Environmental noise can cause an exponential reduction in the mean time to extinction (MTE) of an isolated population. We study this effect on an example of a stochastic birth-death process with rates modulated by a colored Gaussian noise. A path integral formulation yields a transparent way of evaluating the MTE and finding the optimal realization of the environmental noise that determines the most probable path to extinction. The population-size dependence of the MTE changes from exponential in the absence of the environmental noise to a power law for a short-correlated noise and to no dependence for long-correlated noise. We also establish the validity domains of the limits of white noise and adiabatic noise.
We study evolutionary multi-player games in finite populations, subject to fluctuating environments. The population undergoes a birth-death process with absorbing states, and the environment follows a Markovian process, resulting in a fluctuating payoff matrix for the evolutionary game. Our focus is on the fixation or extinction of a single mutant in a population of wildtypes. We show that the nonlinear nature of fitnesses in multi-player games gives rise to an intricate interplay of selection, genetic drift and environmental fluctuations. This generates effects not seen in simpler two-player …
Many populations in nature are fragmented: they consist of local populations occupying separate patches. A local population is prone to extinction due to the shot noise of birth and death processes. A migrating population from another patch can dramatically delay the extinction. What is the optimal migration rate that minimizes the extinction risk of the whole population? Here we answer this question for a connected network of model habitat patches with different carrying capacities.
Extinction of a long-lived isolated stochastic population can be described as an exponentially slow decay of quasi-stationary probability distribution of the population size. We address extinction of a population in a two-population system in the case when the population turnover -- renewal and removal -- is much slower than all other processes. In this case there is a time scale separation in the system which enables one to introduce a short-time quasi-stationary extinction rate W_1 and a long-time quasi-stationary extinction rate W_2, and develop a time-dependent theory of the transition bet…
Established populations often exhibit oscillations in their sizes. If a population is isolated, intrinsic stochasticity of elemental processes can ultimately bring it to extinction. Here we study extinction of oscillating populations in a stochastic version of the Rosenzweig-MacArthur predator-prey model. To this end we extend a WKB approximation (after Wentzel, Kramers and Brillouin) of solving the master equation to the case of extinction from a limit cycle in the space of population sizes. We evaluate the extinction rates and find the most probable paths to extinction by applying Floquet th…
Isolated populations ultimately go extinct because of the intrinsic noise of elementary processes. In multi-population systems extinction of a population may occur via more than one route. We investigate this generic situation in a simple predator-prey (or infected-susceptible) model. The predator and prey populations may coexist for a long time but ultimately both go extinct. In the first extinction route the predators go extinct first, whereas the prey thrive for a long time and then also go extinct. In the second route the prey go extinct first causing a rapid extinction of the predators. A…
This paper deals with extinction of an isolated population caused by intrinsic noise. We model the population dynamics in a "refuge" as a Markov process which involves births and deaths on discrete lattice sites and random migrations between neighboring sites. In extinction scenario I the zero population size is a repelling fixed point of the on-site deterministic dynamics. In extinction scenario II the zero population size is an attracting fixed point, corresponding to what is known in ecology as Allee effect. Assuming a large population size, we develop WKB (Wentzel-Kramers-Brillouin) approx…
We consider a stochastic population model where the intrinsic or demographic noise causes cycling between states before the population eventually goes extinct. A master equation approach coupled with a WKB (Wentzel-Kramers-Brillouin) approximation is used to construct the optimal path to extinction. In addition, a probabilistic argument is used to understand the pre-extinction dynamics and approximate the mean time to extinction. Analytical results agree well with numerical Monte Carlo simulations. A control method is implemented to decrease the mean time to extinction. Analytical results quan…
Standard neutral population genetics theory with a strictly fixed population size has important limitations. An alternative model that allows independently fluctuating population sizes and reproduces the standard neutral evolution is reviewed. We then study a situation such that the competing species are neutral at the equilibrium population size but population size fluctuations nevertheless favor fixation of one species over the other. In this case, a separation of timescales emerges naturally and allows adiabatic elimination of a fast population size variable to deduce the fluctuations-induc…
Many types of bacteria can survive under stress by switching stochastically between two different phenotypes: the "normals" who multiply fast, but are vulnerable to stress, and the "persisters" who hardly multiply, but are resilient to stress. Previous theoretical studies of such bacterial populations have focused on the \emph{fitness}: the asymptotic rate of unbounded growth of the population. Yet for an isolated population of established (and not very large) size, a more relevant measure may be the population \emph{extinction risk} due to the interplay of adverse extrinsic variations and int…
We study the extinction risk of a fragmented population residing on a network of patches coupled by migration, where the local patch dynamics include the Allee effect. We show that mixing between patches dramatically influences the population's viability. Slow migration is shown to always increase the population's global extinction risk compared to the isolated case. At fast migration, we demonstrate that synchrony between patches minimizes the population's extinction risk. Moreover, we discover a critical migration rate that maximizes the extinction risk of the population, and identify an ear…
