  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.
