We consider the classic Moran process modeling the spread of genetic mutations, as extended to structured populations by (Nature, 2005). In this process, individuals are the vertices of a connected graph G. Initially, there is a single mutant vertex, chosen uniformly at random. In each step, a random vertex is selected for reproduction with a probability proportional to its fitness: mutants have fitness r > 1, while non-mutants have fitness 1. The vertex chosen to reproduce places a copy of itself to a uniformly random neighbor in G, replacing the individual that was there. The process ends when the mutation either reaches fixation (i.e., all vertices are mutants), or gets extinct. The principal quantity of interest is the probability with which each of the two outcomes occurs. A problem that has received significant attention recently concerns the existence of families of graphs, called strong amplifiers of selection, for which the fixation probability tends to 1 as the order n of the graph increases, and the existence of strong suppressors of selection, for which this probability tends to 0. For the case of directed graphs, it is known that both strong amplifiers and suppressors exist. For the case of undirected graphs, however, the problem has remained open, and the general belief has been that neither strong amplifiers nor suppressors exist. In this paper we disprove this belief, by providing the first examples of such graphs. The strong amplifier we present has fixation probability 1 − ˜ O(n^ −1 3), and the strong suppressor has fixation probability ˜O(n^−1 4). Both graph constructions are surprisingly simple. We also prove a general upper bound of 1 − ˜Ω(n^−1 3) on the fixation probability of any undirected graph. Hence, our strong amplifier is existentially optimal.
More involved population dynamics have been studied in evolutionary game theory in the context of well-mixed populations @cite_20 @cite_2 , and more recently in populations on graphs @cite_17 @cite_27 . The fitness of an individual is no longer fixed (determined by its type: mutant or non-mutant), but is determined by the expected payoff for the individual when playing a two-player game against a randomly chosen neighbor. In this game mutants play with one strategy and non-mutants with another.
@cite_27 @cite_17 @cite_20 @cite_2
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Evolutionary dynamics have been traditionally studied in the context of homogeneous or spatially extended populations1,2,3,4. Here we generalize population structure by arranging individuals on a graph. Each vertex represents an individual. The weighted edges denote reproductive rates which govern how often individuals place offspring into adjacent vertices. The homogeneous population, described by the Moran process3, is the special case of a fully connected graph with evenly weighted edges. Spatial structures are described by graphs where vertices are connected with their nearest neighbours. We also explore evolution on random and scale-free networks5,6,7. We determine the fixation probability of mutants, and characterize those graphs for which fixation behaviour is identical to that of a homogeneous population7. Furthermore, some graphs act as suppressors and others as amplifiers of selection. It is even possible to find graphs that guarantee the fixation of any advantageous mutant. We also study frequency-dependent selection and show that the outcome of evolutionary games can depend entirely on the structure of the underlying graph. Evolutionary graph theory has many fascinating applications ranging from ecology to multi-cellular organization and economics. Evolutionary dynamics has been traditionally studied in the context of homogeneous populations, mainly described by the Moran process [P. Moran, Random processes in genetics, Proceedings of the Cambridge Philosophical Society 54 (1) (1958) 60-71]. Recently, this approach has been generalized in [E. Lieberman, C. Hauert, M.A. Nowak, Evolutionary dynamics on graphs, Nature 433 (2005) 312-316] by arranging individuals on the nodes of a network (in general, directed). In this setting, the existence of directed arcs enables the simulation of extreme phenomena, where the fixation probability of a randomly placed mutant (i.e., the probability that the offspring of the mutant eventually spread over the whole population) is arbitrarily small or large. On the other hand, undirected networks (i.e., undirected graphs) seem to have a smoother behavior, and thus it is more challenging to find suppressors amplifiers of selection, that is, graphs with smaller greater fixation probability than the complete graph (i.e., the homogeneous population). In this paper we focus on undirected graphs. We present the first class of undirected graphs which act as suppressors of selection, by achieving a fixation probability that is at most one half of that of the complete graph, as the number of vertices increases. Moreover, we provide some generic upper and lower bounds for the fixation probability of general undirected graphs. As our main contribution, we introduce the natural alternative of the model proposed in [E. Lieberman, C. Hauert, M.A. Nowak, Evolutionary dynamics on graphs, Nature 433 (2005) 312-316]. In our new evolutionary model, all individuals interact simultaneously and the result is a compromise between aggressive and non-aggressive individuals. We prove that our new model of mutual influences admits a potential function, which guarantees the convergence of the system for any graph topology and any initial fitness vector of the individuals. Furthermore, we prove fast convergence to the stable state for the case of the complete graph, as well as we provide almost tight bounds on the limit fitness of the individuals. Apart from being important on its own, this new evolutionary model appears to be useful also in the abstract modeling of control mechanisms over invading populations in networks. We demonstrate this by introducing and analyzing two alternative control approaches, for which we bound the time needed to stabilize to the ''healthy'' state of the system. 