Virulence Evolution of Pathogens That Can Grow in Reservoir Environments

Abstract
Many pathogens reside in environmental reservoirs within which they can reproduce and from which they can infect hosts. These facultative pathogens experience different selective pressures in host-associated environments and reservoir environments. Heterogeneous selective pressures have the potential to influence the virulence evolution of these pathogens. Previous research has examined how environmental transmission influences the selective pressures shaping the virulence of pathogens that cannot reproduce in environmental reservoirs, yet many pathogens of humans, crop plants, and livestock can reproduce in these environments. We build on this work to examine how reproduction in reservoirs influences disease dynamics and virulence evolution in a simple facultative pathogen model. We use adaptive dynamics to examine the evolutionary dynamics of facultative pathogens under potential trade-offs between transmission and virulence, shedding and virulence, and reservoir persistence and virulence. We then perform critical function analysis to generalize the results independent of specific trade-off assumptions. We determine that diverse virulence strategies, sometimes resulting from evolutionary bistability or evolutionary branching conditions, are expected for facultative pathogens. Our findings motivate research establishing which trade-offs most strongly influence the virulence evolution of facultative pathogens.