A general framework for modelling trade-offs in adaptive behaviour

An animal's behaviour can influence many variables, such as its energy reserves, its risk of injury or mortality, and its rate of reproduction. To identify the optimal action in a given situation, these various effects can be compared in the common currency of reproductive value. While this idea has been widely used to study trade-offs between pairs of variables, e.g. between energy gain versus survival, here we present a unified framework that makes explicit how these various trade-offs fit together. This unification covers a wide range of biological phenomena, highlighting similarities in their logical structure and helping to identify knowledge gaps. To fill one such gap, we present a new model of foraging under the risk of predation and damage accumulation. We conclude by discussing the use and limitations of state-dependent optimisation theory in predicting biological observations.