  Giant viruses contain large genomes, encode many proteins atypical for
viruses, replicate in large viral factories, and tend to infect protists. The
giant virus replication factories can in turn be infected by so called
virophages, which are smaller viruses that negatively impact giant virus
replication. An example are Mimiviruses that infect the protist Acanthamoeba
and that are themselves infected by the virophage Sputnik. This paper examines
the evolutionary dynamics of this system, using mathematical models. While the
models suggest that the virophage population will evolve to increasing degrees
of giant virus inhibition, it further suggests that this renders the virophage
population prone to extinction due to dynamic instabilities over wide parameter
ranges. Implications and conditions required to avoid extinction are discussed.
Another interesting result is that virophage presence can fundamentally alter
the evolutionary course of the giant virus. While the giant virus is predicted
to evolve towards increasing its basic reproductive ratio in the absence of the
virophage, the opposite is true its presence. Therefore, virophages can not
only benefit the host population directly by inhibiting the giant viruses, but
also indirectly by causing giant viruses to evolve towards weaker phenotypes.
Experimental tests for this model are suggested.
