Population Dynamics in a Changing Environment: Random versus Periodic Switching.

Environmental changes greatly influence the evolution of populations. Here, we study the dynamics of a population of two strains, one growing slightly faster than the other, competing for resources in a time-varying binary environment modeled by a carrying capacity switching either randomly or periodically between states of abundance and scarcity. The population dynamics is characterized by demographic noise (birth and death events) coupled to a varying environment. We elucidate the similarities and differences of the evolution subject to a stochastically and periodically varying environment. Importantly, the population size distribution is generally found to be broader under intermediate and fast random switching than under periodic variations, which results in markedly different asymptotic behaviors between the fixation probability of random and periodic switching. We also determine the detailed conditions under which the fixation probability of the slow strain is maximal.

Fixation properties of rock-paper-scissors games in fluctuating populations.

Rock-paper-scissors games metaphorically model cyclic dominance in ecology and microbiology. In a static environment, these models are characterized by fixation probabilities obeying two different "laws" in large and small well-mixed populations. Here, we investigate the evolution of these three-species models subject to a randomly switching carrying capacity modeling the endless change between states of resources scarcity and abundance. Focusing mainly on the zero-sum rock-paper-scissors game, equivalent to the cyclic Lotka-Volterra model, we study how the coupling of demographic and environmental noise influences the fixation properties. More specifically, we investigate which species is the most likely to prevail in a population of fluctuating size and how the outcome depends on the environmental variability. We show that demographic noise coupled with environmental randomness "levels the field" of cyclic competition by balancing the effect of selection. In particular, we show that fast switching effectively reduces the selection intensity proportionally to the variance of the carrying capacity. We determine the conditions under which new fixation scenarios arise, where the most likely species to prevail changes with the rate of switching and the variance of the carrying capacity. Random switching has a limited effect on the mean fixation time that scales linearly with the average population size. Hence, environmental randomness makes the cyclic competition more egalitarian, but does not prolong the species coexistence. We also show how the fixation probabilities of close-to-zero-sum rock-paper-scissors games can be obtained from those of the zero-sum model by rescaling the selection intensity.

Evolution of a Fluctuating Population in a Randomly Switching Environment.

Environment plays a fundamental role in the competition for resources, and hence in the evolution of populations. Here, we study a well-mixed, finite population consisting of two strains competing for the limited resources provided by an environment that randomly switches between states of abundance and scarcity. Assuming that one strain grows slightly faster than the other, we consider two scenarios-one of pure resource competition, and one in which one strain provides a public good-and investigate how environmental randomness (external noise) coupled to demographic (internal) noise determines the population's fixation properties and size distribution. By analytical means and simulations, we show that these coupled sources of noise can significantly enhance the fixation probability of the slower-growing species. We also show that the population size distribution can be unimodal, bimodal, or multimodal and undergoes noise-induced transitions between these regimes when the rate of switching matches the population's growth rate.

Cooperation dilemma in finite populations under fluctuating environments.

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 probability that cooperators fixate the population and replace all defectors. We analytically and numerically demonstrate that extrinsic noise may drastically enhance the cooperation fixation probability and even change its functional dependence on the population size. These results, which generalize earlier works in population genetics, indicate that extrinsic noise may help sustain and promote a much higher level of cooperation than static settings.

Mobility promotes and jeopardizes biodiversity in rock-paper-scissors games.

Biodiversity is essential to the viability of ecological systems. Species diversity in ecosystems is promoted by cyclic, non-hierarchical interactions among competing populations. Central features of such non-transitive relations are represented by the 'rock-paper-scissors' game, in which rock crushes scissors, scissors cut paper, and paper wraps rock. In combination with spatial dispersal of static populations, this type of competition results in the stable coexistence of all species and the long-term maintenance of biodiversity. However, population mobility is a central feature of real ecosystems: animals migrate, bacteria run and tumble. Here, we observe a critical influence of mobility on species diversity. When mobility exceeds a certain value, biodiversity is jeopardized and lost. In contrast, below this critical threshold all subpopulations coexist and an entanglement of travelling spiral waves forms in the course of time. We establish that this phenomenon is robust; it does not depend on the details of cyclic competition or spatial environment. These findings have important implications for maintenance and temporal development of ecological systems and are relevant for the formation and propagation of patterns in microbial populations or excitable media.

