Invasion and adaptive evolution for individual-based spatially structured populations

The interplay between space and evolution is an important issue in population dynamics, that is in particular crucial in the emergence of polymorphism and spatial patterns. Recently, biological studies suggest that invasion and evolution are closely related. Here we model the interplay between space and evolution starting with an individual-based approach and show the important role of parameter scalings on clustering and invasion. We consider a stochastic discrete model with birth, death, competition, mutation and spatial diffusion, where all the parameters may depend both on the position and on the trait of individuals. The spatial motion is driven by a reflected diffusion in a bounded domain. The interaction is modelled as a trait competition between individuals within a given spatial interaction range. First, we give an algorithmic construction of the process. Next, we obtain large population approximations, as weak solutions of nonlinear reaction-diffusion equations with Neumann's boundary conditions. As the spatial interaction range is fixed, the nonlinearity is nonlocal. Then, we make the interaction range decrease to zero and prove the convergence to spatially localized nonlinear reaction-diffusion equations, with Neumann's boundary conditions. Finally, simulations based on the microscopic individual-based model are given, illustrating the strong effects of the spatial interaction range on the emergence of spatial and phenotypic diversity (clustering and polymorphism) and on the interplay between invasion and evolution. The simulations focus on the qualitative differences between local and nonlocal interactions

Some stochastic models for structured populations : scaling limits and long time behavior

The first chapter concerns monotype population models. We first study general birth and death processes and we give non-explosion and extinction criteria, moment computations and a pathwise representation. We then show how different scales may lead to different qualitative approximations, either ODEs or SDEs. The prototypes of these equations are the logistic (deterministic) equation and the logistic Feller diffusion process. The convergence in law of the sequence of processes is proved by tightness-uniqueness argument. In these large population approximations, the competition between individuals leads to nonlinear drift terms. We then focus on models without interaction but including exceptional events due either to demographic stochasticity or to environmental stochasticity. In the first case, an individual may have a large number of offspring and we introduce the class of continuous state branching processes. In the second case, catastrophes may occur and kill a random fraction of the population and the process enjoys a quenched branching property. We emphasize on the study of the Laplace transform, which allows us to classify the long time behavior of these processes. In the second chapter, we model structured populations by measure-valued stochastic differential equations. Our approach is based on the individual dynamics. The individuals are characterized by parameters which have an influence on their survival or reproduction ability. Some of these parameters can be genetic and are inheritable except when mutations occur, but they can also be a space location or a quantity of parasites. The individuals compete for resources or other environmental constraints. We describe the population by a point measure-valued Markov process. We study macroscopic approximations of this process depending on the interplay between different scalings and obtain in the limit either integro-differential equations or reaction-diffusion equations or nonlinear super-processes. In each case, we insist on the specific techniques for the proof of convergence and for the study of the limiting model. The limiting processes offer different models of mutation-selection dynamics. Then, we study two-level models motivated by cell division dynamics, where the cell population is discrete and characterized by a trait, which may be continuous. In 1 particular, we finely study a process for parasite infection and the trait is the parasite load. The latter grows following a Feller diffusion and is randomly shared in the two daughter cells when the cell divides. Finally, we focus on the neutral case when the rate of division of cells is constant but the trait evolves following a general Markov process and may split in a random number of cells. The long time behavior of the structured population is then linked and derived from the behavior a well chosen SDE (monotype population)

Uniform estimates for metastable transition times in a coupled bistable system

We consider a coupled bistable N-particle system driven by a Brownian noise, with a strong coupling corresponding to the synchronised regime. Our aim is to obtain sharp estimates on the metastable transition times between the two stable states, both for fixed N and in the limit when N tends to infinity, with error estimates uniform in N. These estimates are a main step towards a rigorous understanding of the metastable behavior of infinite dimensional systems, such as the stochastically perturbed Ginzburg-Landau equation. Our results are based on the potential theoretic approach to metastability.Comment: 20 page

