The origin of synergistic symbiosis.

A dominant theme in the history of life has been the evolutionary innovations of cooperative symbioses: the first genomes near the origin of life, integrated prokaryotic cells, the complex symbiotic communities that evolved into modern eukaryotic cells, lichens, mycorrhizae, and so on. In this paper, a model of cooperative symbiosis that shows a threshold condition for the evolution of cooperation is analyzed. The threshold is not easily passed, but cooperative evolution proceeds rapidly once a symbiosis overcomes the threshold. In the model presented here, each species has genetic variability for a symbiotic trait. The trait imposes a reproductive cost on its bearer but enhances the reproduction of its partner species. For example, in the origin of genetic systems, the trait may cause biochemical synergism for the rate of replication of primitive RNA strands as in Eigen and Schuster's hypercycle model. Models of growth are contrasted with synergism, which are most appropriate for the evolution of genetic systems and for mutualisms such as lichens, with the strategic and psychological applications of the Prisoner's Dilemma model.