Coupled environmental and demographic fluctuations shape the evolution of cooperative antimicrobial resistance.

There is a pressing need to better understand how microbial populations respond to antimicrobial drugs, and to find mechanisms to possibly eradicate antimicrobial-resistant cells. The inactivation of antimicrobials by resistant microbes can often be viewed as a cooperative behaviour leading to the coexistence of resistant and sensitive cells in large populations and static environments. This picture is, however, greatly altered by the fluctuations arising in volatile environments, in which microbial communities commonly evolve. Here, we study the eco-evolutionary dynamics of a population consisting of an antimicrobial-resistant strain and microbes sensitive to antimicrobial drugs in a time-fluctuating environment, modelled by a carrying capacity randomly switching between states of abundance and scarcity. We assume that antimicrobial resistance (AMR) is a shared public good when the number of resistant cells exceeds a certain threshold. Eco-evolutionary dynamics is thus characterised by demographic noise (birth and death events) coupled to environmental fluctuations which can cause population bottlenecks. By combining analytical and computational means, we determine the environmental conditions for the long-lived coexistence and fixation of both strains, and characterise a fluctuation-driven AMR eradication mechanism, where resistant microbes experience bottlenecks leading to extinction. We also discuss the possible applications of our findings to laboratory-controlled experiments.

Metastability and anomalous fixation in evolutionary games on scale-free networks.

We study the influence of complex graphs on the metastability and fixation properties of a set of evolutionary processes. In the framework of evolutionary game theory, where the fitness and selection are frequency dependent and vary with the population composition, we analyze the dynamics of snowdrift games (characterized by a metastable coexistence state) on scale-free networks. Using an effective diffusion theory in the weak selection limit, we demonstrate how the scale-free structure affects the system's metastable state and leads to anomalous fixation. In particular, we analytically and numerically show that the probability and mean time of fixation are characterized by stretched-exponential behaviors with exponents depending on the network's degree distribution.

Effect of mobility in the rock-paper-scissor dynamics with high mortality.

In the evolutionary dynamics of a rock-paper-scissor model, the effect of natural death plays a major role in determining the fate of the system. Coexistence, being an unstable fixed point of the model, becomes very sensitive toward this parameter. In order to study the effect of mobility in such a system which has explicit dependence on mortality, we perform Monte Carlo simulation on a two-dimensional lattice having three cyclically competing species. The spatiotemporal dynamics has been studied along with the two-site correlation function. Spatial distribution exhibits emergence of spiral patterns in the presence of mobility. It reveals that the joint effect of death rate and mobility (diffusion) leads to new coexistence and extinction scenarios.

Fixation of a deleterious allele under mutation pressure and finite selection intensity.

The mean fixation time of a deleterious mutant allele is studied beyond the diffusion approximation. As in Kimura's classical work [M. Kimura, Proc. Natl. Acad. Sci. USA. 77, 522 (1980)], that was motivated by the problem of fixation in the presence of amorphic or hypermorphic mutations, we consider a diallelic model at a single locus comprising a wild-type A and a mutant allele A' produced irreversibly from A at small uniform rate v. The relative fitnesses of the mutant homozygotes A'A', mutant heterozygotes A'A and wild-type homozygotes AA are 1-s, 1-h and 1, respectively, where it is assumed that v<0) and discuss three situations: when the mutant is (i) completely dominant (s=h); (ii) completely recessive (h=0), and (iii) semi-dominant (h=s/2). Our theoretical predictions for the mean fixation time and the quasi-stationary distribution of the mutant population in the coexistence state, are shown to be in excellent agreement with numerical simulations. Furthermore, when s is finite, we demonstrate that our results are superior to those of the diffusion theory, while the latter is shown to be an accurate approximation only when N(e)s(2)<<1, where N(e) is the effective population size.

