Composite temporal strategies in pathogen evolution: balancing invasion and persistence

There is ongoing interest in the conditions that favor the evolution of acute, highly transmissible infections in contrast to chronic ones. Earlier studies typically consider the evolution of a trait that is constant over the lifetime of an infection. However, for many pathogens, such traits can vary over this course. Here, we address the evolution of temporal patterns in limited host population sizes, where a trade-off between invasion and persistence can arise. This is of particular relevance to questions on the evolution of acuteness and chronicity. We ask whether population dynamics of transmission at the between-host level could lead pathogen adaptation to favor temporal strategies during the course of infection. To do this, we consider an infection to be composed of multiple stages, allowing each of these to evolve independently under a transmission–duration trade-off. We only consider selection taking place on the between-host level and examine the balance of invasion and persistence (i.e., maximizing replication vs. minimizing vulnerability to extinction), using several fitness-related measures. We find that a composite strategy that is ordered in time can confer higher fitness than any single, constant, strategy. We discuss the relevance of these results for the ordered expression of var genes in Plasmodium falciparum, as well as for infections that characteristically have several stages as in some bacterial pathogens.