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).
In order to fulfill cell proliferation and differentiation through cellular hierarchy, stem cells can undergo either asymmetric or symmetric divisions. Recent studies pay special attention to the effect of different modes of stem cell division on the lifetime risk of cancer, and report that symmetric division is more beneficial to delay the onset of cancer. The fate uncertainty of symmetric division is considered to be the reason for the cancer-delaying effect. In this paper we compare asymmetric and symmetric divisions of stem cells via studying stochastic stem cell models with mutations. Spe…
Although mutations drive the evolutionary process, the rates at which the mutations occur are themselves subject to evolutionary forces. Our purpose here is to understand the role of selection and random genetic drift in the evolution of mutation rates, and we address this question in asexual populations at mutation-selection equilibrium neglecting selective sweeps. Using a multitype branching process, we calculate the fixation probability of a rare nonmutator in a large asexual population of mutators, and find that a nonmutator is more likely to fix when the deleterious mutation rate of the m…
Muller's ratchet describes the irreversible accumulation of deleterious mutations in asexual populations. In well-mixed populations the speed of fitness decline is exponentially small in the population size, and any positive rate of beneficial mutations is sufficient to reverse the ratchet in large populations. The behavior is fundamentally different in populations with spatial structure, because the speed of the ratchet remains nonzero in the infinite size limit when the deleterious mutation rate exceeds a critical value. Based on the relation between the spatial ratchet and directed percolat…
A non-local rare mutations model for quasispecies and Prisoner's dilemma: numerical assessment of qualitative behaviour
Università di Napoli “Parthenope”（那不勒斯帕尔特诺佩大学）；Istituto per le Applicazioni del Calcolo “M. Picone”, Consiglio Nazionale delle Ricerche（意大利国家研究委员会计算应用研究所）
We propose a stochastic model in evolutionary game theory where individuals (or subpopulations) can mutate changing their strategies randomly (but rarely) and explore the external environment. This environment affects the selective pressure by modifying the payoff arising from the interactions between strategies. We derive a Fokker-Plank integro-differential equation and provide Monte-Carlo simulations for the Hawks vs Doves game. In particular we show that, in some cases, taking into account the external environment favors the persistence of the low-fitness strategy.
In an adapted population of mutators in which most mutations are deleterious, a nonmutator that lowers the mutation rate is under indirect selection and can sweep to fixation. Using a multitype branching process, we calculate the fixation probability of a rare nonmutator in a large population of asexual mutators. We show that when beneficial mutations are absent, the fixation probability is a nonmonotonic function of the mutation rate of the mutator: it first increases sublinearly and then decreases exponentially. We also find that beneficial mutations can enhance the fixation probability of a…
In order to accommodate the empirical fact that population structures are rarely simple, modern studies of evolutionary dynamics allow for complicated and highly-heterogeneous spatial structures. As a result, one of the most difficult obstacles lies in making analytical deductions, either qualitative or quantitative, about the long-term outcomes of evolution. The "structure-coefficient" theorem is a well-known approach to this problem for mutation-selection processes under weak selection, but a general method of evaluating the terms it comprises is lacking. Here, we provide such a method for p…
The emergence of a predominant phenotype within a cell population is often triggered by a rare accumulation of DNA mutations in a single cell. For example, tumors may be initiated by a single cell in which multiple mutations cooperate to bypass a cell's defense mechanisms. The risk of such an event is thus determined by the extremal accumulation of mutations across tissue cells. To address this risk, we study the statistics of the maximum mutation numbers in a generic, but tested, model of a renewing cell population. By drawing an analogy between the genealogy of a cell population and the theo…
We present a model for the dynamics of a population of bacteria with a continuum of traits, who compete for resources and exchange horizontally (transfer) an otherwise vertically inherited trait with possible mutations. Competition influences individual demographics, affecting population size, which feeds back on the dynamics of transfer. We consider a stochastic individual-based pure jump process taking values in the space of point measures, and whose jump events describe the individual reproduction, transfer and death mechanisms. In a large population scale, the stochastic process is proved …
Evolutionarily stable strategy (ESS) is an important solution concept in game theory which has been applied frequently to biological models. Informally an ESS is a strategy that if followed by the population cannot be taken over by a mutation strategy that is initially rare. Finding such a strategy has been shown to be difficult from a theoretical complexity perspective. We present an algorithm for the case where mutations are restricted to pure strategies, and present experiments on several game classes including random and a recently-proposed cancer model. Our algorithm is based on a mixed-i…
We consider a stochastic model of population dynamics where each individual is characterised by a trait in {0,1,...,L} and has a natural reproduction rate, a logistic death rate due to age or competition and a probability of mutation towards neighbouring traits at each reproduction event. We choose parameters such that the induced fitness landscape exhibits a valley: mutant individuals with negative fitness have to be created in order for the population to reach a trait with positive fitness. We focus on the limit of large population and rare mutations at several speeds. In particular, when th…
Punctuated equilibrium is a mode of evolution in which phenetic change occurs in rapid bursts that are separated by much longer intervals of stasis during which mutations accumulate but no major phenotypic change occurs. Punctuated equilibrium has been originally proposed within the framework of paleobiology, to explain the lack of transitional forms that is typical of the fossil record. Theoretically, punctuated equilibrium has been linked to self-organized criticality (SOC), a model in which the size of avalanches in an evolving system is power-law distributed, resulting in increasing rarity…
