We reconsider the Moran model in continuous time with population size $N$, two allelic types, and selection. We introduce a new particle representation, which we call the labelled Moran model, and which has the same distribution of type frequencies as the original Moran model, provided the initial values are chosen appropriately. In the new model, individuals are labelled $1,2, \dots, N$; neutral resampling events may take place between arbitrary labels, whereas selective events only occur in the direction of increasing labels. With the help of elementary methods only, we not only recover fixation probabilities, but also obtain detailed insight into the number and nature of the selective events that play a role in the fixation process forward in time.
We consider two versions of stochastic population models with mutation and selection. The first approach relies on a multitype branching process; here, individuals reproduce and change type (i.e., mutate) independently of each other, without restriction on population size. We analyze the equilibrium behaviour of this model, both in the forward and in the backward direction of time; the backward point of view emerges if the ancestry of individuals chosen randomly from the present population is traced back into the past. The second approach is the Moran model with selection. Here, the population…
We study stochastic evolutionary game dynamics in a population of finite size. Individuals in the population are divided into two dynamically evolving groups. The structure of the population is formally described by a Wright-Fisher type Markov chain with a frequency dependent fitness. In a strong selection regime that favors one of the two groups, we obtain qualitatively matching lower and upper bounds for the fixation probability of the advantageous population. In the infinite population limit we obtain an exact result showing that a single advantageous mutant can invade an infinite populatio…
This paper is based on the complete classification of evolutionary scenarios for the Moran process with two strategies given by Taylor et al. (B. Math. Biol. 66(6): 1621--1644, 2004). Their classification is based on whether each strategy is a Nash equilibrium and whether the fixation probability for a single individual of each strategy is larger or smaller than its value for neutral evolution. We improve on this analysis by showing that each evolutionary scenario is characterized by a definite graph shape for the fixation probability function. A second class of results deals with the behavior…
We study fixation probabilities for the Moran stochastic process for the evolution of a population with three or more types of individuals and frequency-dependent fitnesses. Contrarily to the case of populations with two types of individuals, in which fixation probabilities may be calculated by an exact formula, here we must solve a large system of linear equations. We first show that this system always has a unique solution. Other results are upper and lower bounds for the fixation probabilities obtained by coupling the Moran process with three strategies with birth-death processes with only …
The ancestral selection graph in population genetics was introduced by KroneNeuhauser (1997) as an analogue of the coalescent genealogy of a sample of genes from a neutrally evolving population. The number of particles in this graph, followed backwards in time, is a birth and death process with quadratic death and linear birth rates. In this paper an explicit form of the probability distribution of the number of particles is obtained by using the density of the allele frequency in the corresponding diffusion model obtained by Kimura (1955). It is shown that the process of fixation of the allel…
We consider a multi-type Moran model (in continuous time) with selection and type-dependent mutation. This paper is concerned with the evolution of genealogical information forward in time. For this purpose we define and analytically characterize a path-valued Markov process that contains in its state at time $t$ the extended ancestral lines (adding genealogical distances) of the population alive at time $t$. The main result is a representation for the conditional distribution of the extended ancestral lines of a subpopulation alive at a fixed time $T$ (present time) given the type information…
Broom and Rychtář [Proc. R. Soc. A (2008) 464, 2609--2627] found an exact solution for the fixation probabilities of the Moran process for a structured population, in which the interaction structure among individuals is given by the so-called star graph, i.e. one central vertex and $n$ leaves, the leaves connecting only to the center. We generalize on their solution by allowing individuals' fitnesses to depend on the population frequency, and also by allowing a possible change in the order of reproduction and death draws. In their cited paper, Broom and Rychtář considered the birth-death (BD) …
In population genetics, extant samples are usually used for inference of past population genetic forces. With the Kingman coalescent and the backward diffusion equation, inference of the marginal likelihood proceeds from an extant sample backward in time. Conditional on an extant sample, the Moran model can also be used backward in time with identical results, up to a scaling of time. In particular, all three approaches -- the coalescent, the backward diffusion, and the Moran model -- lead to the identical marginal likelihood of the sample. If probabilities of ancestral states are also inferre…
Evolutionary dynamics has been classically studied for homogeneous populations, but now there is a growing interest in the non-homogenous case. One of the most important models has been proposed by Lieberman, Hauert and Nowak, adapting to a weighted directed graph the classical process described by Moran. The Markov chain associated with the graph can be modified by erasing all non-trivial loops in its state space, obtaining the so-called Embedded Markov chain (EMC). The fixation probability remains unchanged, but the expected time to absorption (fixation or extinction) is reduced. In this pap…
Fixation probabilities for multi-allele Moran dynamics with weak selection
