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Register a new userOn the Simpson index for the Moran process with random selection and immigration
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Authors
Arnaud Personne
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Moran or Wright-Fisher processes are probably the most well known model to study the evolution of a population under various effects. Our object of study will be the Simpson index which measures the level of diversity of the population, one of the key parameter for ecologists who study for example forest dynamics. Following ecological motivations, we will consider here the case where there are various species with fitness and immigration parameters being random processes (and thus time evolving). To measure biodiversity, ecologists generally use the Simpson index, who has no closed formula, except in the neutral (no selection) case via a backward approach, and which is difficult to evaluate even numerically when the population size is large. Our approach relies on the large population limit in the weak selection case, and thus to give a procedure which enable us to approximate, with controlled rate, the expectation of the Simpson index at fixed time. Our approach will be forward and valid for all time, which is the main difference with the historical approach of Kingman, or Krone-Neuhauser. We will also study the long time behaviour of the Wright-Fisher process in a simplified setting, allowing us to get a full picture for the approximation of the expectation of the Simpson index.
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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 the sites in $A_m$ in individual $m$, where the individuals are sampled from the current population without replacement. Following the ancestry of these sites backwards in time yields a process on the set of partitions of $S$, which, in the diffusion limit, turns into a marginalised version of the $n$-locus ancestral recombination graph. With the help of an inclusion-exclusion principle, we show that the type distribution corresponding to a given partition may be represented in a systematic way, with the help of so-called recombinators and sampling functions. The same is true of correlation functions (known as linkage disequilibria in genetics) of all orders. We prove that the partitioning process (backward in time) is dual to the Moran population process (forward in time), where the sampling function plays the role of the duality function. This sheds new light on the work of Bobrowski, Wojdyla, and Kimmel (2010). The result also leads to a closed system of ordinary differential equations for the expectations of the sampling functions, which can be translated into expected type distributions and expected linkage disequilibria.
Using graphical methods based on a `lookdown and pruned version of the {em ancestral selection graph}, we obtain a representation of the type distribution of the ancestor in a two-type Wright-Fisher population with mutation and selection, conditional on the overall type frequency in the old population. This extends results from Lenz, Kluth, Baake, and Wakolbinger (Theor. Pop. Biol., 103 (2015), 27-37) to the case of heavy-tailed offspring, directed by a reproduction measure $Lambda$. The representation is in terms of the equilibrium tail probabilities of the line-counting process $L$ of the graph. We identify a strong pathwise Siegmund dual of $L$, and characterise the equilibrium tail probabilities of $L$ in terms of hitting probabilities of the dual process.
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 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 $1-tilde O(n^{-1/3})$, and the strong suppressor has fixation probability $tilde O(n^{-1/4})$. Both graph constructions are surprisingly simple. We also prove a general upper bound of $1-tilde Omega(n^{-1/3})$ on the fixation probability of any undirected graph. Hence, our strong amplifier is existentially optimal.
Consider a two-type Moran population of size $N$ subject to selection and mutation, which is immersed in a varying environment. The population is susceptible to exceptional changes in the environment, which accentuate the selective advantage of the fit individuals. In this setting, we show that the type-composition in the population is continuous with respect to the environment. This allows us to replace the deterministic environment by a random one, which is driven by a subordinator. Assuming that selection, mutation and the environment are weak in relation to $N$, we show that the type-frequency process, with time speed up by $N$, converges as $Ntoinfty$ to a Wright--Fisher-type SDE with a jump term modeling the effect of the environment. Next, we study the asymptotic behavior of the limiting model in the far future and in the distant past, both in the annealed and in the quenched setting. Our approach builds on the genealogical picture behind the model. The latter is described by means of an extension of the ancestral selection graph (ASG). The formal relation between forward and backward objects is given in the form of a moment duality between the type-frequency process and the line-counting process of a pruned version of the ASG. This relation yields characterizations of the annealed and the quenched moments of the asymptotic type distribution. A more involved pruning of the ASG allows us to obtain annealed and quenched results for the ancestral type distribution. In the absence of mutations, one of the types fixates and our results yield expressions for the fixation probabilities.
We consider the mutation--selection differential equation with pairwise interaction (or, equivalently, the diploid mutation--selection equation) and establish the corresponding ancestral process, which is a random tree and a variant of the ancestral selection graph. The formal relation to the forward model is given via duality. To make the tree tractable, we prune branches upon mutations, thus reducing it to its informative parts. The hierarchies inherent in the tree are encoded systematically via tripod trees with weighted leaves; this leads to the stratified ancestral selection graph. The latter also satisfies a duality relation with the mutation--selection equation. Each of the dualities provides a stochastic representation of the solution of the differential equation. This allows us to connect the equilibria and their bifurcations to the long-term behaviour of the ancestral process. Furthermore, with the help of the stratified ancestral selection graph, we obtain explicit results about the ancestral type distribution in the case of unidirectional mutation.
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