Metastability and anomalous fixation in evolutionary games on scale-free networks

We study the influence of complex graphs on the metastability and fixation properties of a set of evolutionary processes. In the framework of evolutionary game theory, where the fitness and selection are frequency-dependent and vary with the population composition, we analyze the dynamics of snowdrift games (characterized by a metastable coexistence state) on scale-free networks. Using an effective diffusion theory in the weak selection limit, we demonstrate how the scale-free structure affects the system's metastable state and leads to anomalous fixation. In particular, we analytically and numerically show that the probability and mean time of fixation are characterized by stretched exponential behaviors with exponents depending on the network's degree distribution.Comment: 5 pages, 4 figures, to appear in Physical Review Letter

Temporal and dimensional effects in evolutionary graph theory

The spread in time of a mutation through a population is studied analytically and computationally in fully-connected networks and on spatial lattices. The time, t_*, for a favourable mutation to dominate scales with population size N as N^{(D+1)/D} in D-dimensional hypercubic lattices and as N ln N in fully-connected graphs. It is shown that the surface of the interface between mutants and non-mutants is crucial in predicting the dynamics of the system. Network topology has a significant effect on the equilibrium fitness of a simple population model incorporating multiple mutations and sexual reproduction. Includes supplementary information.Comment: 6 pages, 4 figures Replaced after final round of peer revie

Preservation of information in a prebiotic package model

The coexistence between different informational molecules has been the preferred mode to circumvent the limitation posed by imperfect replication on the amount of information stored by each of these molecules. Here we reexamine a classic package model in which distinct information carriers or templates are forced to coexist within vesicles, which in turn can proliferate freely through binary division. The combined dynamics of vesicles and templates is described by a multitype branching process which allows us to write equations for the average number of the different types of vesicles as well as for their extinction probabilities. The threshold phenomenon associated to the extinction of the vesicle population is studied quantitatively using finite-size scaling techniques. We conclude that the resultant coexistence is too frail in the presence of parasites and so confinement of templates in vesicles without an explicit mechanism of cooperation does not resolve the information crisis of prebiotic evolution.Comment: 9 pages, 8 figures, accepted version, to be published in PR

Evolutionary dynamics on degree-heterogeneous graphs

The evolution of two species with different fitness is investigated on degree-heterogeneous graphs. The population evolves either by one individual dying and being replaced by the offspring of a random neighbor (voter model (VM) dynamics) or by an individual giving birth to an offspring that takes over a random neighbor node (invasion process (IP) dynamics). The fixation probability for one species to take over a population of N individuals depends crucially on the dynamics and on the local environment. Starting with a single fitter mutant at a node of degree k, the fixation probability is proportional to k for VM dynamics and to 1/k for IP dynamics.Comment: 4 pages, 4 figures, 2 column revtex4 format. Revisions in response to referee comments for publication in PRL. The version on arxiv.org has one more figure than the published PR

Nongaussian fluctuations arising from finite populations: Exact results for the evolutionary Moran process

The appropriate description of fluctuations within the framework of evolutionary game theory is a fundamental unsolved problem in the case of finite populations. The Moran process recently introduced into this context [Nowak et al., Nature (London) 428, 646 (2004)] defines a promising standard model of evolutionary game theory in finite populations for which analytical results are accessible. In this paper, we derive the stationary distribution of the Moran process population dynamics for arbitrary $2\times{}2$ games for the finite size case. We show that a nonvanishing background fitness can be transformed to the vanishing case by rescaling the payoff matrix. In contrast to the common approach to mimic finite-size fluctuations by Gaussian distributed noise, the finite size fluctuations can deviate significantly from a Gaussian distribution.Comment: 4 pages (2 figs). Published in Physical Review E (Rapid Communications

Stochasticity and evolutionary stability

In stochastic dynamical systems, different concepts of stability can be obtained in different limits. A particularly interesting example is evolutionary game theory, which is traditionally based on infinite populations, where strict Nash equilibria correspond to stable fixed points that are always evolutionarily stable. However, in finite populations stochastic effects can drive the system away from strict Nash equilibria, which gives rise to a new concept for evolutionary stability. The conventional and the new stability concepts may apparently contradict each other leading to conflicting predictions in large yet finite populations. We show that the two concepts can be derived from the frequency dependent Moran process in different limits. Our results help to determine the appropriate stability concept in large finite populations. The general validity of our findings is demonstrated showing that the same results are valid employing vastly different co-evolutionary processes

