Participation costs dismiss the advantage of heterogeneous networks in evolution of cooperation

Real social interactions occur on networks in which each individual is connected to some, but not all, of others. In social dilemma games with a fixed population size, heterogeneity in the number of contacts per player is known to promote evolution of cooperation. Under a common assumption of positively biased payoff structure, well-connected players earn much by playing frequently, and cooperation once adopted by well-connected players is unbeatable and spreads to others. However, maintaining a social contact can be costly, which would prevent local payoffs from being positively biased. In replicator-type evolutionary dynamics, it is shown that even a relatively small participation cost extinguishes the merit of heterogeneous networks in terms of cooperation. In this situation, more connected players earn less so that they are no longer spreaders of cooperation. Instead, those with fewer contacts win and guide the evolution. The participation cost, or the baseline payoff, is irrelevant in homogeneous populations but is essential for evolutionary games on heterogeneous networks.Comment: 4 figures + 3 supplementary figure

Most undirected random graphs are amplifiers of selection for Birth-death dynamics, but suppressors of selection for death-Birth dynamics

We analyze evolutionary dynamics on graphs, where the nodes represent individuals of a population. The links of a node describe which other individuals can be displaced by the offspring of the individual on that node. Amplifiers of selection are graphs for which the fixation probability is increased for advantageous mutants and decreased for disadvantageous mutants. A few examples of such amplifiers have been developed, but so far it is unclear how many such structures exist and how to construct them. Here, we show that almost any undirected random graph is an amplifier of selection for Birth-death updating, where an individual is selected to reproduce with probability proportional to its fitness and one of its neighbors is replaced by that offspring at random. If we instead focus on death-Birth updating, in which a random individual is removed and its neighbors compete for the empty spot, then the same ensemble of graphs consists of almost only suppressors of selection for which the fixation probability is decreased for advantageous mutants and increased for disadvantageous mutants. Thus, the impact of population structure on evolutionary dynamics is a subtle issue that will depend on seemingly minor details of the underlying evolutionary process

Counterintuitive properties of the fixation time in network-structured populations

Evolutionary dynamics on graphs can lead to many interesting and counterintuitive findings. We study the Moran process, a discrete time birth-death process, that describes the invasion of a mutant type into a population of wild-type individuals. Remarkably, the fixation probability of a single mutant is the same on all regular networks. But non-regular networks can increase or decrease the fixation probability. While the time until fixation formally depends on the same transition probabilities as the fixation probabilities, there is no obvious relation between them. For example, an amplifier of selection, which increases the fixation probability and thus decreases the number of mutations needed until one of them is successful, can at the same time slow down the process of fixation. Based on small networks, we show analytically that (i) the time to fixation can decrease when links are removed from the network and (ii) the node providing the best starting conditions in terms of the shortest fixation time depends on the fitness of the mutant. Our results are obtained analytically on small networks, but numerical simulations show that they are qualitatively valid even in much larger populations

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

Evolutionary game theory: Temporal and spatial effects beyond replicator dynamics

Evolutionary game dynamics is one of the most fruitful frameworks for studying evolution in different disciplines, from Biology to Economics. Within this context, the approach of choice for many researchers is the so-called replicator equation, that describes mathematically the idea that those individuals performing better have more offspring and thus their frequency in the population grows. While very many interesting results have been obtained with this equation in the three decades elapsed since it was first proposed, it is important to realize the limits of its applicability. One particularly relevant issue in this respect is that of non-mean-field effects, that may arise from temporal fluctuations or from spatial correlations, both neglected in the replicator equation. This review discusses these temporal and spatial effects focusing on the non-trivial modifications they induce when compared to the outcome of replicator dynamics. Alongside this question, the hypothesis of linearity and its relation to the choice of the rule for strategy update is also analyzed. The discussion is presented in terms of the emergence of cooperation, as one of the current key problems in Biology and in other disciplines.Comment: Review, 48 pages, 26 figure

Spatial Invasion of Cooperation

The evolutionary puzzle of cooperation describes situations where cooperators provide a fitness benefit to other individuals at some cost to themselves. Under Darwinian selection, the evolution of cooperation is a conundrum, whereas non-cooperation (or defection) is not. In the absence of supporting mechanisms, cooperators perform poorly and decrease in abundance. Evolutionary game theory provides a powerful mathematical framework to address the problem of cooperation using the prisoner's dilemma. One well-studied possibility to maintain cooperation is to consider structured populations, where each individual interacts only with a limited subset of the population. This enables cooperators to form clusters such that they are more likely to interact with other cooperators instead of being exploited by defectors. Here we present a detailed analysis of how a few cooperators invade and expand in a world of defectors. If the invasion succeeds, the expansion process takes place in two stages: first, cooperators and defectors quickly establish a local equilibrium and then they uniformly expand in space. The second stage provides good estimates for the global equilibrium frequencies of cooperators and defectors. Under hospitable conditions, cooperators typically form a single, ever growing cluster interspersed with specks of defectors, whereas under more hostile conditions, cooperators form isolated, compact clusters that minimize exploitation by defectors. We provide the first quantitative assessment of the way cooperators arrange in space during invasion and find that the macroscopic properties and the emerging spatial patterns reveal information about the characteristics of the underlying microscopic interactions.MathematicsOrganismic and Evolutionary Biolog