Environmental feedback maintains cooperation in phage $\Phi_6$

The evolution and maintenance of cooperation is a fundamental problem in evolutionary biology. Because cooperative behaviors impose a cost, Cooperators are vulnerable to exploitation by Defectors that do not pay the cost to cooperate but still benefit from the cooperation of others. The bacteriophage $\Phi_6$ exhibits cooperative and defective phenotypes in infection: during replication, phages produce essential proteins in the host cell cytoplasm. Coinfection between multiple phages is possible. A given phage cannot guarantee exclusive access to its own proteins, so Cooperators contribute to the common pool of proteins while Defectors contribute less and instead appropriate proteins from Cooperators. Previous experimental work found that $\Phi_6$ was trapped in a prisoner's dilemma, predicting that the cooperative phenotype should disappear. Here we propose that environmental feedback, or interplay between phage and host densities, can maintain cooperation in $\Phi_6$ populations by modulating the rate of co-infection and shifting the advantages of cooperation vs. defection. We build and analyze an ODE model and find that for a wide range of parameter values, environmental feedback allows Cooperation to survive.