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. We model evolution according to an asymmetric game as occurring in multiple finite populations, one for each role in the game, and study the effect of subjecting individuals to stochastic strategy mutations. We show that, when these mutations occur sufficiently infrequently, the dynamics over all population states simplify to an ergodic Markov chain over just the pure population states (where each population is monomorphic). This makes calculation of the stationary distribution computationally feasible. The transition probabilities of this embedded Markov chain involve fixation probabilities of mutants in single populations. The asymmetry of the underlying game leads to fixation probabilities that are derived from frequency-independent selection, in contrast to the analogous single-population symmetric-game case (Fudenberg and Imhof, 2006). This frequency independence is useful in that it allows us to employ results from the population genetics literature to calculate the stationary distribution of the evolutionary process, giving sharper, and sometimes even analytic, results. We demonstrate the utility of this approach by applying it to a battle-of-the-sexes game, a Crawford–Sobel signalling game, and the beer-quiche game of Cho and Kreps (1987). This text offers a systematic, rigorous, and unified presentation of evolutionary game theory, covering the core developments of the theory from its inception in biology in the 1970s through recent advances. Evolutionary game theory, which studies the behavior of large populations of strategically interacting agents, is used by economists to make predictions in settings where traditional assumptions about agents' rationality and knowledge may not be justified. Recently, computer scientists, transportation scientists, engineers, and control theorists have also turned to evolutionary game theory, seeking tools for modeling dynamics in multiagent systems. Population Games and Evolutionary Dynamics provides a point of entry into the field for researchers and students in all of these disciplines. The text first considers population games, which provide a simple, powerful model for studying strategic interactions among large numbers of anonymous agents. It then studies the dynamics of behavior in these games. By introducing a general model of myopic strategy revision by individual agents, the text provides foundations for two distinct approaches to aggregate behavior dynamics: the deterministic approach, based on differential equations, and the stochastic approach, based on Markov processes. Key results on local stability, global convergence, stochastic stability, and nonconvergence are developed in detail. Ten substantial appendixes present the mathematical tools needed to work in evolutionary game theory, offering a practical introduction to the methods of dynamic modeling. Accompanying the text are more than 200 color illustrations of the mathematics and theoretical results; many were created using the Dynamo software suite, which is freely available on the author's Web site. Readers are encouraged to use Dynamo to run quick numerical experiments and to create publishable figures for their own research. Individuals of different types, may it be genetic, cultural, or else, with different levels of fitness often compete for reproduction and survival. A fitter type generally has higher chances of disseminating their copies to other individuals. The fixation probability of a single mutant type introduced in a population of wild-type individuals quantifies how likely the mutant type spreads. How much the excess fitness of the mutant type increases its fixation probability, namely, the selection pressure, is important in assessing the impact of the introduced mutant. Previous studies mostly based on undirected and unweighted contact networks of individuals showed that the selection pressure depends on the structure of networks and the rule of reproduction. Real networks underlying ecological and social interactions are usually directed or weighted. Here we examine how the selection pressure is modulated by directionality of interactions under several update rules. Our conclusions are twofold. First, directionality discounts the selection pressure for different networks and update rules. Second, given a network, the update rules in which death events precede reproduction events significantly decrease the selection pressure than the other rules. We study evolutionary game dynamics in finite populations. We analyze an evolutionary process, which we call pairwise comparison, for which we adopt