Adaptation in a stochastic multi-resources chemostat model

Abstract: We are interested in modeling the Darwinian evolution resulting from the interplay of phenotypic variation and natural selection through ecological interactions, in the specific scales of the biological framework of adaptive dynamics. Adaptive dynamics so far has been put on a rigorous footing only for direct competition models (Lotka-Volterra models) involving a competition kernel which describes the competition pressure from one individual to another one. We extend this to a multi-resources chemostat model, where the competition between individuals results from the sharing of several resources which have their own dynamics. Starting from a stochastic birth and death process model, we prove that, when advantageous mutations are rare, the population behaves on the mutational time scale as a jump process moving between equilibrium states (the polymorphic evolution sequence of the adaptive dynamics literature). An essential technical ingredient is the study of the long time behavior of a chemostat multi-resources dynamical system. In the small mutational steps limit this process in turn gives rise to a differential equation in phenotype space called canonical equation of adaptive dynamics. From this canonical equation and still assuming small mutation steps, we prove a rigorous characterization of the evolutionary branching points.

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The goal of this paper is to study the Darwinian evolution of genetic parameters of bacteria in the chemostat model. In particular, it is studied how mutations and competition for nutrients lead to their progressive adaptation. In comparison with previous works, this model is more realistic, because it takes into account explicit dynamics for several resources. The outcome of such consideration is a model of competitive interactions between bacteria, driving selection, with a more complicated nonlinearity than the Lotka-Volterra direct competition. The adaptive dynamics of this model is studied combining large populations, rare and small mutations. Bacteria dynamics follows a birth and death process with reproduction due to resources consumption and mutations. The resource process is deterministic between birth and death events in the bacteria population; then is introduced a multi-resource chemostat with deterministic systems as a large-size approximation of these probabilistic models. Studying the behavior of such deterministic nonlinear systems, the authors prove the convergence to a unique equilibrium as soon as some non-degeneracy assumptions hold. It is also proved that (1) all traits with zero density at equilibrium actually go extinct after a time of the order the logarithm of the population size; (2) the time of exit from a neighborhood of the equilibrium grows exponentially in the population size. Finally, it is studied the individual-based process on the evolutionary time scale in three steps: (1) large population and rare mutation, but the size of each mutation does not go to 0; (2) scaling of small mutations in the PES, letting the size of each mutation go to 0 (it should be noticed that in this limit the dynamics of coexisting traits is governed by an ODE extending the canonical equation of adaptive dynamics); (3) taking into account this canonical equation and assuming small mutation steps, it is proved that the coexistence is maintained after evolutionary branching, and that the distance between the branches increases.

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