Prophage Provide a Safe Haven for Adaptive Exploration in Temperate Viruses

Prophage sequences constitute a substantial fraction of the temperate virus gene pool. Although subject to mutational decay, prophage sequences can also be an important source of adaptive mutations for these viral populations. Here we develop a life-history model for temperate viruses, including both the virulent (lytic) and the temperate phases of the life cycle. We then examine the survival of mutations that increase fitness during the lytic phase (attachment rate, burst size), increase fitness in the temperate phase (increasing host survival), or affect transitions between the two phases (integration or induction probability). We find that beneficial mutations are much more likely to survive, ultimately, if they first occur in the prophage state. This conclusion applies even to traits that are only expressed during the lytic phase, and arises due to the substantially lower variance in the offspring distribution during the temperate cycle. This observation, however, is balanced by the fact that many more mutations can be generated during lytic replication. Overall we predict that the prophage state provides a refuge, relatively shielded from genetic drift, in which temperate viruses can explore possible adaptive steps.