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The evolution of coercive policing in genetically mixed groups: the case of plasmid copy number control

Policing is a widespread mechanism regulating cooperation in both human and animal social groups. Policing can promote the evolution and maintenance of cooperation among non-relatives by tying the reproductive success of individuals to the efficiency and success of the group. In this paper, we investigate the evolution of reproductive policing using a multi-scale computational model inspired by the copy number control system of conjugative bacterial plasmids. Our results show that the repression of competition through policing can evolve across a very broad range of migration (plasmid conjugation) rates, improving system-level performance and bringing efficiency gains to the group beyond those achievable by pure self-restraint. Reproductive policing acts to increase genetic relatedness by reducing variation in group size which, in turn, reduces the heterogeneity of the plasmid population. When among-group migration is high, coercive policing strategies are favoured, characterized by high levels of policing coupled with relatively lower obedience. Coercive policing strategies preferentially limit the reproduction of rival lineages while, at the same time, maintaining effective collective reproductive control. Author Summary The emergence and maintenance of cooperation is a topic of great importance in evolutionary biology. The evolution of cooperation has been explained in the context of kin selection when there is sufficient genetic relatedness among interacting individuals. When there is insufficient relatedness, the presence of alternative mechanisms, such as mutual policing, can promote the evolution and maintenance of cooperation by tying the reproductive success of individual to the efficiency and success of the group. In this paper, we investigate the evolution of reproductive policing using an agent-based computational model inspired by a simple and elegant biological example: replication control among conjugative plasmids, a class of molecular symbionts of bacterial hosts. Our results show that the repression of competition through policing evolves and improves plasmid group performance beyond levels achievable by self-restraint, across a very broad range of migration rates. Under conditions of high migration (frequent conjugation), we observe the evolution of coercive policing strategies that limit the reproduction of rival lineages by investing disproportionately in policing relative to their obedience to the policing trait.