The effect of temporal fluctuations on the evolution of host tolerance to parasitism.

There are many mechanisms that hosts can evolve to defend against parasites, two of which are resistance and tolerance. These defences often have different evolutionary behaviours, and it is important to consider how each individual mechanism may respond to changes in environment. In particular, host defence through tolerance is predicted to be unlikely to lead to variation, despite many observations of diversity in both animal and plant systems. Hence understanding the drivers of diversity in host defence and parasite virulence is vital for predicting future evolutionary changes in infectious disease dynamics. It has been suggested that heterogeneous environments might generally promote diversity, but the effect of temporal fluctuations has received little attention theoretically or empirically, and there has been no examination of how temporal fluctuations affects the evolution of host tolerance. In this study, we use a mathematical model to investigate the evolution of host tolerance in a temporally fluctuating environment. We show that investment in tolerance increases in more variable environments, giving qualitatively different evolutionary behaviours when compared to resistance. Once seasonality is introduced evolutionary branching though tolerance can occur and create diversity within the population, although potentially only temporarily. This branching behaviour arises due to the emergence of a negative feedback with the maximum infected density on a cycle, which is strongest when the infected population is large. This work reinforces the qualitative differences between tolerance and resistance evolution, but also provides theoretical evidence for the theory that heterogeneous environments promote host-parasite diversity, hence constant environment assumptions may omit important evolutionary outcomes.