Coevolutionary games - a mini review

Prevalence of cooperation within groups of selfish individuals is puzzling in that it contradicts with the basic premise of natural selection. Favoring players with higher fitness, the latter is key for understanding the challenges faced by cooperators when competing with defectors. Evolutionary game theory provides a competent theoretical framework for addressing the subtleties of cooperation in such situations, which are known as social dilemmas. Recent advances point towards the fact that the evolution of strategies alone may be insufficient to fully exploit the benefits offered by cooperative behavior. Indeed, while spatial structure and heterogeneity, for example, have been recognized as potent promoters of cooperation, coevolutionary rules can extend the potentials of such entities further, and even more importantly, lead to the understanding of their emergence. The introduction of coevolutionary rules to evolutionary games implies, that besides the evolution of strategies, another property may simultaneously be subject to evolution as well. Coevolutionary rules may affect the interaction network, the reproduction capability of players, their reputation, mobility or age. Here we review recent works on evolutionary games incorporating coevolutionary rules, as well as give a didactic description of potential pitfalls and misconceptions associated with the subject. In addition, we briefly outline directions for future research that we feel are promising, thereby particularly focusing on dynamical effects of coevolutionary rules on the evolution of cooperation, which are still widely open to research and thus hold promise of exciting new discoveries.Comment: 24 two-column pages, 10 figures; accepted for publication in BioSystem

Red Queen Coevolution on Fitness Landscapes

Species do not merely evolve, they also coevolve with other organisms. Coevolution is a major force driving interacting species to continuously evolve ex- ploring their fitness landscapes. Coevolution involves the coupling of species fit- ness landscapes, linking species genetic changes with their inter-specific ecological interactions. Here we first introduce the Red Queen hypothesis of evolution com- menting on some theoretical aspects and empirical evidences. As an introduction to the fitness landscape concept, we review key issues on evolution on simple and rugged fitness landscapes. Then we present key modeling examples of coevolution on different fitness landscapes at different scales, from RNA viruses to complex ecosystems and macroevolution.Comment: 40 pages, 12 figures. To appear in "Recent Advances in the Theory and Application of Fitness Landscapes" (H. Richter and A. Engelbrecht, eds.). Springer Series in Emergence, Complexity, and Computation, 201

Mutualism and evolutionary multiplayer games: revisiting the Red King

Coevolution of two species is typically thought to favour the evolution of faster evolutionary rates helping a species keep ahead in the Red Queen race, where `it takes all the running you can do to stay where you are'. In contrast, if species are in a mutualistic relationship, it was proposed that the Red King effect may act, where it can be beneficial to evolve slower than the mutualistic species. The Red King hypothesis proposes that the species which evolves slower can gain a larger share of the benefits. However, the interactions between the two species may involve multiple individuals. To analyse such a situation, we resort to evolutionary multiplayer games. Even in situations where evolving slower is beneficial in a two-player setting, faster evolution may be favoured in a multiplayer setting. The underlying features of multiplayer games can be crucial for the distribution of benefits. They also suggest a link between the evolution of the rate of evolution and group size

Conformity enhances network reciprocity in evolutionary social dilemmas

The pursuit of highest payoffs in evolutionary social dilemmas is risky and sometimes inferior to conformity. Choosing the most common strategy within the interaction range is safer because it ensures that the payoff of an individual will not be much lower than average. Herding instincts and crowd behavior in humans and social animals also compel to conformity on their own right. Motivated by these facts, we here study the impact of conformity on the evolution of cooperation in social dilemmas. We show that an appropriate fraction of conformists within the population introduces an effective surface tension around cooperative clusters and ensures smooth interfaces between different strategy domains. Payoff-driven players brake the symmetry in favor of cooperation and enable an expansion of clusters past the boundaries imposed by traditional network reciprocity. This mechanism works even under the most testing conditions, and it is robust against variations of the interaction network as long as degree-normalized payoffs are applied. Conformity may thus be beneficial for the resolution of social dilemmas.Comment: 8 two-column pages, 5 figures; accepted for publication in Journal of the Royal Society Interfac