We study the combined impact of a colored environmental noise and demographic noise on the extinction risk of a long-lived and well-mixed isolated stochastic population which exhibits the Allee effect. The environmental noise modulates the population birth and death rates. Assuming that the Allee effect is strong, and the environmental noise is positively correlated and Gaussian, we derive a Fokker-Planck equation for the joint probability distribution of the population sizes and environmental fluctuations. In WKB approximation this equation reduces to an effective two-dimensional Hamiltonian …
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…
The extinction time of an isolated population can be exponentially reduced by a periodic modulation of its environment. We investigate this effect using, as an example, a stochastic branching-annihilation process with a time-dependent branching rate. The population extinction is treated in eikonal approximation, where it is described as an instanton trajectory of a proper reaction Hamiltonian. The modulation of the environment perturbs this trajectory and synchronizes it with the modulation phase. We calculate the corresponding change in the action along the instanton using perturbation techni…
How high should be the rate of immigration into a stochastic population in order to significantly reduce the probability of observing the population extinct? Is there any relation between the population size distributions with and without immigration? Under what conditions can one justify the simple patch occupancy models which ignore the population distribution and its dynamics in a patch, and treat a patch simply as either occupied or empty? We address these questions by exactly solving a simple stochastic model obtained by adding a steady immigration to a variant of the Verhulst model: a pr…
We consider non-demographic noise in the form of uncertainty in the reaction step size, and reveal a dramatic effect this noise may have on the stability of self-regulating populations. Employing the reaction scheme mA->kA, but allowing, e.g., the product number k to be a-priori unknown and sampled from a given distribution, we show that such non-demographic noise can greatly reduce the population's extinction risk compared to the fixed k case. Our analysis is tested against numerical simulations, and by using empirical data of different species, we argue that certain distributions may be more…
We investigate extinction of a long-lived self-regulating stochastic population, caused by intrinsic (demographic) noise. Extinction typically occurs via one of two scenarios depending on whether the absorbing state n=0 is a repelling (scenario A) or attracting (scenario B) point of the deterministic rate equation. In scenario A the metastable stochastic population resides in the vicinity of an attracting fixed point next to the repelling point n=0. In scenario B there is an intermediate repelling point n=n_1 between the attracting point n=0 and another attracting point n=n_2 in the vicinity o…
We investigate how a catastrophic event (modeled as a temporary fall of the reproduction rate) increases the extinction probability of an isolated self-regulated stochastic population. Using a variant of the Verhulst logistic model as an example, we combine the probability generating function technique with an eikonal approximation to evaluate the exponentially large increase in the extinction probability caused by the catastrophe. This quantity is given by the eikonal action computed over "the optimal path" (instanton) of an effective classical Hamiltonian system with a time-dependent Hamilto…
Evolutionary adaptation through bet-hedging in finite populations under fluctuating environments
Grupo Interdisciplinar de Sistemas Complejos (GISC) and Departamento de Estructura de la Materia, Física Térmica y Electrónica, Universidad Complutense de Madrid（马德里康普顿斯大学复杂系统跨学科小组及物质结构、热学与电子学系）
Many species are unsustainable at small population densities (Allee Effect), i.e., below a threshold named Allee threshold, the population decreases instead of growing. In a closed local population, environmental fluctuations always lead to extinction. Here, we show how, in spatially extended habitats, dispersal can lead to a sustainable population in a region, provided the amplitude of environmental fluctuations is below an extinction threshold. We have identified two types of sustainable populations: high-density and low-density populations (through a mean-field approximation, valid in the l…
We present a novel approach allowing the study of rare events like fixation under fluctuating environments, modeled as extrinsic noise, in evolutionary processes characterized by the dominance of one species. Our treatment consists of mapping the system onto an auxiliary model, exhibiting metastable species coexistence, that can be analyzed semiclassically. This approach enables us to study the interplay between extrinsic and demographic noise on the statistics of interest. We illustrate our theory by considering the paradigmatic prisoner's dilemma game whose evolution is described by the prob…
In recent years non-demographic variability has been shown to greatly affect dynamics of stochastic populations. For example, non-demographic noise in the form of a bursty reproduction process with an a-priori unknown burst size, or environmental variability in the form of time-varying reaction rates, have been separately found to dramatically impact the extinction risk of isolated populations. In this work we investigate the extinction risk of an isolated population under the combined influence of these two types of non-demographic variation. Using the so-called momentum-space WKB approach we…
Mathematical models of population extinction have a variety of applications in such areas as ecology, paleontology and conservation biology. Here we propose and investigate two types of sub-exponential models of population extinction.The results of performed analysis show that the principle of minimum of information loss is the underlying law for the evolution of a broad class of models of population extinction. Finally, we propose a possible application of this modeling framework to mechanisms underlying time perception.