Center for Advanced Systems Understanding (CASUS) – Helmholtz-Zentrum Dresden-Rossendorf (HZDR)（高级系统理解中心（CASUS）– 德累斯顿-罗斯多夫亥姆霍兹中心（HZDR））；Universidade Federal de Minas Gerais（米纳斯吉拉斯联邦大学）；ICTP South American Institute for Fundamental Research & Instituto de Física Teórica, Universidade Estadual Paulista - UNESP（国际理论物理中心南美基础研究所与理论物理研究所，圣保罗州立大学（UNESP））；Department of Ecology, Institute of Biosciences, University of São Paulo（圣保罗大学生物科学学院生态学系）
Evolutionary graph theory models the effects of natural selection and random drift on structured populations of competing mutant and non-mutant individuals. Recent studies have found that fixation times in such systems often have right-skewed distributions. Little is known, however, about how these distributions and their skew depend on mutant fitness. Here we calculate the fitness dependence of the fixation-time distribution for the Moran Birth-death process in populations modeled by two extreme networks: the complete graph and the one-dimensional ring lattice, obtaining exact solutions in th…
The Moran process is a classic stochastic process that models invasion dynamics on graphs. A single "mutant" (e.g., a new opinion, strain, social trait etc.) invades a population of residents spread over the nodes of a graph. The mutant fitness advantage $δ\geq 0$ determines how aggressively mutants propagate to their neighbors. The quantity of interest is the fixation probability, i.e., the probability that the initial mutant eventually takes over the whole population. However, in realistic settings, the invading mutant has an advantage only in certain locations. E.g., a bacterial mutation al…
The Moran model with recombination is considered, which describes the evolution of the genetic composition of a population under recombination and resampling. There are $n$ sites (or loci), a finite number of letters (or alleles) at every site, and we do not make any scaling assumptions. In particular, we do not assume a diffusion limit. We consider the following marginal ancestral recombination process. Let $S = \{1,...,n\}$ and $\mathcal A=\{A_1, ..., A_m\}$ be a partition of $S$. We concentrate on the joint probability of the letters at the sites in $A_1$ in individual $1$, $...$, and at th…
A two-types, discrete-time population model with finite, constant size is constructed, allowing for a general form of frequency-dependent selection and skewed offspring distribution. Selection is defined based on the idea that individuals first choose a (random) number of $\textit{potential}$ parents from the previous generation and then, from the selected pool, they inherit the type of the fittest parent. The probability distribution function of the number of potential parents per individual thus parametrises entirely the selection mechanism. Using sampling- and moment-duality, weak convergen…
We consider the classic Moran process modeling the spread of genetic mutations, as extended to structured populations by Lieberman et al.\ (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 ther…
The Moran process is a random process that models the spread of genetic mutations through graphs. If the graph is connected, the process eventually reaches "fixation", where every vertex is a mutant, or "extinction", where no vertex is a mutant. Our main result is an almost-tight bound on expected absorption time. For all epsilon > 0, we show that the expected absorption time on an n-vertex graph is o(n^(3+epsilon)). In fact, we show that it is at most n^3 * exp(O((log log n)^3)) and that there is a family of graphs where it is Omega(n^3). In the course of proving our main result, we also esta…
Many mathematical models of evolution assume that all individuals experience the same environment. Here, we study the Moran process in heterogeneous environments. The population is of finite size with two competing types, which are exposed to a fixed number of environmental conditions. Reproductive rate is determined by both the type and the environment. We first calculate the condition for selection to favor the mutant relative to the resident wild type. In large populations, the mutant is favored if and only if the mutant's spatial average reproductive rate exceeds that of the resident. But …
We consider a Moran model with recombination in a haploid population of size $N$. At each birth event, with probability $1-ρ_N R$ the offspring copies one parent's chromosome, and with probability $ρ_N R$ she inherits a chromosome that is a mosaic of both parental chromosomes. We assume that at time $0$ each individual has her chromosome painted in a different color and we study the color partition of the chromosome that is asymptotically fixed in a large population, when we look at a portion of the chromosome such that $ρ:= \lim_{N\to \infty} \frac{ρ_N N}{2}\to \infty$. To do so, we follow ba…
The fixation probability of a single mutant invading a population of residents is among the most widely-studied quantities in evolutionary dynamics. Amplifiers of natural selection are population structures that increase the fixation probability of advantageous mutants, compared to well-mixed populations. Extensive studies have shown that many amplifiers exist for the Birth-death Moran process, some of them substantially increasing the fixation probability or even guaranteeing fixation in the limit of large population size. On the other hand, no amplifiers are known for the death-Birth Moran p…
We present a new model for seed banks, where direct ancestors of individuals may have lived in the near as well as the very far past. The classical Wright-Fisher model, as well as a seed bank model with bounded age distribution considered by Kaj, Krone and Lascoux (2001) are special cases of our model. We discern three parameter regimes of the seed bank age distribution, which lead to substantially different behaviour in terms of genetic variability, in particular with respect to fixation of types and time to the most recent common ancestor. We prove that for age distributions with finite mean…
The evolutionary advantage of replacers in the Moran process
Computer Science Institute, Charles University（查尔斯大学计算机科学学院）；Department of Organismic and Evolutionary Biology, Harvard University（哈佛大学生物演化系）；Department of Mathematics, Harvard University（哈佛大学数学系）