Universality of weak selection

Weak selection, which means a phenotype is slightly advantageous over another, is an important limiting case in evolutionary biology. Recently it has been introduced into evolutionary game theory. In evolutionary game dynamics, the probability to be imitated or to reproduce depends on the performance in a game. The influence of the game on the stochastic dynamics in finite populations is governed by the intensity of selection. In many models of both unstructured and structured populations, a key assumption allowing analytical calculations is weak selection, which means that all individuals perform approximately equally well. In the weak selection limit many different microscopic evolutionary models have the same or similar properties. How universal is weak selection for those microscopic evolutionary processes? We answer this question by investigating the fixation probability and the average fixation time not only up to linear, but also up to higher orders in selection intensity. We find universal higher order expansions, which allow a rescaling of the selection intensity. With this, we can identify specific models which violate (linear) weak selection results, such as the one--third rule of coordination games in finite but large populations.Comment: 12 pages, 3 figures, accepted for publication in Physical Review

The statistical mechanics of a polygenic characterunder stabilizing selection, mutation and drift

By exploiting an analogy between population genetics and statistical mechanics, we study the evolution of a polygenic trait under stabilizing selection, mutation, and genetic drift. This requires us to track only four macroscopic variables, instead of the distribution of all the allele frequencies that influence the trait. These macroscopic variables are the expectations of: the trait mean and its square, the genetic variance, and of a measure of heterozygosity, and are derived from a generating function that is in turn derived by maximizing an entropy measure. These four macroscopics are enough to accurately describe the dynamics of the trait mean and of its genetic variance (and in principle of any other quantity). Unlike previous approaches that were based on an infinite series of moments or cumulants, which had to be truncated arbitrarily, our calculations provide a well-defined approximation procedure. We apply the framework to abrupt and gradual changes in the optimum, as well as to changes in the strength of stabilizing selection. Our approximations are surprisingly accurate, even for systems with as few as 5 loci. We find that when the effects of drift are included, the expected genetic variance is hardly altered by directional selection, even though it fluctuates in any particular instance. We also find hysteresis, showing that even after averaging over the microscopic variables, the macroscopic trajectories retain a memory of the underlying genetic states.Comment: 35 pages, 8 figure

Shift in critical temperature for random spatial permutations with cycle weights

We examine a phase transition in a model of random spatial permutations which originates in a study of the interacting Bose gas. Permutations are weighted according to point positions; the low-temperature onset of the appearance of arbitrarily long cycles is connected to the phase transition of Bose-Einstein condensates. In our simplified model, point positions are held fixed on the fully occupied cubic lattice and interactions are expressed as Ewens-type weights on cycle lengths of permutations. The critical temperature of the transition to long cycles depends on an interaction-strength parameter $\alpha$. For weak interactions, the shift in critical temperature is expected to be linear in $\alpha$ with constant of linearity $c$. Using Markov chain Monte Carlo methods and finite-size scaling, we find $c = 0.618 \pm 0.086$. This finding matches a similar analytical result of Ueltschi and Betz. We also examine the mean longest cycle length as a fraction of the number of sites in long cycles, recovering an earlier result of Shepp and Lloyd for non-spatial permutations.Comment: v2 incorporated reviewer comments. v3 removed two extraneous figures which appeared at the end of the PDF

Evolutionary game theory in growing populations

Existing theoretical models of evolution focus on the relative fitness advantages of different mutants in a population while the dynamic behavior of the population size is mostly left unconsidered. We here present a generic stochastic model which combines the growth dynamics of the population and its internal evolution. Our model thereby accounts for the fact that both evolutionary and growth dynamics are based on individual reproduction events and hence are highly coupled and stochastic in nature. We exemplify our approach by studying the dilemma of cooperation in growing populations and show that genuinely stochastic events can ease the dilemma by leading to a transient but robust increase in cooperationComment: 4 pages, 2 figures and 2 pages supplementary informatio