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Stochastic growth rates for life histories with rare migration or diapause

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 Added by David Steinsaltz
 Publication date 2015
  fields Biology
and research's language is English




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The growth of a population divided among spatial sites, with migration between the sites, is sometimes modelled by a product of random matrices, with each diagonal elements representing the growth rate in a given time period, and off-diagonal elements the migration rate. If the sites are reinterpreted as age classes, the same model may apply to a single population with age-dependent mortality and reproduction. We consider the case where the off-diagonal elements are small, representing a situation where there is little migration or, alternatively, where a deterministic life-history has been slightly disrupted, for example by introducing a rare delay in development. We examine the asymptotic behaviour of the long-term growth rate. We show that when the highest growth rate is attained at two different sites in the absence of migration (which is always the case when modelling a single age-structured population) the increase in stochastic growth rate due to a migration rate $epsilon$ is like $(log epsilon^{-1})^{-1}$ as $epsilondownarrow 0$, under fairly generic conditions. When there is a single site with the highest growth rate the behavior is more delicate, depending on the tails of the growth rates. For the case when the log growth rates have Gaussian-like tails we show that the behavior near zero is like a power of $epsilon$, and derive upper and lower bounds for the power in terms of the difference in the growth rates and the distance between the sites.



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The growth of a population divided among spatial sites, with migration between the sites, is sometimes modelled by a product of random matrices, with each diagonal elements representing the growth rate in a given time period, and off-diagonal elements the migration rate. The randomness of the matrices then represents stochasticity of environmental conditions. We consider the case where the off-diagonal elements are small, representing a situation where migration has been introduced into an otherwise sessile meta-population. We examine the asymptotic behaviour of the long-term growth rate. When there is a single site with the highest growth rate, under the assumption of Gaussian log growth rates at the individual sites (or having Gaussian-like tails) we show that the behavior near zero is like a power of $epsilon$, and derive upper and lower bounds for the power in terms of the difference in the growth rates and the distance between the sites. In particular, when the difference in mean log growth rate between two sites is sufficiently small, or the variance of the difference between the sites sufficiently large, migration will always be favored by natural selection, in the sense that introducing a small amount of migration will increase the growth rate of the population relative to the zero-migration case.
We follow up on a companion work that considered growth rates of populations growing at different sites, with different randomly varying growth rates at each site, in the limit as migration between sites goes to 0. We extend this work here to the special case where the maximum average log growth rate is achieved at two different sites. The primary motivation is to cover the case where `sites are understood as age classes for the same individuals. The theory then calculates the effect on growth rate of introducing a rare delay in development, a diapause, into an otherwise fixed-length semelparous life history. Whereas the increase in stochastic growth rate due to rare migrations was found to grow as a power of the migration rate, we show that under quite general conditions that in the diapause model --- or in the migration model with two or more sites having equal individual stochastic growth rates --- the increase in stochastic growth rate due to diapause at rate $epsilon$ behaves like $(log epsilon^{-1})^{-1}$ as $epsilondownarrow 0$. In particular, this implies that a small random disruption to the deterministic life history will always be favored by natural selection, in the sense that it will increase the stochastic growth rate relative to the zero-delay deterministic life history.
We consider stochastic matrix models for population driven by random environments which form a Markov chain. The top Lyapunov exponent $a$, which describes the long-term growth rate, depends smoothly on the demographic parameters (represented as matrix entries) and on the parameters that define the stochastic matrix of the driving Markov chain. The derivatives of $a$ -- the stochastic elasticities -- with respect to changes in the demographic parameters were derived by cite{tuljapurkar1990pdv}. These results are here extended to a formula for the derivatives with respect to changes in the Markov chain driving the environments. We supplement these formulas with rigorous bounds on computational estimation errors, and with rigorous derivations of both the new and the old formulas.
The stationary distribution of the diffusion limit of the 2-island, 2-allele Wright-Fisher with small but otherwise arbitrary mutation and migration rates is investigated. Following a method developed by Burden and Tang (2016, 2017) for approximating the forward Kolmogorov equation, the stationary distribution is obtained to leading order as a set of line densities on the edges of the sample space, corresponding to states for which one island is bi-allelic and the other island is non-segregating, and a set of point masses at the corners of the sample space, corresponding to states for which both islands are simultaneously non-segregating. Analytic results for the corner probabilities and line densities are verified independently using the backward generator and for the corner probabilities using the coalescent.
Stochastic epidemic models on networks are inherently high-dimensional and the resulting exact models are intractable numerically even for modest network sizes. Mean-field models provide an alternative but can only capture average quantities, thus offering little or no information about variability in the outcome of the exact process. In this paper we conjecture and numerically prove that it is possible to construct PDE-limits of the exact stochastic SIS epidemics on regular and ErdH{o}s-Renyi networks. To do this we first approximate the exact stochastic process at population level by a Birth-and-Death process (BD) (with a state space of $O(N)$ rather than $O(2^N)$) whose coefficients are determined numerically from Gillespie simulations of the exact epidemic on explicit networks. We numerically demonstrate that the coefficients of the resulting BD process are density-dependent, a crucial condition for the existence of a PDE limit. Extensive numerical tests for Regular and ErdH{o}s-Renyi networks show excellent agreement between the outcome of simulations and the numerical solution of the Fokker-Planck equations. Apart from a significant reduction in dimensionality, the PDE also provides the means to derive the epidemic outbreak threshold linking network and disease dynamics parameters, albeit in an implicit way. Perhaps more importantly, it enables the formulation and numerical evaluation of likelihoods for epidemic and network inference as illustrated in a worked out example.
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