Speed of coming down from infinity for birth and death processes

We finely describe the speed of "coming down from infinity" for birth and death processes which eventually become extinct. Under general assumptions on the birth and death rates, we firstly determine the behavior of the successive hitting times of large integers. We put in light two different regimes depending on whether the mean time for the process to go from $n+1$ to $n$ is negligible or not compared to the mean time to reach $n$ from infinity. In the first regime, the coming down from infinity is very fast and the convergence is weak. In the second regime, the coming down from infinity is gradual and a law of large numbers and a central limit theorem for the hitting times sequence hold. By an inversion procedure, we deduce that the process is a.s. equivalent to a non-increasing function when the time goes to zero. Our results are illustrated by several examples including applications to population dynamics and population genetics. The particular case where the death rate varies regularly is studied in details.Comment: 30 pages. arXiv admin note: text overlap with arXiv:1310.740

A host-parasite multilevel interacting process and continuous approximations

We are interested in modeling some two-level population dynamics, resulting from the interplay of ecological interactions and phenotypic variation of individuals (or hosts) and the evolution of cells (or parasites) of two types living in these individuals. The ecological parameters of the individual dynamics depend on the number of cells of each type contained by the individual and the cell dynamics depends on the trait of the invaded individual. Our models are rooted in the microscopic description of a random (discrete) population of individuals characterized by one or several adaptive traits and cells characterized by their type. The population is modeled as a stochastic point process whose generator captures the probabilistic dynamics over continuous time of birth, mutation and death for individuals and birth and death for cells. The interaction between individuals (resp. between cells) is described by a competition between individual traits (resp. between cell types). We look for tractable large population approximations. By combining various scalings on population size, birth and death rates and mutation step, the single microscopic model is shown to lead to contrasting nonlinear macroscopic limits of different nature: deterministic approximations, in the form of ordinary, integro- or partial differential equations, or probabilistic ones, like stochastic partial differential equations or superprocesses. The study of the long time behavior of these processes seems very hard and we only develop some simple cases enlightening the difficulties involved

Estimates for the density of a nonlinear Landau process

The aim of this paper is to obtain estimates for the density of the law of a specific nonlinear diffusion process at any positive bounded time. This process is issued from kinetic theory and is called Landau process, by analogy with the associated deterministic Fokker-Planck-Landau equation. It is not Markovian, its coefficients are not bounded and the diffusion matrix is degenerate. Nevertheless, the specific form of the diffusion matrix and the nonlinearity imply the non-degeneracy of the Malliavin matrix and then the existence and smoothness of the density. In order to obtain a lower bound for the density, the known results do not apply. However, our approach follows the main idea consisting in discretizing the interval time and developing a recursive method. To this aim, we prove and use refined results on conditional Malliavin calculus. The lower bound implies the positivity of the solution of the Landau equation, and partially answers to an analytical conjecture. We also obtain an upper bound for the density, which again leads to an unusual estimate due to the bad behavior of the coefficients

Quasi-stationary distributions and population processes

This survey concerns the study of quasi-stationary distributions with a specific focus on models derived from ecology and population dynamics. We are concerned with the long time behavior of different stochastic population size processes when 0 is an absorbing point almost surely attained by the process. The hitting time of this point, namely the extinction time, can be large compared to the physical time and the population size can fluctuate for large amount of time before extinction actually occurs. This phenomenon can be understood by the study of quasi-limiting distributions. In this paper, general results on quasi-stationarity are given and examples developed in detail. One shows in particular how this notion is related to the spectral properties of the semi-group of the process killed at 0. Then we study different stochastic population models including nonlinear terms modeling the regulation of the population. These models will take values in countable sets (as birth and death processes) or in continuous spaces (as logistic Feller diffusion processes or stochastic Lotka-Volterra processes). In all these situations we study in detail the quasi-stationarity properties. We also develop an algorithm based on Fleming-Viot particle systems and show a lot of numerical pictures