The Stability of Cooperation Involving Variable Investment

In this paper, we attempt to reconcile the results of several studies which have investigated the evolution of cooperation between non-relatives in systems where investment in partners can vary. In contrast to previous proposals, we show for the first time that variable-investment cooperation can be readily maintained in inter-species mutualistic relationships even in the absence of spatial structure, but that the stability of this interaction is dependent on the particular investment–response rule that is employed. By allowing the evolution of investment–response parameters in both inter- and intra-specific versions of the continuous variable-investment Prisonersʹ Dilemma we show that, in the absence of further factors, the raise-the stakes (RTS) strategy is likely to evolve into a simpler, variable investment form of Give-as-Good-as-you-Get that initially offers a high fixed amount and subsequently matches its partnerʹs investment. Nevertheless, we demonstrate that genuine RTS-like strategies will still be selected if two intuitively reasonable conditions hold: if individuals are limited in terms of the total they can invest in cooperative actions over their lifetimes, and if there are always some individuals in a population that cannot cooperate.