedback loops, whereby the outcomes of previous agonistic interactions (e.g. access to food) impact factors that might be important in subsequent interactions (e.g. body condition). We provide a conceptual framework which illustrates that there are many potential routes through which feedbacks can occur. Such feedbacks suggest that the factors that determine outcomes of dominance interactions are highly intertwined and are likely to rarely act independently of one-another. Further, we generalise our framework to include multi-generational feed-forward mechanisms and highlight how interaction outcomes in one generation can influence the factors determining interaction outcomes of their offspring via a range of parental effects. This general framework describes how interaction outcomes and the factors determining them are linked within generations via feedback loops, and between generations via feed-forward mechanisms. We then highlight methodological approaches that will facilitate the study of feedback loops and dominance dynamics. Lastly, we discuss how our framework can shape future research, including investigating how feedbacks in dominance hierarchies produce ‘self-organised’ structure, exploring how interaction outcomes are integrated to form dominance hierarchies, and the routes of parental influence on the dominance status of offspring. Ultimately, by considering dominance interactions as part of a dynamic system, that also feeds forward into subsequent generations, we will better understand the factors that structure dominance hierarchies in animal groups.