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Giant viruses contain large genomes, encode many proteins atypical forviruses, replicate in large viral factories, and tend to infect protists. Thegiant virus replication factories can in turn be infected by so calledvirophages, which are smaller viruses that negatively impact giant virusreplication. An example are Mimiviruses that infect the protist Acanthamoebaand that are themselves infected by the virophage Sputnik. This paper examinesthe evolutionary dynamics of this system, using mathematical models. While themodels suggest that the virophage population will evolve to increasing degreesof giant virus inhibition, it further suggests that this renders the virophagepopulation prone to extinction due to dynamic instabilities over wide parameterranges. Implications and conditions required to avoid extinction are discussed.Another interesting result is that virophage presence can fundamentally alterthe evolutionary course of the giant virus. While the giant virus is predictedto evolve towards increasing its basic reproductive ratio in the absence of thevirophage, the opposite is true its presence. Therefore, virophages can notonly benefit the host population directly by inhibiting the giant viruses, butalso indirectly by causing giant viruses to evolve towards weaker phenotypes.Experimental tests for this model are suggested.

Evolutionary Dynamics of Giant Viruses and their Virophages