Epidemic evolutionarily stable strategies within an age-structured host population

To understand infectious disease dynamics, we need to understand the inextricably intertwined nature of the ecology and evolution of pathogens and hosts. Epidemiological dynamics of many infectious diseases have highlighted the importance of considering the demographics of the societies in which they spread, particularly with respect to age structure. In addition, the waves of the recent COVID-19 pandemic driven by variant replacements at an unprecedented speed show that it is vital to consider the evolutionary aspects. The classic trade-off theory of virulence addresses aspects of pathogen evolution, but here we explore in more detail the possibility of society-specific evolutionarily stable strategies (ESS) during an unfolding pandemic. Theory posits the existence under some conditions of an ESS representing the evolutionary endpoint of change. By using a demographically realistic model incorporating infection rates that vary with age, we outline which evolutionary scenarios are plausible. Focusing on the rate of infection and duration of infectivity, we ask whether an ESS exists, what characterizes it, and as a result which long-term public-health consequences may be expected. We demonstrate that the ESS of an evolving pathogen depends upon the background age-dependent frailty and mortality rates. Our findings shed important light on the plausible long-term trajectories of highly evolvable novel pathogens.