Exclusion of the fittest predicts microbial community diversity in fluctuating environments.

Microorganisms live in environments that inevitably fluctuate between mild and harsh conditions. As harsh conditions may cause extinctions, the rate at which fluctuations occur can shape microbial communities and their diversity, but we still lack an intuition on how. Here, we build a mathematical model describing two microbial species living in an environment where substrate supplies randomly switch between abundant and scarce. We then vary the rate of switching as well as different properties of the interacting species, and measure the probability of the weaker species driving the stronger one extinct. We find that this probability increases with the strength of demographic noise under harsh conditions and peaks at either low, high, or intermediate switching rates depending on both species' ability to withstand the harsh environment. This complex relationship shows why finding patterns between environmental fluctuations and diversity has historically been difficult. In parameter ranges where the fittest species was most likely to be excluded, however, the beta diversity in larger communities also peaked. In sum, how environmental fluctuations affect interactions between a few species pairs predicts their effect on the beta diversity of the whole community.

How does homophily shape the topology of a dynamic network?

We consider a dynamic network of individuals that may hold one of two different opinions in a two-party society. As a dynamical model, agents can endlessly create and delete links to satisfy a preferred degree, and the network is shaped by homophily, a form of social interaction. Characterized by the parameter J∈[-1,1], the latter plays a role similar to Ising spins: agents create links to others of the same opinion with probability (1+J)/2 and delete them with probability (1-J)/2. Using Monte Carlo simulations and mean-field theory, we focus on the network structure in the steady state. We study the effects of J on degree distributions and the fraction of cross-party links. While the extreme cases of homophily or heterophily (J=±1) are easily understood to result in complete polarization or anti-polarization, intermediate values of J lead to interesting features of the network. Our model exhibits the intriguing feature of an "overwhelming transition" occurring when communities of different sizes are subject to sufficient heterophily: agents of the minority group are oversubscribed and their average degree greatly exceeds that of the majority group. In addition, we introduce an original measure of polarization which displays distinct advantages over the commonly used average edge homogeneity.

Oscillatory dynamics in rock-paper-scissors games with mutations.

We study the oscillatory dynamics in the generic three-species rock-paper-scissors games with mutations. In the mean-field limit, different behaviors are found: (a) for high mutation rate, there is a stable interior fixed point with coexistence of all species; (b) for low mutation rates, there is a region of the parameter space characterized by a limit cycle resulting from a Hopf bifurcation; (c) in the absence of mutations, there is a region where heteroclinic cycles yield oscillations of large amplitude (not robust against noise). After a discussion on the main properties of the mean-field dynamics, we investigate the stochastic version of the model within an individual-based formulation. Demographic fluctuations are therefore naturally accounted and their effects are studied using a diffusion theory complemented by numerical simulations. It is thus shown that persistent erratic oscillations (quasi-cycles) of large amplitude emerge from a noise-induced resonance phenomenon. We also analytically and numerically compute the average escape time necessary to reach a (quasi-)cycle on which the system oscillates at a given amplitude.

Self-organization of mobile populations in cyclic competition.

The formation of out-of-equilibrium patterns is a characteristic feature of spatially extended, biodiverse, ecological systems. Intriguing examples are provided by cyclic competition of species, as metaphorically described by the 'rock-paper-scissors' game. Both experimentally and theoretically, such non-transitive interactions have been found to induce self-organization of static individuals into noisy, irregular clusters. However, a profound understanding and characterization of such patterns is still lacking. Here, we theoretically investigate the influence of individuals' mobility on the spatial structures emerging in rock-paper-scissors games. We devise a quantitative approach to analyze the spatial patterns self-forming in the course of the stochastic time evolution. For a paradigmatic model originally introduced by May and Leonard, within an interacting particle approach, we demonstrate that the system's behavior-in the proper continuum limit-is aptly captured by a set of stochastic partial differential equations. The system's stochastic dynamics is shown to lead to the emergence of entangled rotating spiral waves. While the spirals' wavelength and spreading velocity is demonstrated to be accurately predicted by a (deterministic) complex Ginzburg-Landau equation, their entanglement results from the inherent stochastic nature of the system. These findings and our methods have important applications for understanding the formation of noisy patterns, e.g. in ecological and evolutionary contexts, and are also of relevance for the kinetics of (bio)-chemical reactions.

