We examine birth--death processes with state dependent transition probabilities and at least one absorbing boundary. In evolution, this describes selection acting on two different types in a finite population where reproductive events occur successively. If the two types have equal fitness the system performs a random walk. If one type has a fitness advantage it is favored by selection, which introduces a bias (asymmetry) in the transition probabilities. How long does it take until advantageous mutants have invaded and taken over? Surprisingly, we find that the average time of such a process can increase, even if the mutant type always has a fitness advantage. We discuss this finding for the Moran process and develop a simplified model which allows a more intuitive understanding. We show that this effect can occur for weak but non--vanishing bias (selection) in the state dependent transition rates and infer the scaling with system size. We also address the Wright-Fisher model commonly used in population genetics, which shows that this stochastic slowdown is not restricted to birth-death processes.
We study the stochastic dynamics of evolutionary games, and focus on the so-called `stochastic slowdown' effect, previously observed in (Altrock et. al, 2010) for simple evolutionary dynamics. Slowdown here refers to the fact that a beneficial mutation may take longer to fixate than a neutral one. More precisely, the fixation time conditioned on the mutant taking over can show a maximum at intermediate selection strength. We show that this phenomenon is present in the prisoner's dilemma, and also discuss counterintuitive slowdown and speedup in coexistence games. In order to establish the micr…
We consider the lifetimes of metastable states in bistable evolutionary games (coordination games), and examine how they are affected by spatial structure. A semiclassical approximation based on a path integral method is applied to stochastic evolutionary game dynamics with and without spatial structure, and the lifetimes of the metastable states are evaluated. It is shown that the population dependence of the lifetimes is qualitatively different in these two models. Our result indicates that spatial structure can accelerate the transitions between metastable states.
Frequency dependent selection and demographic fluctuations play important roles in evolutionary and ecological processes. Under frequency dependent selection, the average fitness of the population may increase or decrease based on interactions between individuals within the population. This should be reflected in fluctuations of the population size even in constant environments. Here, we propose a stochastic model, which naturally combines these two evolutionary ingredients by assuming frequency dependent competition between different types in an individual-based model. In contrast to previous…
Population structure can have a significant effect on evolution. For some systems with sufficient symmetry, analytic results can be derived within the mathematical framework of evolutionary graph theory which relate to the outcome of the evolutionary process. However, for more complicated heterogeneous structures, computationally intensive methods are required such as individual-based stochastic simulations. By adapting methods from statistical physics, including moment closure techniques, we first show how to derive existing homogenised pair approximation models and the exact neutral drift mo…
Phenotypic plasticity and its evolution may help evolutionary rescue in a novel and stressful environment, especially if environmental novelty reveals cryptic genetic variation that enables the evolution of increased plasticity. However, the environmental stochasticity ubiquitous in natural systems may alter these predictions because high plasticity may amplify phenotype-environment mismatches. Although previous studies have highlighted this potential detrimental effect of plasticity in stochastic environments, they have not investigated how it affects extinction risk in the context of evoluti…
Agent-based stochastic models for finite populations have recently received much attention in the game theory of evolutionary dynamics. Both the ultimate fixation and the pre-fixation transient behavior are important to a full understanding of the dynamics. In this paper, we study the transient dynamics of the well-mixed Moran process through constructing a landscape function. It is shown that the landscape playing a central theoretical "device" that integrates several lines of inquiries: the stable behavior of the replicator dynamics, the long-time fixation, and continuous diffusion approxima…
This manuscript contains nothing new, but synthesizes known results: For the theoretical population geneticist with a probabilistic background, we provide a summary of some key results on stochastic differential equations. For the evolutionary game theorist, we give a new perspective on the derivations of results obtained when using discrete birth-death processes. For the theoretical biologist familiar with deterministic modeling, we outline how to derive and work with stochastic versions of classical ecological and evolutionary processes.
