Deterministic evolutionary theory robustly predicts that populations displaying altruistic behaviours will be driven to extinction by mutant cheats that absorb common benefits but do not themselves contribute. Here we show that when demographic stochasticity is accounted for, selection can in fact act in the reverse direction to that predicted deterministically, instead favouring cooperative behaviors that appreciably increase the carrying capacity of the population. Populations that exist in larger numbers experience a selective advantage by being more stochastically robust to invasions than smaller populations, and this advantage can persist even in the presence of reproductive costs. We investigate this general effect in the specific context of public goods production and find conditions for stochastic selection reversal leading to the success of public good producers. This insight, developed here analytically, is missed by both the deterministic analysis as well as standard game theoretic models that enforce a fixed population size. The effect is found to be amplified by space; in this scenario we find that selection reversal occurs within biologically reasonable parameter regimes for microbial populations. Beyond the public good problem, we formulate a general mathematical framework for models that may exhibit stochastic selection reversal. In this context, we describe a stochastic analogue to r-K theory, by which small populations can evolve to higher densities in the absence of disturbance.
We explore a class of hybrid (piecewise deterministic) systems characterized by a large number of individuals inhabiting an environment whose state is described by a set of continuous variables. We use analytical and numerical methods from nonequilibrium statistical mechanics to study the influence that intrinsic noise has on the qualitative behavior of the system. We discuss the application of these concepts to the case of semiarid ecosystems. Using a system-size expansion we calculate the power spectrum of the fluctuations in the system. This predicts the existence of noise-induced oscillati…
Recent theoretical studies have shown that demographic stochasticity can greatly increase the tendency of asexually reproducing phenotypically diverse organisms to spontaneously evolve into localised clusters, suggesting a simple mechanism for sympatric speciation. Here we study the role of demographic stochasticity in a model of competing organisms subject to assortative mating. We find that in models with sexual reproduction, noise can also lead to the formation of phenotypic clusters in parameter ranges where deterministic models would lead to a homogeneous distribution. In some cases, nois…
We study simple stochastic scenarios, based on birth-and-death Markovian processes, that describe populations with Allee effect, to account for the role of demographic stochasticity. In the mean-field deterministic limit we recover well-known deterministic evolution equations widely employed in population ecology. The mean-time to extinction is in general obtained by the Wentzel-Kramers-Brillouin (WKB) approximation for populations with strong and weak Allee effects. An exact solution for the mean time to extinction can be found via a recursive equation for special cases of the stochastic dyna…
We present an explicit unified stochastic model of fluctuations in population size due to random birth, death, density-dependent competition and environmental fluctuations. Stochastic dynamics provide insight into small populations, including processes such as extinction, that cannot be correctly treated by deterministic methods. We present exact analytical and simulation-based results for extinction times of our stochastic model and compare the different effects of environmental stochasticity and intrinsic demographic stochasticity. We use both the discrete master equation approach and an exa…
It has been shown that differences in fecundity variance can influence the probability of invasion of a genotype in a population, i.e. a genotype with lower variance in offspring number can be favored in finite populations even if it has a somewhat lower mean fitness than a competitor. In this paper, Gillespie's results are extended to population genetic systems with explicit age structure, where the demographic variance (variance in growth rate) calculated in the work of Engen and colleagues is used as a generalization of "variance in offspring number" to predict the interaction between deter…
Recent collapses of many fisheries across the globe have challenged the mathematical approach to these systems through classic bioeconomic models. Decimated populations did not recover as fast as predicted by these models and depensatory effects were introduced to better fit the dynamics at low population abundances. Alternative to depensation, modeling captures by non-linear harvesting functions produces equivalent outcomes at small abundances, and the dynamics undergoes a bifurcation leading to population collapse and recovery once catching efforts are above or below certain thresholds, resp…
We study the phenomenon of cyclic dominance in the paradigmatic Rock--Paper--Scissors model, as occurring in both stochastic individual-based models of finite populations and in the deterministic replicator equations. The mean-field replicator equations are valid in the limit of large populations and, in the presence of mutation and unbalanced payoffs, they exhibit an attracting limit cycle. The period of this cycle depends on the rate of mutation; specifically, the period grows logarithmically as the mutation rate tends to zero. We find that this behaviour is not reproduced in stochastic simu…