Survival behavior in the cyclic Lotka-Volterra model with a randomly switching reaction rate.

We study the influence of a randomly switching reproduction-predation rate on the survival behavior of the nonspatial cyclic Lotka-Volterra model, also known as the zero-sum rock-paper-scissors game, used to metaphorically describe the cyclic competition between three species. In large and finite populations, demographic fluctuations (internal noise) drive two species to extinction in a finite time, while the species with the smallest reproduction-predation rate is the most likely to be the surviving one (law of the weakest). Here we model environmental (external) noise by assuming that the reproduction-predation rate of the strongest species (the fastest to reproduce and predate) in a given static environment randomly switches between two values corresponding to more and less favorable external conditions. We study the joint effect of environmental and demographic noise on the species survival probabilities and on the mean extinction time. In particular, we investigate whether the survival probabilities follow the law of the weakest and analyze their dependence on the external noise intensity and switching rate. Remarkably, when, on average, there is a finite number of switches prior to extinction, the survival probability of the predator of the species whose reaction rate switches typically varies nonmonotonically with the external noise intensity (with optimal survival about a critical noise strength). We also outline the relationship with the case where all reaction rates switch on markedly different time scales.

Eco-evolutionary dynamics of a population with randomly switching carrying capacity.

Environmental variability greatly influences the eco-evolutionary dynamics of a population, i.e. it affects how its size and composition evolve. Here, we study a well-mixed population of finite and fluctuating size whose growth is limited by a randomly switching carrying capacity. This models the environmental fluctuations between states of resources abundance and scarcity. The population consists of two strains, one growing slightly faster than the other, competing under two scenarios: one in which competition is solely for resources, and one in which the slow (cooperating) strain produces a public good (PG) that benefits also the fast (free-riding) strain. We investigate how the coupling of demographic and environmental (external) noise affects the population's eco-evolutionary dynamics. By analytical and computational means, we study the correlations between the population size and its composition, and discuss the social-dilemma-like 'eco-evolutionary game' characterizing the PG production. We determine in what conditions it is best to produce a PG; when cooperating is beneficial but outcompeted by free riding, and when the PG production is detrimental for cooperators. Within a linear noise approximation to populations of varying size, we also accurately analyse the coupled effects of demographic and environmental noise on the size distribution.

Heterogeneous out-of-equilibrium nonlinear q-voter model with zealotry.

We study the dynamics of the out-of-equilibrium nonlinear q-voter model with two types of susceptible voters and zealots, introduced in Mellor et al. [Europhys. Lett. 113, 48001 (2016)EULEEJ0295-507510.1209/0295-5075/113/48001]. In this model, each individual supports one of two parties and is either a susceptible voter of type q_{1} or q_{2}, or is an inflexible zealot. At each time step, a q_{i}-susceptible voter (i=1,2) consults a group of q_{i} neighbors and adopts their opinion if all group members agree, while zealots are inflexible and never change their opinion. This model violates detailed balance whenever q_{1}≠q_{2} and is characterized by two distinct regimes of low and high density of zealotry. Here, by combining analytical and numerical methods, we investigate the nonequilibrium stationary state of the system in terms of its probability distribution, nonvanishing currents, and unequal-time two-point correlation functions. We also study the switching time properties of the model by exploiting an approximate mapping onto the model of Mobilia [Phys. Rev. E 92, 012803 (2015)PLEEE81539-375510.1103/PhysRevE.92.012803] that satisfies the detailed balance, and we outline some properties of the model near criticality.

