The Price equation and evolutionary epidemiology

The Price equation has found widespread application in many areas of evolutionary biology, including the evolutionary epidemiology of infectious diseases. In this paper, we illustrate the utility of this approach to modelling disease evolution by first deriving a version of Price’s equation that can be applied in continuous time and to populations with overlapping generations. We then show how this version of Price’s equation provides an alternative perspective on pathogen evolution by considering the epidemiological meaning of each of its terms. Finally, we extend these results to the case where population size is small and generates demographic stochasticity. We show that the particular partitioning of evolutionary change given by Price’s equation is also a natural way to partition the evolutionary consequences of demographic stochasticity, and demonstrate how such stochasticity tends to weaken selection on birth rate (e.g. the transmission rate of an infectious disease) and enhance selection on mortality rate (e.g. factors, like virulence, that cause the end of an infection). In the long term, if there is a trade-off between virulence and transmission across parasite strains, the weaker selection on transmission and stronger selection on virulence that arises from demographic stochasticity will tend to drive the evolution of lower levels of virulence. This article is part of the theme issue ‘Fifty years of the Price equation’.

Price's equation made clear

Price's equation provides a very simple—and very general—encapsulation of evolutionary change. It forms the mathematical foundations of several topics in evolutionary biology, and has also been applied outwith evolutionary biology to a wide range of other scientific disciplines. However, the equation's combination of simplicity and generality has led to a number of misapprehensions as to what it is saying and how it is supposed to be used. Here, I give a simple account of what Price's equation is, how it is derived, what it is saying and why this is useful. In particular, I suggest that Price's equation is useful not primarily as a predictor of evolutionary change but because it provides a general theory of selection. As an illustration, I discuss some of the insights Price's equation has brought to the study of social evolution. This article is part of the theme issue ‘Fifty years of the Price equation’.