the ubiquitous Fermi distribution function from statistical mechanics. The inverse temperature in this process controls the intensity of selection, leading to a unified framework for evolutionary dynamics at all intensities of selection, from random drift to imitation dynamics. We derive a simple closed formula that determines the feasibility of cooperation in finite populations, whenever cooperation is modeled in terms of any symmetric two-person game. In contrast with previous results, the present formula is valid at all intensities of selection and for any initial condition. We investigate the evolutionary dynamics of cooperators in finite populations, and study the interplay between intensity of selection and the remnants of interior fixed points in infinite populations, as a function of a given initial number of cooperators, showing how this interplay strongly affects the approach to fixation of a given trait in finite populations, leading to counterintuitive results at different intensities of selection. Every form of behavior is shaped by trial and error. Such stepwise adaptation can occur through individual learning or through natural selection, the basis of evolution. Since the work of Maynard Smith and others, it has been realized how game theory can model this process. Evolutionary game theory replaces the static solutions of classical game theory by a dynamical approach centered not on the concept of rational players but on the population dynamics of behavioral programs. In this book the authors investigate the nonlinear dynamics of the self-regulation of social and economic behavior, and of the closely related interactions among species in ecological communities. Replicator equations describe how successful strategies spread and thereby create new conditions that can alter the basis of their success, i.e., to enable us to understand the strategic and genetic foundations of the endless chronicle of invasions and extinctions that punctuate evolution. In short, evolutionary game theory describes when to escalate a conflict, how to elicit cooperation, why to expect a balance of the sexes, and how to understand natural selection in mathematical terms. Comprehensive treatment of ecological and game theoretic dynamics Invasion dynamics and permanence as key concepts Explanation in terms of games of things like competition between species Darwinian dynamics based on mutation and selection form the core of mathematical models for adaptation and coevolution of biological populations. The evolutionary outcome is often not a fitness-maximizing equilibrium but can include oscillations and chaos. For studying frequency-dependent selection, game-theoretic arguments are more appropriate than optimization algorithms. Replicator and adaptive dynamics describe short- and long-term evolution in phenotype space and have found applications ranging from animal behavior and ecology to speciation, macroevolution, and human language. Evolutionary game theory is an essential component of a mathematical and computational approach to biology. Evolutionary graph theory (EGT), studies the ability of a mutant gene to overtake a nite structured population. In this review, we describe the original framework for EGT and the major work that has followed it. This review looks at the calculation of the probability" - the probability of a mutant taking over a population and focuses on game-theoretic applications. We look at varying topics such as alternate evolutionary dynamics, time to xation, special topological cases, and game theoretic results. Throughout the review, we examine several interesting open problems that warrant further research. A new approach to understanding evolution [Val09], namely viewing it through the lens of computation, has already started yielding new insights, e.g., natural selection under sexual reproduction can be interpreted as the Multiplicative Weight Update (MWU) Algorithm in coordination games played among genes [CLPV14]. Using this machinery, we study the role of mutation in changing environments in the presence of sexual reproduction. Following [WVA05], we model changing environments via a Markov chain, with the states representing environments, each with its own fitness matrix. In this setting, we show that in the absence of mutation, the population goes extinct, but in the presence of mutation, the population survives with positive probability. On the way to proving the above theorem, we need to establish some facts about dynamics in games. We provide the first, to our knowledge, polynomial convergence bound for noisy MWU in a coordination game. Finally, we also show that in static environments, sexual evolution with mutation converges, for any level of mutation. In a recent series of papers a surprisingly strong connection was discovered between standard evolutionary models of natural selection and Multiplicative Weights Updates Algorithm, a ubiquitous model of online learning and optimization. These papers establish that, under specific assumptions, mathematical models of biological evolution can be reduced to studying discrete replicator dynamics, a close variant of MWUA, in coordination games. This connection allows for introducing insights from game theoretic dynamics into the field of mathematical biology. Using these results as a stepping stone, we show that mathematical models of haploid evolution