Speciation is of fundamental importance to understanding the huge diversity of life on Earth. In contrast to current phenomenological models, we develop a biophysically motivated approach to study speciation involving the co-evolution of protein binding DNA for two geographically isolated populations. Our results predict that, despite neutral diffusion of hybrids in trait space, smaller populations have a higher rate of speciation, due to sequence entropy poising populations more closely to incompatible regions of phenotype space. A key lesson of this work is that non-trivial contributions of …
The central goal of a dynamical theory of evolution is to abstract the mean evolutionary trajectory in the trait space by considering ecological processes at the level of the individual. In this work, we develop such a theory for a new class of deterministic individual based models describing individual births and deaths, which captures the essential features of standard stochastic individual-based models and become identical with the latter under maximal competition. The key motivation is to derive the canonical equation of adaptive dynamics from this microscopic ecological model, which can b…
Frequency-dependent selection reflects the interaction between different species as they battle for limited resources in their environment. In a stochastic evolutionary game the species relative fitnesses guides the evolutionary dynamics which fluctuate due to random drift. Dependence of species selection advantages on the environment introduces additional possibilities for the evolutionary dynamics. We analyse a simple model in which a random environment allows competing species to coexist for a long time before a fixation of a single species happens. In our analysis we use stability in a lin…
Fixation times in evolutionary games under weak selection
在进化博弈动力学中，繁殖成功率随着进化博弈中的表现而增加。如果策略$A$的表现优于策略$B$，则策略$A$将在人群中传播。在随机动力学下，单个突变体迟早会接管整个种群或灭绝。我们分析了与此过程相关的平均退出时间（或平均固定时间）。我们分析表明，这些时间依赖于一个令人惊讶的简单的方式下弱选择，即强随机性的游戏的回报矩阵：回报差$Δπ$是一个线性函数的$A$个人$i$，$Δπ= u i + v$的数量。无条件平均退出时间仅取决于常数项$v$。假设单个$A$突变体接管群体，则相应的条件平均退出时间仅取决于密度依赖项$u$。我们证明了这一发现的两个常用的微观进化过程。
The Letter presents a novel way to connect random walks, stochastic differential equations, and evolutionary game theory. We introduce a new concept of potential function for discrete-space stochastic systems. It is based on a correspondence between one-dimensional stochastic differential equations and random walks, which may be exact not only in the continuous limit but also in finite-state spaces. Our method is useful for computation of fixation probabilities in discrete stochastic dynamical systems with two absorbing states. We apply it to evolutionary games, formulating two simple and intu…
We study the multi-strategy stochastic evolutionary game with death-birth updating in expanding spatial populations of size $N\to \infty$. The model is a voter model perturbation. For typical populations, we require perturbation strengths satisfying $1/N\ll w\ll 1$. Under appropriate conditions on the space, the limiting density processes of strategy are proven to obey the replicator equation, and the normalized fluctuations converge to a Gaussian process with the Wright-Fisher covariance function in the limiting densities. As an application, we resolve in the positive a conjecture from the bi…
One of the properties that make ecological systems so unique is the range of complex behavioural patterns that can be exhibited by even the simplest communities with only a few species. Much of this complexity is commonly attributed to stochastic factors which have very high-degrees of freedom. Orthodox study of the evolution of these simple networks has generally been limited in its ability to explain complexity, since it restricts evolutionary adaptation to an inertia-free process with few degrees of freedom in which only gradual, moderately complex behaviours are possible. We propose a mode…
The Moran process is one of an basic mathematical structure in the evolutionary game theory. In this work, we introduce the formulation of the path integral approach for evolutionary game theory based on the Moran process. We derive the transition probability by the path integral from the initial state to the final state with updating rule of the Moran process. In this framework, the transition probability is the sum of all the evolutionary paths. The path integral formula of the transition probability maybe expected to be a new mathematical tool to explore the stochastic game evolutionary dyn…
We propose a numerical approach to study the invasion fitness of a mutant and to determine evolutionary singular strategies in evolutionary structured models in which the competitive exclusion principle holds. Our approach is based on a dual representation, which consists of the modelling of the small size mutant population by a stochastic model and the computation of its corresponding deterministic model. The use of the deterministic model greatly facilitates the numerical determination of the feasibility of invasion as well as the convergence-stability of the evolutionary singular strategy. …
Spatial evolutionary games model individuals who are distributed in a spatial domain and update their strategies upon playing a normal form game with their neighbors. We derive integro-differential equations as deterministic approximations of the microscopic updating stochastic processes. This generalizes the known mean-field ordinary differential equations and provide a powerful tool to investigate the spatial effects in populations evolution. The deterministic equations allow to identify many interesting features of the evolution of strategy profiles in a population, such as standing and tra…
Effects of Stochastic Games on Evolutionary Dynamics in Structured Populations