For a beneficial allele which enters a large unstructured population and eventually goes to fixation, it is known that the time to fixation is approximately $2\log(α)/α$ for a large selection coefficient $α$. For a population that is distributed over finitely many colonies, with migration between these colonies, we detect various regimes of the migration rate $μ$ for which the fixation times have different asymptotics as $α\to \infty$. If $μ$ is of order $α$, the allele fixes (as in the spatially unstructured case) in time $\sim 2\log(α)/α$. If $μ$ is of order $α^γ, 0\leq γ\leq 1$, the fixatio…
Population structures can be crucial determinants of evolutionary processes. For the Moran process on graphs certain structures suppress selective pressure, while others amplify it (Lieberman et al. 2005 Nature 433 312-316). Evolutionary amplifiers suppress random drift and enhance selection. Recently, some results for the most powerful known evolutionary amplifier, the superstar, have been invalidated by a counter example (Díaz et al. 2013 Proc. R. Soc. A 469 20130193). Here we correct the original proof and derive improved upper and lower bounds, which indicate that the fixation probability …
We consider the Moran process in a graph called the "star" and obtain the asymptotic expression for the fixation probability of a single mutant when the size of the graph is large. The expression obtained corrects the previously known expression announced in reference [E Lieberman, C Hauert, and MA Nowak. Evolutionary dynamics on graphs. Nature, 433(7023):312-316, 2005] and further studied in [M. Broom and J. Rychtar. An analysis of the fixation probability of a mutant on special classes of non-directed graphs. Proc. R. Soc. A-Math. Phys. Eng. Sci., 464(2098):2609-2627, 2008]. We also show tha…
The Moran process, as studied by [Lieberman, E., Hauert, C. and Nowak, M. Evolutionary dynamics on graphs. Nature 433, pp. 312-316 (2005)], 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 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…
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 o…
The Moran process on graphs is a popular model to study the dynamics of evolution in a spatially structured population. Exact analytical solutions for the fixation probability and time of a new mutant have been found for only a few classes of graphs so far. Simulations are time-expensive and many realizations are necessary, as the variance of the fixation times is high. We present an algorithm that numerically computes these quantities for arbitrary small graphs by an approach based on the transition matrix. The advantage over simulations is that the calculation has to be executed only once. B…
We define the incentive process, a natural generalization of the Moran process incorporating evolutionary updating mechanisms corresponding to well-known evolutionary dynamics, such as the logit, projection, and best-reply dynamics. Fixation probabilities and internal stable states are given for a variety of incentives, including new closed-forms, as well as results relating fixation probabilities for members of two one-parameter families of incentive processes. We show that the behaviors of the incentive process can deviate significantly from the analogous properties of deterministic evolutio…
We consider the Moran model in continuous time with two types, mutation, and selection. We concentrate on the ancestral line and its stationary type distribution. Building on work by Fearnhead (J. Appl. Prob. 39 (2002), 38-54) and Taylor (Electron. J. Probab. 12 (2007), 808-847), we characterise this distribution via the fixation probability of the offspring of all individuals of favourable type (regardless of the offspring's types). We concentrate on a finite population and stay with the resulting discrete setting all the way through. This way, we extend previous results and gain new insight …
Temporal environmental variations are ubiquitous in nature, yet most of the theoretical works in population genetics and evolution assume fixed environment. Here we analyze the effect of variations in carrying capacity on the fate of a mutant type. We consider a two-state Moran model, where selection intensity at equilibrium may differ (in amplitude and in sign) from selection during periods of sharp growth and sharp decline. Using Kimura's diffusion approximation we present simple formulae for effective population size and effective selection, and use it to calculate the chance of ultimate fi…
This work is a systematic study of discrete Markov chains that are used to describe the evolution of a two-types population. Motivated by results valid for the well-known Moran (M) and Wright-Fisher (WF) processes, we define a general class of Markov chains models which we term the Kimura class. It comprises the majority of the models used in population genetics, and we show that many well-known results valid for M and WF processes are still valid in this class. In all Kimura processes, a mutant gene will either fixate or become extinct, and we present a necessary and sufficient condition for …
The mean fixation time of a deleterious mutant allele is studied beyond the diffusion approximation. As in Kimura's classical work [M. Kimura, Proc. Natl. Acad. Sci. U.S.A. Vol.77, 522 (1980)], that was motivated by the problem of fixation in the presence of amorphic or hypermorphic mutations, we consider a diallelic model at a single locus comprising a wild-type A and a mutant allele A' produced irreversibly from A at small uniform rate v. The relative fitnesses of the mutant homozygotes A'A', mutant heterozygotes A'A and wild-type homozygotes AA are 1-s, 1-h and 1, respectively, where it is …
Tumors initiate when a population of proliferating cells accumulates a certain number and type of genetic and/or epigenetic alterations. The population dynamics of such sequential acquisition of (epi)genetic alterations has been the topic of much investigation. The phenomenon of stochastic tunneling, where an intermediate mutant in a sequence does not reach fixation in a population before generating a double mutant, has been studied using a variety of computational and mathematical methods. However, the field still lacks a comprehensive analytical description since theoretical predictions of f…