Coevolution of Cooperation and Partner Rewiring Range in Spatial Social Networks

In recent years, there has been growing interest in the study of coevolutionary games on networks. Despite much progress, little attention has been paid to spatially embedded networks, where the underlying geographic distance, rather than the graph distance, is an important and relevant aspect of the partner rewiring process. It thus remains largely unclear how individual partner rewiring range preference, local vs. global, emerges and affects cooperation. Here we explicitly address this issue using a coevolutionary model of cooperation and partner rewiring range preference in spatially embedded social networks. In contrast to local rewiring, global rewiring has no distance restriction but incurs a one-time cost upon establishing any long range link. We find that under a wide range of model parameters, global partner switching preference can coevolve with cooperation. Moreover, the resulting partner network is highly degree-heterogeneous with small average shortest path length while maintaining high clustering, thereby possessing small-world properties. We also discover an optimum availability of reputation information for the emergence of global cooperators, who form distant partnerships at a cost to themselves. From the coevolutionary perspective, our work may help explain the ubiquity of small-world topologies arising alongside cooperation in the real world

How the other half lives: CRISPR-Cas's influence on bacteriophages

CRISPR-Cas is a genetic adaptive immune system unique to prokaryotic cells used to combat phage and plasmid threats. The host cell adapts by incorporating DNA sequences from invading phages or plasmids into its CRISPR locus as spacers. These spacers are expressed as mobile surveillance RNAs that direct CRISPR-associated (Cas) proteins to protect against subsequent attack by the same phages or plasmids. The threat from mobile genetic elements inevitably shapes the CRISPR loci of archaea and bacteria, and simultaneously the CRISPR-Cas immune system drives evolution of these invaders. Here we highlight our recent work, as well as that of others, that seeks to understand phage mechanisms of CRISPR-Cas evasion and conditions for population coexistence of phages with CRISPR-protected prokaryotes.Comment: 24 pages, 8 figure

Extending the Modern Synthesis: The evolution of ecosystems

The Modern Evolutionary Synthesis formalizes the role of variation, heredity, differential reproduction and mutation in population genetics. Here we explore a mathematical structure, based on the asymptotic limit theorems of information theory, that instantiates the punctuated dynamic relations of organisms and their embedding environments. The mathematical overhead is considerable, and we conclude that the model must itself be extended even further to allow the possibility of the transfer of heritage information between different classes of organisms. In essence, we provide something of a formal roadmap for the modernization of the Modern Synthesis

Leaders should not be conformists in evolutionary social dilemmas

The most common assumption in evolutionary game theory is that players should adopt a strategy that warrants the highest payoff. However, recent studies indicate that the spatial selection for cooperation is enhanced if an appropriate fraction of the population chooses the most common rather than the most profitable strategy within the interaction range. Such conformity might be due to herding instincts or crowd behavior in humans and social animals. In a heterogeneous population where individuals differ in their degree, collective influence, or other traits, an unanswered question remains who should conform. Selecting conformists randomly is the simplest choice, but it is neither a realistic nor the optimal one. We show that, regardless of the source of heterogeneity and game parametrization, socially the most favorable outcomes emerge if the masses conform. On the other hand, forcing leaders to conform significantly hinders the constructive interplay between heterogeneity and coordination, leading to evolutionary outcomes that are worse still than if conformists were chosen randomly. We conclude that leaders must be able to create a following for network reciprocity to be optimally augmented by conformity. In the opposite case, when leaders are castrated and made to follow, the failure of coordination impairs the evolution of cooperation.Comment: 7 two-column pages, 4 figures; accepted for publication in Scientific Reports [related work available at arXiv:1412.4113

A SLOWLY EVOLVING HOST MOVES FIRST IN SYMBIOTIC INTERACTIONS

Symbiotic relationships, both parasitic and mutualistic, are ubiquitous in nature. Understanding how these symbioses evolve, from bacteria and their phages to humans and our gut microflora, is crucial in understanding how life operates. Often, symbioses consist of a slowly evolving host species with each host only interacting with its own subpopulation of symbionts. The Red Queen hypothesis describes coevolutionary relationships as constant arms races with each species rushing to evolve an advantage over the other, suggesting that faster evolution is favored. Here, we use a simple game theoretic model of host–symbiont coevolution that includes population structure to show that if the symbionts evolve much faster than the host, the equilibrium distribution is the same as it would be if it were a sequential game where the host moves first against its symbionts. For the slowly evolving host, this will prove to be advantageous in mutualisms and a handicap in antagonisms. The result follows from rapid symbiont adaptation to its host and is robust to changes in the parameters, even generalizing to continuous and multiplayer games. Our findings provide insight into a wide range of symbiotic phenomena and help to unify the field of coevolutionary theory.National Institutes of Health (U.S.) (K99 Pathways to Independence Award