In large clonal populations, several clones generally compete which results in complex evolutionary and ecological dynamics: experiments show successive selective sweeps of favorable mutations as well as long-term coexistence of multiple clonal strains. The mechanisms underlying either coexistence or fixation of several competing strains have rarely been studied altogether. Conditions for coexistence have mostly been studied by population and community ecology, while rates of invasion and fixation have mostly been studied by population genetics. In order to provide a global understanding of th…
Microbial populations adapt to their environment by acquiring advantageous mutations, but in the early twentieth century, questions about how these organisms acquire mutations arose. The experiment of Salvador Luria and Max Delbrück that won them a Nobel Prize in 1969 confirmed that mutations don't occur out of necessity, but instead can occur many generations before there is a selective advantage, and thus organisms follow Darwinian evolution instead of Lamarckian. Since then, new areas of research involving microbial evolution have spawned as a result of their experiment. Determining the mut…
We study a simple model of DNA evolution in a growing population of cells. Each cell contains a nucleotide sequence which randomly mutates at cell division. Cells divide according to a branching process. Following typical parameter values in bacteria and cancer cell populations, we take the mutation rate to zero and the final number of cells to infinity. We prove that almost every site (entry of the nucleotide sequence) is mutated in only a finite number of cells, and these numbers are independent across sites. However independence breaks down for the rare sites which are mutated in a positive…
We consider an asexually reproducing population on a finite type space whose evolution is driven by exponential birth, death and competition rates, as well as the possibility of mutation at a birth event. On the individual-based level this population can be modelled as a measure-valued Markov process. Multiple variations of this system have been studied in the simultaneous limit of large populations and rare mutations, where the regime is chosen such that mutations are separated. We consider the deterministic system, resulting from the large population limit, and then let the mutation probabil…
Mutation and drift play opposite roles in genetics. While mutation creates diversity, drift can cause gene variants to disappear, especially when they are rare. In the absence of natural selection and migration, the balance between the drift and mutation in a well-mixed population defines its diversity. The Moran model captures the effects of these two evolutionary forces and has a counterpart in social dynamics, known as the Voter model with external opinion influencers. Two extreme outcomes of the Voter model dynamics are consensus and coexistence of opinions, which correspond to low and hig…
It is known that learning of players who interact in a repeated game can be interpreted as an evolutionary process in a population of ideas. These analogies have so far mostly been established in deterministic models, and memory loss in learning has been seen to act similarly to mutation in evolution. We here propose a representation of reinforcement learning as a stochastic process in finite "populations of ideas". The resulting birth-death dynamics has absorbing states and allows for the extinction or fixation of ideas, marking a key difference to mutation-selection processes in finite popul…
In this paper, we study the number of equilibria of the replicator-mutator dynamics for both deterministic and random multi-player two-strategy evolutionary games. For deterministic games, using Decartes' rule of signs, we provide a formula to compute the number of equilibria in multi-player games via the number of change of signs in the coefficients of a polynomial. For two-player social dilemmas (namely, the Prisoner's Dilemma, Snowdrift, Stag Hunt, and Harmony), we characterize (stable) equilibrium points and analytically calculate the probability of having a certain number of equilibria wh…
Population expansions trigger many biomedical and ecological transitions, from tumor growth to invasions of non-native species. Although population spreading often selects for more invasive phenotypes, we show that this outcome is far from inevitable. In cooperative populations, mutations reducing dispersal have a competitive advantage. Such mutations then steadily accumulate at the expansion front bringing invasion to a halt. Our findings are a rare example of evolution driving the population into an unfavorable state and could lead to new strategies to combat unwelcome invaders. In addition,…
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…
We present a mechanistic formalism for the study of evolutionary dynamics models based on the diffusion approximation described by the Kimura Equation. In this formalism, the central component is the fitness potential, from which we obtain an expression for the amount of work necessary for a given type to reach fixation. In particular, within this interpretation, we develop a graphical analysis --- similar to the one used in classical mechanics --- providing the basic tool for a simple heuristic that describes both the short and long term dynamics. As a by-product, we provide a new definition …
People tend to have their social interactions with members of their own community. Such group-structured interactions can have a profound impact on the behaviors that evolve. Group structure affects the way people cooperate, and how they reciprocate each other's cooperative actions. Past work has shown that population structure and reciprocity can both promote the evolution of cooperation. Yet the impact of these mechanisms has been typically studied in isolation. In this work, we study how the two mechanisms interact. Using a game-theoretic model, we explore how people engage in reciprocal co…
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…
In this paper, we investigate the consequences of dormancy in the `rare mutation' and `large population' regime of stochastic adaptive dynamics. Starting from an individual-based micro-model, we first derive the polymorphic evolution sequence of the population, based on previous work by Baar and Bovier (2018). After passing to a second `small mutations' limit, we arrive at the canonical equation of adaptive dynamics, and state a corresponding criterion for evolutionary branching, extending a previous result of Champagnat and Méléard (2011). The criterion allows a quantitative and qualitative a…