When predicting the fate and consequences of recurring deleterious mutations in self-fertilising populations most models developed make the assumption that populations have discrete non-overlapping generations. This makes them biologically irrelevant when considering perennial species with over-lapping generations and where mating occurs independently of the age group. Previous models studying the effect of perennial life-histories on the genetic properties of populations in the presence of self-fertilisation have done so considering age-dependent selection and have found that, contrary to emp…
Demographic (shot) noise in population dynamics scales with the square root of the population size. This process is very important, as it yields an absorbing state at zero field, but simulating it, especially on spatial domains, is a non-trivial task. Here we compare the results of two operator-splitting techniques suggested for simulating the corresponding Langevin equation, one by Pechenik and Levine (PL) and the other by Dornic, Chaté and Muñoz (DCM). We identify an anomalously strong bias toward the active phase in the numerical scheme of DCM, a bias which is not present in the alternative…
In large but finite populations, weak demographic stochasticity due to random birth and death events can lead to population extinction. The process is analogous to the escaping problem of trapped particles under random forces. Methods widely used in studying such physical systems, for instance, Wentzel-Kramers-Brillouin (WKB) and Fokker-Planck methods, can be applied to solve similar biological problems. In this article, we comparatively analyse applications of WKB and Fokker-Planck methods to some typical stochastic population dynamical models, including the logistic growth, endemic SIR, pred…
We study the fixation probability of a mutant type when introduced into a resident population. As opposed to the usual assumption of constant pop- ulation size, we allow for stochastically varying population sizes. This is implemented by a stochastic competitive Lotka-Volterra model. The compe- tition coefficients are interpreted in terms of inverse payoffs emerging from an evolutionary game. Since our study focuses on the impact of the competition values, we assume the same birth and death rates for both types. In this gen- eral framework, we derive an approximate formula for the fixation pro…
Understanding the causes and effects of spatial aggregation is one of the most fundamental problems in ecology. Aggregation is an emergent phenomenon arising from the interactions between the individuals of the population, able to sense only -at most- local densities of their cohorts. Thus, taking into account the individual-level interactions and fluctuations is essential to reach a correct description of the population. Classic deterministic equations are suitable to describe some aspects of the population, but leave out features related to the stochasticity inherent to the discreteness of t…
The scarcity of water characterising drylands forces vegetation to adopt appropriate survival strategies. Some of these generate water-vegetation feedback mechanisms that can lead to spatial self-organisation of vegetation, as it has been shown with models representing plants by a density of biomass, varying continuously in time and space. However, although plants are usually quite plastic they also display discrete qualities and stochastic behaviour. These features may give rise to demographic noise, which in certain cases can influence the qualitative dynamics of ecosystem models. In the pre…
In contrast to the neutral population cycles of the deterministic mean-field Lotka--Volterra rate equations, including spatial structure and stochastic noise in models for predator-prey interactions yields complex spatio-temporal structures associated with long-lived erratic population oscillations. Environmental variability in the form of quenched spatial randomness in the predation rates results in more localized activity patches. Population fluctuations in rare favorable regions in turn cause a remarkable increase in the asymptotic densities of both predators and prey. Very intriguing featu…
This paper is concerned with a reaction--diffusion system modeling the fixation and the invasion in a population of a gene drive (an allele biasing inheritance, increasing its own transmission to offspring). In our model, the gene drive has a negative effect on the fitness of individuals carrying it, and is therefore susceptible of decreasing the total carrying capacity of the population locally in space. This tends to generate an opposing demographic advection that the gene drive has to overcome in order to invade. While previous reaction--diffusion models neglected this aspect, here we focus…
We study the effects of demographic stochasticity on the long-term dynamics of biological coevolution models of community assembly. The noise is induced in order to check the validity of deterministic population dynamics. While mutualistic communities show little dependence on the stochastic population fluctuations, predator-prey models show strong dependence on the stochasticity, indicating the relevance of the finiteness of the populations. For a predator-prey model, the noise causes drastic decreases in diversity and total population size. The communities that emerge under influence of the …