Multiple-scale mobility is ubiquitous in nature and has become instrumental for understanding and modeling animal foraging behavior. However, the impact of individual movements on the long-term stability of populations remains largely unexplored. We analyze deterministic and stochastic Lotka Volterra systems, where mobile predators consume scarce resources (prey) confined in patches. In fragile systems (that is, those unfavorable to species coexistence), the predator species has a maximized abundance and is resilient to degraded prey conditions when individual mobility is multiple scaled. With…
We study a generic reaction-diffusion model for single-species population dynamics that includes reproduction, death, and competition. The population is assumed to be confined in a refuge beyond which conditions are so harsh that they lead to certain extinction. Standard continuum mean field models in one dimension yield a critical refuge length L_c such that a population in a refuge larger than this is assured survival. Herein we extend the model to take into account the discreteness and finiteness of the population, which leads us to a stochastic description. We present a particular critical…
We investigate the dynamics of the voter model in which the population itself changes endogenously via the birth-death process. There are two species of voters, labeled A and B, and the population of each species can grow or shrink by the birth-death process at equal rates $b$. Individuals of opposite species also undergo voter model dynamics in which an AB pair can equiprobably become AA or BB with rate $v$---neutral evolution. In the limit $b/v\to\infty$, the distribution of consensus times varies as $t^{-3}$ and the probability that the population size equals $n$ at the moment of consensus …
Competitive birth-death processes often exhibit an oscillatory behavior. We investigate a particular case where the oscillation cycles are marginally stable on the mean-field level. An iconic example of such a system is the Lotka-Volterra model of predator-prey competition. Fluctuation effects due to discreteness of the populations destroy the mean-field stability and eventually drive the system toward extinction of one or both species. We show that the corresponding extinction time scales as a certain power-law of the population sizes. This behavior should be contrasted with the extinction of…
Population genetics struggles to model extinction; standard models track the relative rather than absolute fitness of genotypes, while the exceptions describe only the short-term transition from imminent doom to evolutionary rescue. But extinction can result from failure to adapt not only to catastrophes, but also to a backlog of environmental challenges. We model long-term evolution to long series of small challenges, where fitter populations reach higher population sizes. The population's long-term fitness dynamic is well approximated by a simple stochastic Markov chain model. Long-term pers…
We consider control of reaction and population systems by deterministically imposed transitions between the states with different numbers of particles or individuals. Even where the imposed transitions are significantly less frequent than spontaneous transitions, they can exponentially strongly modify the rates of rare events, including switching between metastable states or population extinction. We also study optimal control of rare events, and specifically, optimal control of disease extinction for a limited vaccine supply. A comparison is made with control of rare events by modulating the …
Species extinction occurs regularly and unavoidably in ecological systems. The time scales for extinction can broadly vary and inform on the ecosystem's stability. We study the spatio-temporal extinction dynamics of a paradigmatic population model where three species exhibit cyclic competition. The cyclic dynamics reflects the non-equilibrium nature of the species interactions. While previous work focusses on the coarsening process as a mechanism that drives the system to extinction, we found that unexpectedly the dynamics to extinction is much richer. We observed three different types of dyna…
Game theory ideas provide a useful framework for studying evolutionary dynamics in a well-mixed environment. This approach, however, typically enforces a strictly fixed overall population size, deemphasizing natural growth processes. We study a competitive Lotka-Volterra model, with number fluctuations, that accounts for natural population growth and encompasses interaction scenarios typical of evolutionary games. We show that, in an appropriate limit, the model describes standard evolutionary games with both genetic drift and overall population size fluctuations. However, there are also regim…
Population extinction is a rare event which requires overcoming an effective barrier. We show that the extinction rate can be fragile: a small change in the system parameters leads to an exponentially strong change of the rate, with the barrier height depending on the parameters nonanalytically. General conditions of the fragility are established. The fragility is found in one of the best-known models of epidemiology, the SIS model. The analytical expressions are compared with simulations.