imply the extinction of genetic diversity in the long term limit, a widely believed conjecture in genetics. In game theoretic terms we show that in the case of coordination games, under minimal genericity assumptions, discrete replicator dynamics converge to pure Nash equilibria for all but a zero measure of initial conditions. This result holds despite the fact that mixed Nash equilibria can be exponentially (or even uncountably) many, completely dominating in number the set of pure Nash equilibria. Thus, in haploid organisms the long term preservation of genetic diversity needs to be safeguarded by other evolutionary mechanisms such as mutations and speciation. This work extends what is known so far for a basic model of evolutionary antagonism in undirected networks (graphs). More specifically, this work studies the generalized Moran process, as introduced by Lieberman, Hauert, and Nowak [Nature, 433:312-316, 2005], where the individuals of a population reside on the vertices of an undirected connected graph. The initial population has a single mutant of a fitness value r (typically r>1), residing at some vertex v of the graph, while every other vertex is initially occupied by an individual of fitness 1. At every step of this process, an individual (i.e. vertex) is randomly chosen for reproduction with probability proportional to its fitness, and then it places a copy of itself on a random neighbor, thus replacing the individual that was residing there. The main quantity of interest is the fixation probability, i.e. the probability that eventually the whole graph is occupied by descendants of the mutant. In this work we concentrate on the fixation probability when the mutant is initially on a specific vertex v, thus refining the older notion of which studied the fixation probability when the initial mutant is placed at a random vertex. We then aim at finding graphs that have many "strong starts" (or many "weak starts") for the mutant. Thus we introduce a parameterized notion of selective amplifiers (resp. selective suppressors) of evolution. We prove the existence of strong selective amplifiers (i.e. for h(n)=Θ(n) vertices v the fixation probability of v is at least @math for a function c(r) that depends only on r), and the existence of quite strong selective suppressors. Regarding the traditional notion of fixation probability from a random start, we provide strong upper and lower bounds: first we demonstrate the non-existence of "strong universal" amplifiers, and second we prove the Thermal Theorem which states that for any undirected graph, when the mutant starts at vertex v, the fixation probability at least @math . This theorem (which extends the "Isothermal Theorem" of for regular graphs) implies an almost tight lower bound for the usual notion of fixation probability. Our proof techniques are original and are based on new domination arguments which may be of general interest in Markov Processes that are of the general birth-death type. The Moran process, as studied by [10], is a stochastic process modeling the spread of genetic mutations in populations. In this process, agents of a two-type population (i.e. mutants and residents) are associated with the vertices of a graph. Initially, only one vertex chosen uniformly at random (u.a.r.) is a mutant, with fitness (r > 0 ), while all other individuals are residents, with fitness 1. In every step, an individual is chosen with probability proportional to its fitness, and its state (mutant or resident) is passed on to a neighbor which is chosen u.a.r. In this paper, we introduce and study for the first time a generalization of the model of [10] by assuming that different types of individuals perceive the population through different graphs defined on the same vertex set, namely (G_R = (V, E_R) ) for residents and (G_M = (V, E_M) ) for mutants. In this model, we study the fixation probability, namely the probability that eventually only mutants remain in the population, for various pairs of graphs. Evolutionary models on graphs, as an extension of the Moran process, have two major implementations: birth–death (BD) models (or the invasion process) and death–birth (DB) models (or voter models). The isothermal theorem states that the fixation probability of mutants in a large group of graph structures (known as isothermal graphs, which include regular graphs) coincides with that for the mixed population. This result has been proved by (2005 Nature 433, 312–316. (doi:10.1038 nature03204)) in the case of BD processes, where mutants differ from the wild-types by their birth rate (and not by their death rate). In this paper, we discuss to what extent the isothermal theorem can be formulated for DB processes, proving that it only holds for mutants that differ from the wild-type by their death rate (and not by their birth rate). For more general BD and DB processes with arbitrary birth and death rates of mutants, we show that the fixation probabilities of mutants are different from those obtained in the mass-action populations. We focus on spatial lattices and show that the difference between BD and DB processes on one- and two-dimensional lattices is non-small even for large population sizes. We support these results with a generating function approach that can be generalized to arbitrary graph structures. Finally, we discuss several biological applications of the