Levels of selection in positive-strand virus dynamics

Conflicting selection pressures occurring over the life cycle of an organism constitute serious challenges to the robustness of replication. Viruses present a credible model system for analysing problems that arise through evolutionary conflicts of interest. We present a multi-level selection model for the life cycle of positive-strand RNA viruses. The model combines within-cell replication kinetics and protein synthesis, and between-cell population dynamics of virion production and transmission. We show how these two levels of within-host selection interact to produce tradeoffs in the life history strategy of a virus without consideration of host mortality. We find that viruses evolve towards intermediate rather than maximum encapsidation rates. This can be interpreted as selection for intermediate virulence through cellular persistence. We characterize a theoretical persistence threshold arising from the trade-off between genome replication and genetic translation within the cell. We present counter-intuitive relationships whereby increasing genome decay rates and rates of encapsidation lead to increases in the abundance of virus-encoded proteins. Data from poliovirus suggest that viruses might be unable to resolve the vertical conflicts of interests among different levels of selection.