No Arabic abstract
In this paper I will review twenty years of work on the question: When is there coexistence in stochastic spatial models? The answer, announced in Durrett and Levin [Theor. Pop. Biol. 46 (1994) 363--394], and that we explain in this paper is that this can be determined by examining the mean-field ODE. There are a number of rigorous results in support of this picture, but we will state nine challenging and important open problems, most of which date from the 1990s.
The aim of this paper is to tackle part of the program set by Diekmann et al. in their seminal paper Diekmann et al. (2001). We quote It remains to investigate whether, and in what sense, the nonlinear determin-istic model formulation is the limit of a stochastic model for initial population size tending to infinity We set a precise and general framework for a stochastic individual based model : it is a piecewise deterministic Markov process defined on the set of finite measures. We then establish a law of large numbers under conditions easy to verify. Finally we show how this applies to old and new examples.
We discuss similarities and differences between systems of interacting players maximizing their individual payoffs and particles minimizing their interaction energy. Long-run behavior of stochastic dynamics of spatial games with multiple Nash equilibria is analyzed. In particular, we construct an example of a spatial game with three strategies, where stochastic stability of Nash equilibria depends on the number of players and the kind of dynamics.
Field theory tools are applied to analytically study fluctuation and correlation effects in spatially extended stochastic predator-prey systems. In the mean-field rate equation approximation, the classic Lotka-Volterra model is characterized by neutral cycles in phase space, describing undamped oscillations for both predator and prey populations. In contrast, Monte Carlo simulations for stochastic two-species predator-prey reaction systems on regular lattices display complex spatio-temporal structures associated with persistent erratic population oscillations. The Doi-Peliti path integral representation of the master equation for stochastic particle interaction models is utilized to arrive at a field theory action for spatial Lotka-Volterra models in the continuum limit. In the species coexistence phase, a perturbation expansion with respect to the nonlinear predation rate is employed to demonstrate that spatial degrees of freedom and stochastic noise induce instabilities toward structure formation, and to compute the fluctuation corrections for the oscillation frequency and diffusion coefficient. The drastic downward renormalization of the frequency and the enhanced diffusivity are in excellent qualitative agreement with Monte Carlo simulation data.
Motivated by models of cancer formation in which cells need to acquire $k$ mutations to become cancerous, we consider a spatial population model in which the population is represented by the $d$-dimensional torus of side length $L$. Initially, no sites have mutations, but sites with $i-1$ mutations acquire an $i$th mutation at rate $mu_i$ per unit area. Mutations spread to neighboring sites at rate $alpha$, so that $t$ time units after a mutation, the region of individuals that have acquired the mutation will be a ball of radius $alpha t$. We calculate, for some ranges of the parameter values, the asymptotic distribution of the time required for some individual to acquire $k$ mutations. Our results, which build on previous work of Durrett, Foo, and Leder, are essentially complete when $k = 2$ and when $mu_i = mu$ for all $i$.
Classical ecological theory predicts that environmental stochasticity increases extinction risk by reducing the average per-capita growth rate of populations. To understand the interactive effects of environmental stochasticity, spatial heterogeneity, and dispersal on population growth, we study the following model for population abundances in $n$ patches: the conditional law of $X_{t+dt}$ given $X_t=x$ is such that when $dt$ is small the conditional mean of $X_{t+dt}^i-X_t^i$ is approximately $[x^imu_i+sum_j(x^j D_{ji}-x^i D_{ij})]dt$, where $X_t^i$ and $mu_i$ are the abundance and per capita growth rate in the $i$-th patch respectivly, and $D_{ij}$ is the dispersal rate from the $i$-th to the $j$-th patch, and the conditional covariance of $X_{t+dt}^i-X_t^i$ and $X_{t+dt}^j-X_t^j$ is approximately $x^i x^j sigma_{ij}dt$. We show for such a spatially extended population that if $S_t=(X_t^1+...+X_t^n)$ is the total population abundance, then $Y_t=X_t/S_t$, the vector of patch proportions, converges in law to a random vector $Y_infty$ as $ttoinfty$, and the stochastic growth rate $lim_{ttoinfty}t^{-1}log S_t$ equals the space-time average per-capita growth rate $sum_imu_iE[Y_infty^i]$ experienced by the population minus half of the space-time average temporal variation $E[sum_{i,j}sigma_{ij}Y_infty^i Y_infty^j]$ experienced by the population. We derive analytic results for the law of $Y_infty$, find which choice of the dispersal mechanism $D$ produces an optimal stochastic growth rate for a freely dispersing population, and investigate the effect on the stochastic growth rate of constraints on dispersal rates. Our results provide fundamental insights into ideal free movement in the face of uncertainty, the persistence of coupled sink populations, the evolution of dispersal rates, and the single large or several small (SLOSS) debate in conservation biology.