College of Artificial Intelligence, Southwest University（西南大学人工智能学院）；Business College, Southwest University（西南大学商学院）；Faculty of Natural Sciences and Mathematics, University of Maribor（马里博尔大学自然科学与数学学院）；Community Healthcare Center Dr. Adolf Drolc Maribor（马里博尔市阿道夫·德罗尔克博士社区健康中心）；Department of Physics, Kyung Hee University（庆熙大学物理系）；University College, Korea University（韩国大学大学院）；Institute of Technical Physics and Materials Science, Centre for Energy Research（能源研究中心技术物理与材料科学研究所）
This paper proposed a discrete stochastic dynamic programming (SDP) model for sustainable ecosystem (SE) planning of the Loess Plateau in Northwestern, China, and analyzed the ecological resource planning by the evolutionary game model in the decision-making process. The main objective is to explore a new approach of SE planning from a viewpoint of discrete SDP and evolutionary game theory, with a specific application in the area of ecological resource planning such as water management problems. In contrast to previous work, the proposed SDP method focuses on the transition probability matrix …
Trusting in others and reciprocating that trust with trustworthy actions are crucial to successful and prosperous societies. The Trust Game has been widely used to quantitatively study trust and trustworthiness, involving a sequential exchange between an investor and a trustee. The deterministic evolutionary game theory predicts no trust and no trustworthiness whereas the behavioural experiments with the one-shot anonymous Trust Game show that people substantially trust and respond trustworthily. To explain these discrepancies, previous works often turn to additional mechanisms, which are borr…
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…
Evolutionary game dynamics describes the spreading of successful strategies in a population of reproducing individuals. Typically, the microscopic definition of strategy spreading is stochastic, such that the dynamics becomes deterministic only in infinitely large populations. Here, we introduce a new microscopic birth--death process that has a fully deterministic strong selection limit in well--mixed populations of any size. Additionally, under weak selection, from this new process the frequency dependent Moran process is recovered. This makes it a natural extension of the usual evolutionary …
The evolutionary process has been modelled in many ways using both stochastic and deterministic models. We develop an algebraic model of evolution in a population of asexually reproducing organisms in which we represent a stochastic walk in phenotype space, constrained to the edges of an underlying graph representing the genotype, with a time-homogeneous Markov Chain. We show its equivalence to a more standard, explicit stochastic model and show the algebraic model's superiority in computational efficiency. Because of this increase in efficiency, we offer the ability to simulate the evolution …
Deterministic evolutionary game dynamics can lead to stable coexistences of different types. Stochasticity, however, drives the loss of such coexistences. This extinction is usually accompanied by population size fluctuations. We investigate the most probable extinction trajectory under such fluctuations by mapping a stochastic evolutionary model to a problem of classical mechanics using the Wentzel-Kramers-Brillouin (WKB) approximation. Our results show that more abundant types in a coexistence can be more likely to go extinct first well agreed with previous results, and also the distance bet…
How the neutral diversity is affected by selection and adaptation is investigated in an eco-evolutionary framework. In our model, we study a finite population in continuous time, where each individual is characterized by a trait under selection and a completely linked neutral marker. Population dynamics are driven by births and deaths, mutations at birth, and competition between individuals. Trait values influence ecological processes (demographic events, competition), and competition generates selection on trait variation, thus closing the eco-evolutionary feedback loop. The demographic effec…
We consider a population living in a patchy environment that varies stochastically in space and time. The population is composed of two morphs (that is, individuals of the same species with different genotypes). In terms of survival and reproductive success, the associated phenotypes differ only in their habitat selection strategies. We compute invasion rates corresponding to the rates at which the abundance of an initially rare morph increases in the presence of the other morph established at equilibrium. If both morphs have positive invasion rates when rare, then there is an equilibrium dist…
Evolutionary game dynamics in finite populations is typically subject to noise, inducing effects which are not present in deterministic systems, including fixation and extinction. In the first part of this paper we investigate the phenomenon of drift reversal in finite populations, taking into account that drift is a local quantity in strategy space. Secondly, we study a simple imitation dynamics, and show that it can lead to fixation at internal mixed-strategy fixed points even in finite populations. Imitation in infinite populations is adequately described by conventional replicator dynamics…
The study of biological cells in terms of mesoscopic, nonequilibrium, nonlinear, stochastic dynamics of open chemical systems provides a paradigm for other complex, self-organizing systems with ultra-fast stochastic fluctuations, short-time deterministic nonlinear dynamics, and long-time evolutionary behavior with exponentially distributed rare events, discrete jumps among punctuated equilibria, and catastrophe.