results. Rock is wrapped by paper, paper is cut by scissors and scissors are crushed by rock. This simple game is popular among children and adults to decide on trivial disputes that have no obvious winner, but cyclic dominance is also at the heart of predator–prey interactions, the mating strategy of side-blotched lizards, the overgrowth of marine sessile organisms and competition in microbial populations. Cyclical interactions also emerge spontaneously in evolutionary games entailing volunteering, reward, punishment, and in fact are common when the competing strategies are three or more, regardless of the particularities of the game. Here, we review recent advances on the rock–paper–scissors (RPS) and related evolutionary games, focusing, in particular, on pattern formation, the impact of mobility and the spontaneous emergence of cyclic dominance. We also review mean-field and zero-dimensional RPS models and the application of the complex Ginzburg–Landau equation, and we highlight the importance and usefulness of statistical physics for the successful study of large-scale ecological systems. Directions for future research, related, for example, to dynamical effects of coevolutionary rules and invasion reversals owing to multi-point interactions, are also outlined. We consider a noncooperative interaction among a large population of mobiles that interfere with each other through many local interactions. The first objective of this paper is to extend the evolutionary game framework to allow an arbitrary number of mobiles that are involved in a local interaction. We allow for interactions between mobiles that are not necessarily reciprocal. We study 1) multiple-access control in a slotted Aloha-based wireless network and 2) power control in wideband code-division multiple-access wireless networks. We define and characterize the equilibrium (called evolutionarily stable strategy) for these games and study the influence of wireless channels and pricing on the evolution of dynamics and the equilibrium. We consider the classic Moran process modeling the spread of genetic mutations, as extended to structured populations by (Nature, 2005). In this process, individuals are the vertices of a connected graph G. Initially, there is a single mutant vertex, chosen uniformly at random. In each step, a random vertex is selected for reproduction with a probability proportional to its fitness: mutants have fitness r > 1, while non-mutants have fitness 1. The vertex chosen to reproduce places a copy of itself to a uniformly random neighbor in G, replacing the individual that was there. The process ends when the mutation either reaches fixation (i.e., all vertices are mutants), or gets extinct. The principal quantity of interest is the probability with which each of the two outcomes occurs. A problem that has received significant attention recently concerns the existence of families of graphs, called strong amplifiers of selection, for which the fixation probability tends to 1 as the order n of the graph increases, and the existence of strong suppressors of selection, for which this probability tends to 0. For the case of directed graphs, it is known that both strong amplifiers and suppressors exist. For the case of undirected graphs, however, the problem has remained open, and the general belief has been that neither strong amplifiers nor suppressors exist. In this paper we disprove this belief, by providing the first examples of such graphs. The strong amplifier we present has fixation probability 1 − ˜ O(n^ −1 3), and the strong suppressor has fixation probability ˜O(n^−1 4). Both graph constructions are surprisingly simple. We also prove a general upper bound of 1 − ˜Ω(n^−1 3) on the fixation probability of any undirected graph. Hence, our strong amplifier is existentially optimal. Abstract In this paper we investigate the evolutionary dynamics of strategic behavior in the game of poker by means of data gathered from a large number of real world poker games. We perform this study from an evolutionary game theoretic perspective using two Replicator Dynamics models. First we consider the basic selection model on this data, secondly we use a model which includes both selection and mutation. We investigate the dynamic properties by studying how rational players switch between different strategies under different circumstances, what the basins of attraction of the equilibria look like, and what the stability properties of the attractors are. We illustrate the dynamics using a simplex analysis. Our experimental results confirm existing domain knowledge of the game, namely that certain strategies are clearly inferior while others can be successful given certain game conditions. A finite-population dynamic evolutionary model is presented, which shows that increasing the individual capacity of sending pre-play signals (without any pre-defined meaning), opens a route for cooperation. The population dynamics leads individuals to discriminate between different signals and react accordingly to the signals received. The proportion of time that the population spends in different states can be calculated analytically. We show that increasing the number of different signals benefits cooperative strategies, illustrating how cooperators may take profit from a diverse signaling portfolio to forecast future behaviors and avoid being cheated by defectors.