RNA Bacteriophage Φ6
Evolution of sex and its consequences
Game theory and virus interactions
Evolutionary ecology of host shifts
Evolution of sex and its consequences
We use the RNA bacteriophage Φ6 as a model to study the costs
and benefits of genetic exchange (sex). When multiple viruses co-infect
the same host cell, sex produces hybrid progeny containing a mixture
of genetic information from the co-infecting parents (Fig. 1).

Figure 1: Colorized electron microscope image
of herpesviruses (yellow and green spheres) coinfecting a
single cell. During coinfection, sexual reproduction can produce
viral progeny that contain a mixture of genes found in the
coinfecting parents.
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Sex may benefit viruses by promoting genetic variation, which might
allow sexual populations to evolve faster than asexual ones. In
contrast, sex requires co-infection and may exert a cost because
it increases competition between viruses for limited intracellular
resources.
One experiment allowed replicate populations of Φ6 to evolve
in the presence and absence of sex for hundreds of viral generations
(Turner and Chao 1998, Genetics 150:523-532). Unexpectedly, sex
was costly because sexually-evolved viruses became attenuated (weakened)
in their ability to infect the host alone. This indicates that the
cost of intra-host competition can outweigh any of the potential
benefits associated with sex.
Ongoing projects examine whether sex is advantageous in purging
epistatic (interacting) mutations from the virus genome, whether
sex is costly in terms of breaking apart co-adapted loci, and how
reproductive system (sexuality versus asexuality) influences the
rate of molecular evolution and prevalence for epistasis to evolve.
Game theory and virus interactions
“Name me somebody that is not a parasite, and I’ll
go out and say a
prayer for him” – Bob Dylan (Visions of Johanna, 1966)
The manufacture of diffusible, and hence shared, intracellular
products during virus co-infection allows for the conflicting strategies
of cooperation and defection (selfishness). Whereas a viral genotype
that synthesizes larger quantities of product is effectively a cooperator,
a genotype that synthesizes less but specializes in sequestering
a larger share of the products is a defector.

Figure 2: Expected fitness values for a game
in which opponents utilize conflicting strategies of cooperation
and defection. Entries in the payoff matrix represent the
fitness to an individual adopting the strategy on the left,
if the opponent adopts the strategy above. Defectors gain
a fitness advantage (1 + s2) that allows them to invade a
population of cooperators. If the cost of defection is too
strong, (1 ? c) < (1 ? s1), cooperators may also invade
and the two strategies are driven to a stable polymorphism.
Prisoner’s dilemma occurs if it always pays to be selfish,
(1 ? c) > (1 ? s1); defection sweeps through the population
despite the greater fitness payoff had all individuals cooperated.
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We use phage Φ6 to study the evolution of viral conflicts.
One study showed that viruses cultured under high levels of co-infection
evolve selfish strategies, but their fixation in the population
causes mean fitness to decline (Turner and Chao 1999, Nature 398:441-443).
These data conform to the prisoner’s dilemma of game theory
(Fig. 2), where selfishness evolves despite the greater fitness
payoff if all players cooperate.
Ongoing projects examine the generality of the prisoner’s
dilemma result in Φ6; for instance, evolved cooperator viruses
can re-invade populations of defectors, allowing the two strategies
to coexist in a stable polymorphism (Turner and Chao 2003, American
Naturalist 161:497-505). Current research examines the molecular
mechanism(s) that may afford selfish genotypes an advantage during
intra-host competition. We are also examining whether viruses evolve
specific mechanisms to exclude large numbers of viruses from co-infecting
the same cell, in order to reduce intracellular competition (Turner
et al. 1999, Journal of Virology 73:2420-2424).
Evolutionary ecology of host shifts
A virus’s ecological niche is governed in part by its host
range, the hosts in which a virus can produce viable progeny. Viruses
may expand their host range through mutations that facilitate entry
into new host environments. Although host shifts allow a virus (or
any other parasite) to expand its ecological niche, traits governing
the infection of multiple host types can decrease fitness in the
original or alternate host environments.
We use phage Φ6 and Pseudomonas bacteria as a model to test basic
questions relating to evolutionary ecology of host shifts. Ongoing
projects include measuring the mutation spectrum associated with
expanded host range, growth tradeoffs for host range mutants across
host environments, and strength of selection resulting from simultaneous
adaptation of viruses to multiple habitats.
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