Several modifications of the famous mathematical Game of Life are introduced by making Game of Life rules stochastic and mutual influence of cells in their 8-neighborhood on a rectangular lattice spatially non-uniform. Results are reported of experimental investigation of evolutionary dynamics of the introduced models. A number of new phenomena in the evolutionary dynamics of the models and collective behavior of patterns they generate are revealed, described and illustrated: formation of maze-like patterns as fixed points of the models, "self-controlled growth", "eternal life" in a bounded sp…
We present a general framework to describe the evolutionary dynamics of an arbitrary number of types in finite populations based on stochastic differential equations (SDE). For large, but finite populations this allows to include demographic noise without requiring explicit simulations. Instead, the population size only rescales the amplitude of the noise. Moreover, this framework admits the inclusion of mutations between different types, provided that mutation rates, $μ$, are not too small compared to the inverse population size 1/N. This ensures that all types are almost always represented i…
We study the interplay of population growth and evolutionary dynamics using a stochastic model based on birth and death events. In contrast to the common assumption of an independent population size, evolution can be strongly affected by population dynamics in general. Especially for fast reproducing microbes which are subject to selection, both types of dynamics are often closely intertwined. We illustrate this by considering different growth scenarios. Depending on whether microbes die or stop to reproduce (dormancy), qualitatively different behaviors emerge. For cooperating bacteria, a perm…
We are interested in the impact of natural selection in a prey-predator community. We introduce an individual-based model of the community that takes into account both prey and predator phenotypes. Our aim is to understand the phenotypic coevolution of prey and predators. The community evolves as a multi-type birth and death process with mutations. We first consider the infinite particle approximation of the process without mutation. In this limit, the process can be approximated by a system of differential equations. We prove the existence of a unique globally asymptotically stable equilibriu…
Languages and genes are both transmitted from generation to generation, with opportunity for differential reproduction and survivorship of forms. Here we apply a rigorous inference framework, drawn from population genetics, to distinguish between two broad mechanisms of language change: drift and selection. Drift is change that results from stochasticity in transmission and it may occur in the absence of any intrinsic difference between linguistic forms; whereas selection is truly an evolutionary force arising from intrinsic differences -- for example, when one form is preferred by members of …
The observed cooperation on the level of genes, cells, tissues, and individuals has been the object of intense study by evolutionary biologists, mainly because cooperation often flourishes in biological systems in apparent contradiction to the selfish goal of survival inherent in Darwinian evolution. In order to resolve this paradox, evolutionary game theory has focused on the Prisoner's Dilemma (PD), which incorporates the essence of this conflict. Here, we encode strategies for the iterated Prisoner's Dilemma (IPD) in terms of conditional probabilities that represent the response of decision…
In times of plenty expectations rise, just as in times of crisis they fall. This can be mathematically described as a Win-Stay-Lose-Shift strategy with dynamic aspiration levels, where individuals aspire to be as wealthy as their average neighbor. Here we investigate this model in the realm of evolutionary social dilemmas on the square lattice and scale-free networks. By using the master equation and Monte Carlo simulations, we find that cooperators coexist with defectors in the whole phase diagram, even at high temptations to defect. We study the microscopic mechanism that is responsible for …
Random walks on multidimensional nonlinear landscapes are of interest in many areas of science and engineering. In particular, properties of adaptive trajectories on fitness landscapes determine population fates and thus play a central role in evolutionary theory. The topography of fitness landscapes and its effect on evolutionary dynamics have been extensively studied in the literature. We will survey the current research knowledge in this field, focusing on a recently developed systematic approach to characterizing path lengths, mean first-passage times, and other statistics of the path ense…
Evolutionary dynamics in finite populations is known to fixate eventually in the absence of mutation. We here show that a similar phenomenon can be found in stochastic game dynamical batch learning, and investigate fixation in learning processes in a simple 2x2 game, for two-player games with cyclic interaction, and in the context of the best-shot network game. The analogues of finite populations in evolution are here finite batches of observations between strategy updates. We study when and how such fixation can occur, and present results on the average time-to-fixation from numerical simulat…
We consider a stochastic model for the evolution of a discrete population structured by a trait with values on a finite grid of the torus, and with mutation and selection. Traits are vertically inherited unless a mutation occurs, and influence the birth and death rates. We focus on a parameter scaling where population is large, individual mutations are small but not rare, and the grid mesh for the trait values is much smaller than the size of mutation steps. When considering the evolution of the population in a long time scale, the contribution of small sub-populations may strongly influence t…