Logistic dynamics of small populations with demographic stochasticity
Consejo Nacional de Investigaciones Científicas y Técnicas（国家科学研究委员会）；Centro Atómico Bariloche and Instituto Balseiro, Comisión Nacional de Energía Atómica and Universidad Nacional de Cuyo（巴里洛切原子中心与巴尔塞罗研究所，国家原子能委员会与门多萨国立大学）
Frequency dependent selection and demographic fluctuations play important roles in evolutionary and ecological processes. Under frequency dependent selection, the average fitness of the population may increase or decrease based on interactions between individuals within the population. This should be reflected in fluctuations of the population size even in constant environments. Here, we propose a stochastic model, which naturally combines these two evolutionary ingredients by assuming frequency dependent competition between different types in an individual-based model. In contrast to previous…
Community ecology has traditionally relied on the competitive exclusion principle, a piece of common wisdom in conceptual frameworks developed to describe species assemblages. Key concepts in community ecology, such as limiting similarity and niche partitioning, are based on competitive exclusion. However, this classical paradigm in ecology relies on implications derived from simple, deterministic models. Here we show how the predictions of a symmetric, deterministic model about the way extinctions proceed can be utterly different from the results derived from the same model when ecological dr…
We describe a continuous-time modelling framework for biological population dynamics that accounts for demographic noise. In the spirit of the methodology used by statistical physicists, transitions between the states of the system are caused by individual events while the dynamics are described in terms of the time-evolution of a probability density function. In general, the application of the diffusion approximation still leaves a description that is quite complex. However, in many biological applications one or more of the processes happen slowly relative to the system's other processes, an…
Stochasticity can play an important role in the dynamics of biologically relevant populations. These span a broad range of scales: from intra-cellular populations of molecules to population of cells and then to groups of plants, animals and people. Large deviations in stochastic population dynamics - such as those determining population extinction, fixation or switching between different states - are presently in a focus of attention of statistical physicists. We review recent progress in applying different variants of dissipative WKB approximation (after Wentzel, Kramers and Brillouin) to thi…
Stochastic Pharmacokinetic Escape: A Field-Theoretic Approach to Fluctuation-Induced Tumor Relapse
Physics Engineering Department, Universidad de Guanajuato（瓜纳华托大学物理工程学院）；Physics Department, Universidad de Guanajuato（瓜纳华托大学物理系）；De Duve Institute, Université Catholique de Louvain（杜维研究所，天主教鲁汶大学）
The "Kill the Winner" hypothesis is an attempt to address the problem of diversity in biology. It argues that host-specific predators control the population of each prey, preventing a winner from emerging and thus maintaining the coexistence of all species in the system. We develop a stochastic model for the "Kill the Winner" paradigm and show that the stable coexistence state of the deterministic "Kill the Winner" model is destroyed by demographic stochasticity, through a cascade of extinction events. We formulate an individual-level stochastic model in which predator-prey coevolution promote…
We study the role of demographic fluctuations in typical endemics as exemplified by the stochastic SIRS model. The birth-death master equation of the model is simulated using exact numerics and analysed within the linear noise approximation. The endemic fixed point is unstable to internal demographic noise, and leads to sustained oscillations. This is ensured when the eigenvalues ($λ$) of the linearised drift matrix are complex, which in turn, is possible only if detailed balance is violated. In the oscillatory state, the phases decorrelate asymptotically, distinguishing such oscillations from…
Identifying the critical domain size necessary for a population to persist is an important question in ecology. Both demographic and environmental stochasticity impact a population's ability to persist. Here we explore ways of including this variability. We study populations which have traditionally been modelled using a deterministic integrodifference equation (IDE) framework, with distinct dispersal and sedentary stages. Individual based models (IBMs) are the most intuitive stochastic analogues to IDEs but yield few analytic insights. We explore two alternate approaches; one is a scaling up …
Demographic noise has profound effects on evolutionary and population dynamics, as well as on chemical reaction systems and models of epidemiology. Such noise is intrinsic and due to the discreteness of the dynamics in finite populations. We here show that similar noise-sustained trajectories arise in game dynamical learning, where the stochasticity has a different origin: agents sample a finite number of moves of their opponents in-between adaptation events. The limit of infinite batches results in deterministic modified replicator equations, whereas finite sampling leads to a stochastic dyna…