Coexistence versus extinction in the stochastic cyclic Lotka-Volterra model.

Cyclic dominance of species has been identified as a potential mechanism to maintain biodiversity, see, e.g., B. Kerr, M. A. Riley, M. W. Feldman and B. J. M. Bohannan [Nature 418, 171 (2002)] and B. Kirkup and M. A. Riley [Nature 428, 412 (2004)]. Through analytical methods supported by numerical simulations, we address this issue by studying the properties of a paradigmatic non-spatial three-species stochastic system, namely, the "rock-paper-scissors" or cyclic Lotka-Volterra model. While the deterministic approach (rate equations) predicts the coexistence of the species resulting in regular (yet neutrally stable) oscillations of the population densities, we demonstrate that fluctuations arising in the system with a finite number of agents drastically alter this picture and are responsible for extinction: After long enough time, two of the three species die out. As main findings we provide analytic estimates and numerical computation of the extinction probability at a given time. We also discuss the implications of our results for a broad class of competing population systems.

Bottleneck-induced transitions in a minimal model for intracellular transport.

We consider the influence of disorder on the nonequilibrium steady state of a minimal model for intracellular transport. In this model particles move unidirectionally according to the totally asymmetric exclusion process (TASEP) and are coupled to a bulk reservoir by Langmuir kinetics. Our discussion focuses on localized point defects acting as a bottleneck for the particle transport. Combining analytic methods and numerical simulations, we identify a rich phase behavior as a function of the defect strength. Our analytical approach relies on an effective mean-field theory obtained by splitting the lattice into two subsystems, which are effectively connected exploiting the local current conservation. Introducing the key concept of a carrying capacity, the maximal current that can flow through the bulk of the system (including the defect), we discriminate between the cases where the defect is irrelevant and those where it acts as a bottleneck and induces various novel phases (called bottleneck phases). Contrary to the simple TASEP in the presence of inhomogeneities, many scenarios emerge and translate into rich underlying phase diagrams, the topological properties of which are discussed.

Fluctuations and correlations in lattice models for predator-prey interaction.

Including spatial structure and stochastic noise invalidates the classical Lotka-Volterra picture of stable regular population cycles emerging in models for predator-prey interactions. Growth-limiting terms for the prey induce a continuous extinction threshold for the predator population whose critical properties are in the directed percolation universality class. We discuss the robustness of this scenario by considering an ecologically inspired stochastic lattice predator-prey model variant where the predation process includes next-nearest-neighbor interactions. We find that the corresponding stochastic model reproduces the above scenario in dimensions 1< d < or =4, in contrast with the mean-field theory, which predicts a first-order phase transition. However, the mean-field features are recovered upon allowing for nearest-neighbor particle exchange processes, provided these are sufficiently fast.

Voting and catalytic processes with inhomogeneities.

We consider the dynamics of the voter model and of the monomer-monomer catalytic process in the presence of many "competing" inhomogeneities and show, through exact calculations and numerical simulations, that their presence results in a non-trivial fluctuating steady state whose properties are studied and turn out to specifically depend on the dimensionality of the system, the strength of the inhomogeneities, and their separating distances. In fact, in arbitrary dimensions, we obtain an exact (yet formal) expression of the order parameters (magnetization and concentration of adsorbed particles) in the presence of an arbitrary number n of inhomogeneities ("zealots" in the voter language) and formal similarities with suitable electrostatic systems are pointed out. In the non-trivial cases n = 1,2, we explicitly compute the static and long-time properties of the order parameters and therefore capture the generic features of the systems. When n > 2 , the problems are studied through numerical simulations. In one spatial dimension, we also compute the expressions of the stationary order parameters in the completely disordered case, where n is arbitrary large. Particular attention is paid to the spatial dependence of the stationary order parameters and formal connections with electrostatics.