Theoretical ecologists have long sought to understand how the persistence of populations depends on biotic and abiotic factors. Classical work showed that demographic stochasticity causes the mean time to extinction to increase exponentially with population size, whereas variation in environmental conditions can lead to a power law scaling. Recent work has focused especially on the influence of the autocorrelation structure ("color") of environmental noise. In theoretical physics, there is a burst of research activity in analyzing large fluctuations in stochastic population dynamics. This rese…
We study inhomogeneous host-pathogen dynamics to model the global amphibian population extinction in a lake basin system. The lake basin system is modeled as quenched disorder. In this model we show that once the pathogen arrives at the lake basin it spreads from one lake to another, eventually spreading to the entire lake basin system in a wave like pattern. The extinction time has been found to depend on the steady state host population and pathogen growth rate. Linear estimate of the extinction time is computed. The steady state host population shows a threshold behavior in the interaction …
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…
Population dynamics reflects an underlying birth-death process, where the rates associated with different events may depend on external environmental conditions and on the population density. A whole family of simple and popular deterministic models (like logistic growth) support a transcritical bifurcation point between an extinction phase and an active phase. Here we provide a comprehensive analysis of the phases of that system, taking into account both the endogenous demographic noise (random birth and death events) and the effect of environmental stochasticity that causes variations in bir…
Extinction appears ubiquitously in many fields, including chemical reactions, population biology, evolution, and epidemiology. Even though extinction as a random process is a rare event, its occurrence is observed in large finite populations. Extinction occurs when fluctuations due to random transitions act as an effective force which drives one or more components or species to vanish. Although there are many random paths to an extinct state, there is an optimal path that maximizes the probability to extinction. In this article, we show that the optimal path is associated with the dynamical sy…
We study the joint effect of thermal bath fluctuations and an external noise tuning activity of cytotoxic cells on the triggered immune response in a growing cancerous tissue. The immune response is assumed to be primarily mediated by effector cells that develop a cytotoxic activity against the abnormal tissue. The kinetics of such a reaction is represented by an enzymatic-like Michaelis-Menten two step process. Effective free-energy surface for the process is further parametrized by the fluctuating energy barrier between the states of high and low concentration of cancerous cells. By analyzin…
Biological populations are subject to fluctuating environmental conditions. Different adaptive strategies can allow them to cope with these fluctuations: specialization to one particular environmental condition, adoption of a generalist phenotype that compromise between conditions, or population-wise diversification (bet-hedging). Which strategy provides the largest selective advantage in the long run depends on the range of accessible phenotypes and the statistics of the environmental fluctuations. Here, we analyze this problem in a simple mathematical model of population growth. First, we re…
We introduce a new model for plant metapopulations with a seed bank component, living in a fragmented environment in which local extinction events are frequent. This model is an intermediate between population dynamics models with a seed bank component, based on the classical Wright-Fisher model, and Stochastic Patch Occupancy Models (SPOMs) used in metapopulation ecology. Its main feature is the use of "ghost" individuals, which can reproduce but with a very strong selective disadvantage against "real" individuals, to artificially ensure a constant population size. We show the existence of an…
Understanding the mechanisms governing population extinctions is of key importance to many problems in ecology and evolution. Stochastic factors are known to play a central role in extinction, but the interactions between a population's demographic stochasticity and environmental noise remain poorly understood. Here, we model environmental forcing as a stochastic fluctuation between two states, one with a higher death rate than the other. We find that in general there exists a rate of fluctuations that minimizes the mean time to extinction, a phenomenon previously dubbed "resonant activation."…
We introduce a model of biological evolution where species evolve in response to biotic interactions and a fluctuating environmental stress. The species may either become extinct or mutate to acquire a new fitness value when the effective stress level is greater than their individual fitness. The model exhibits a phase transition to a completely extinct phase as the environmental stress or the mutation rate is varied. We discuss the generic conditions for which this transition is continuous. The model is exactly solvable and the critical behavior is characterized by an unusual dynamic exponent…
The question of whether a population will persist or go extinct is of key interest throughout ecology and biology. Various mathematical techniques allow us to generate knowledge regarding individual behaviour, which can be analysed to obtain predictions about the ultimate survival or extinction of the population. A common model employed to describe population dynamics is the lattice-based random walk model with crowding (exclusion). This model can incorporate behaviour such as birth, death and movement, while including natural phenomena such as finite size effects. Performing sufficiently many…
