Hamilton's rule, gradual evolution, and the optimal (feedback) control of phenotypically plastic traits

Most traits expressed by organisms, such as gene expression profiles, developmental trajectories, behavioural sequences and reaction norms are function-valued traits (colloquially phenotypically plastic traits), since they vary across an individual's age and in response to various internal and/or external factors (state variables). Furthermore, most organisms live in populations subject to limited genetic mixing and are thus likely to interact with their relatives. We here formalise selection on genetically determined function-valued traits of individuals interacting in a group-structured population, by deriving the marginal version of Hamilton's rule for function-valued traits. This rule simultaneously gives a condition for the invasion of an initially rare mutant function-valued trait and its ultimate fixation in the population (invasion thus implies substitution). Hamilton's rule thus underlies the gradual evolution of function-valued traits and gives rise to necessary first-order conditions for their uninvadability (evolutionary stability). We develop a novel analysis using optimal control theory and differential game theory, to simultaneously characterise and compare the first-order conditions of (i) open-loop traits - functions of time (or age) only, and (ii) closed-loop (state-feedback) traits - functions of both time and state variables. We show that closed-loop traits can be represented as the simpler open-loop traits when individuals do not interact or when they interact with clonal relatives. Our analysis delineates the role of state-dependence and interdependence between individuals for trait evolution, which has implications to both life-history theory and social evolution. (C) 2021 The Authors. Published by Elsevier Ltd.

Automated summary

This study formalizes selection on genetically determined function-valued traits of individuals interacting in a group-structured population, deriving the marginal version of Hamilton's rule. It explores the conditions for the invasion of mutant function-valued traits and their eventual fixation in the population. Using optimal control theory and differential game theory, the study characterizes open-loop and closed-loop traits, showing the impact of state-dependence and individual interdependence on trait evolution.