Adaptation and enslavement in endosymbiont-host associations.

The evolutionary persistence of symbiotic associations is a puzzle. Adaptation should eliminate cooperative traits if it is possible to enjoy the advantages of cooperation without reciprocating-a facet of cooperation known in game theory as the Prisoner's Dilemma. Despite this barrier, symbioses are widespread and may have been necessary for the evolution of complex life. The discovery of strategies such as tit-for-tat has been presented as a general solution to the problem of cooperation. However, this only holds for within-species cooperation, where a single strategy will come to dominate the population. In a symbiotic association each species may have a different strategy, and the theoretical analysis of the single-species problem is no guide to the outcome. We present basic analysis of two-species cooperation and show that a species with a fast adaptation rate is enslaved by a slowly evolving one. Paradoxically, the rapidly evolving species becomes highly cooperative, whereas the slowly evolving one gives little in return. This helps understand the occurrence of endosymbioses where the host benefits, but the symbionts appear to gain little from the association.