Motivated by modeling the dynamics of a population living in a flowing medium where the environmental factors are random in space, we have studied an asymmetric variant of the one-dimensional contact process, where the quenched random reproduction rates are systematically greater in one direction than in the opposite one. The spatial disorder turns out to be a relevant perturbation but, according to results of Monte Carlo simulations, the behavior of the model at the extinction transition is different from the (infinite randomness) critical behavior of the disordered, symmetric contact process…
The environment in which a population evolves can have a crucial impact on selection. We study evolutionary dynamics in finite populations of fixed size in a changing environment. The population dynamics are driven by birth and death events. The rates of these events may vary in time depending on the state of the environment, which follows an independent Markov process. We develop a general theory for the fixation probability of a mutant in a population of wild-types, and for mean unconditional and conditional fixation times. We apply our theory to evolutionary games for which the payoff struc…
Species extinction is a core process that affects the diversity of life on Earth. Competition between species in a population is considered by ecological niche-based theories as a key factor leading to different severity of species extinctions. There are population dynamics models that describe a simple and easily understandable mechanism for resource competition. However, these models can not efficiently characterize and quantify new emergent extinctions in a large population appearing due to environmental forcing. To address this issue we develop a stochastic physics-inspired approach to ana…
Time-dependent distribution of the global extinction of megafauna is compared with the growth of human population. There is no correlation between the two processes. Furthermore, the size of human population and its growth rate were far too small to have any significant impact on the environment and on the life of megafauna.
In this work we suggest and consider an original, simple mathematical model of a "quasi-rapid" extinction population dynamics. It describes a decrease and final extinction of the population of one prey species by a "quasi-rapid" interaction with one predator species with increasing population. This "quasi-rapid" interaction means ecologically that prey species behaves practically quite passively (since there is no time for any reaction, i.e. defense), like an appropriate environment, in respect to "quasi-rapid" activity of the predator species that can have different "quasi-rapid" hunting abil…
Recently, a first step was made by the authors towards a systematic investigation of the effect of reaction-step-size noise - uncertainty in the step size of the reaction - on the dynamics of stochastic populations. This was done by investigating the effect of bursty influx on the switching dynamics of stochastic populations. Here we extend this formalism to account for bursty reproduction processes, and improve the accuracy of the formalism to include subleading-order corrections. Bursty reproduction appears in various contexts, where notable examples include bursty viral production from infe…
Computer simulations carried out by Alroy (2001) are examined. Contrary to the claim of their Author, there is no convincing evidence that the extinction of megafauna was caused by humans. Intentionally or unintentionally, attempt was made to force the human-induced extinction of megafauna by assuming an absurdly fast growth of the hypothetical human population in North America. The assumed growth rate was around two-orders of magnitude larger than normally expected. It is well known that the past growth of human population was slow, but in these simulations, the growth of the hypothetical hum…
Most infectious diseases including more than half of known human pathogens are not restricted to just one host, yet much of the mathematical modeling of infections has been limited to a single species. We investigate consequences of a single epidemic propagating in multiple species and compare and contrast it with the endemic steady state of the disease. We use the two-species Susceptible-Infected-Recovered (SIR) model to calculate the severity of post-epidemic collapses in populations of two host species as a function of their initial population sizes, the times individuals remain infectious,…
Many mathematical frameworks of evolutionary game dynamics assume that the total population size is constant and that selection affects only the relative frequency of strategies. Here, we consider evolutionary game dynamics in an extended Wright-Fisher process with variable population size. In such a scenario, it is possible that the entire population becomes extinct. Survival of the population may depend on which strategy prevails in the game dynamics. Studying cooperative dilemmas, it is a natural feature of such a model that cooperators enable survival, while defectors drive extinction. Alt…
Temporal environmental variations are ubiquitous in nature, yet most of the theoretical works in population genetics and evolution assume fixed environment. Here we analyze the effect of variations in carrying capacity on the fate of a mutant type. We consider a two-state Moran model, where selection intensity at equilibrium may differ (in amplitude and in sign) from selection during periods of sharp growth and sharp decline. Using Kimura's diffusion approximation we present simple formulae for effective population size and effective selection, and use it to calculate the chance of ultimate fi…
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…
