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.
Several birth-and-death processes on graphs have been studied, beside the Moran process, which vary in the order in which the birth and death events occur in a step, and on whether the selection bias introduced by fitness is applied in the choice of the vertex that reproduces or the one that dies. The Voter model @cite_26 is a well-known such process. It has been observe that small differences in these processes have significant impact on the fixation probability @cite_13 @cite_4 @cite_11 . In particular, for dynamics where, unlike in Moran process, the vertex to die is chosen first, it is known that simple strong suppressors exits, such as the star @math with fixation probability @math , while there are no strong amplifiers @cite_19 . Although not elaborated in the current paper, our results carry over to the process where the vertex to reproduce is chosen uniformly at random, and its offspring replaces a neighbor chosen with probability inversely proportional to its fitness. This corresponds to a setting where advantageous mutants do not reproduce more often, but they live longer.
@cite_26 @cite_4 @cite_19 @cite_13 @cite_11
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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. 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. The Moran process, as studied by Lieberman, Hauert, and Nowak, is a randomised algorithm modelling the spread of genetic mutations in populations. The algorithm runs on an underlying graph where individuals correspond to vertices. Initially, one vertex (chosen uniformly at random) possesses a mutation, with fitness r > 1. All other individuals have fitness 1. During each step of the algorithm, an individual is chosen with probability proportional to its fitness, and its state (mutant or nonmutant) is passed on to an out-neighbour which is chosen uniformly at random. If the underlying graph is strongly connected, then the algorithm will eventually reach fixation, in which all individuals are mutants, or extinction, in which no individuals are mutants. An infinite family of directed graphs is said to be strongly amplifying if, for every r > 1, the extinction probability tends to 0 as the number of vertices increases. A formal definition is provided in the article. Strong amplification is a rather surprising property—it means that in such graphs, the fixation probability of a uniformly placed initial mutant tends to 1 even though the initial mutant only has a fixed selective advantage of r > 1 (independently of n). The name “strongly amplifying” comes from the fact that this selective advantage is “amplified.” Strong amplifiers have received quite a bit of attention, and proposed two potentially strongly amplifying families—superstars and metafunnels. Heuristic arguments have been published, arguing that there are infinite families of superstars that are strongly amplifying. The same has been claimed for metafunnels. In this article, we give the first rigorous proof that there is an infinite family of directed graphs that is strongly amplifying. We call the graphs in the family “megastars.” When the algorithm is run on an n-vertex graph in this family, starting with a uniformly chosen mutant, the extinction probability is roughly n − 1 2 (up to logarithmic factors). We prove that all infinite families of superstars and metafunnels have larger extinction probabilities (as a function of n). Finally, we prove that our analysis of megastars is fairly tight—there is no infinite family of megastars such that the Moran algorithm gives a smaller extinction probability (up to logarithmic factors). Also, we provide a counterexample which clarifies the literature concerning the isothermal theorem of 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.