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Antimicrobial resistance (AMR) is a global threat and combating its spread is of paramount importance. AMR often results from a cooperative behaviour with shared protection against drugs. Microbial communities generally evolve in volatile environments and spatial structures. Migration, fluctuations, and environmental variability thus have significant impacts on AMR, whose maintenance in static environments is generally promoted by migration. Here, we demonstrate that this picture changes dramatically in time-fluctuating spatially structured environments. To this end, we consider a two-dimensional metapopulation model consisting of demes in which drug-resistant and sensitive cells evolve in a time-changing environment in the presence of a toxin against which protection can be shared. Cells migrate between neighbouring demes and hence connect them. When the environment varies neither too quickly nor too slowly, the dynamics is characterised by bottlenecks causing fluctuation-driven local extinctions, a mechanism countered by migration that rescues AMR. Through simulations and mathematical analysis, we investigate how migration and environmental variability influence the probability of resistance eradication. We determine the near-optimal conditions for the fluctuation-driven AMR eradication, and show that slow but non-zero migration speeds up the clearance of resistance and can enhance its eradication probability. We discuss our study's impact on laboratory-controlled experiments., Comment: 24+9 pages, 8+3 figures, 1 table. Simulation data and codes for all figures, as well as all 5 Supplementary Movies, are electronically available from the Open Science Framework (OSF) repository. DOI: https://doi.org/10.17605/OSF.IO/EPB28

There is mounting evidence that species interactions often involve long-term memory, with highly-varying waiting times between successive events and long-range temporal correlations. Accounting for memory undermines the common Markovian assumption, and dramatically impacts key ingredients of population dynamics including birth, foraging, predation, and competition processes. Here, we investigate a critical aspect of population dynamics, namely non-Markovian multi-species competition. This is done in the realm of the zero-sum rock-paper-scissors (zRPS) model that is broadly used in the life sciences to metaphorically describe cyclic competition between three interacting species. We develop a general non-Markovian formalism for multi-species dynamics, allowing us to determine the regions of the parameter space where each species dominates. In particular, when the dynamics are Markovian, the waiting times are exponentially distributed and the fate of the zRPS model in large well-mixed populations follows the so-called ``law of the weakest'' (LOW), predicting that the species with the lowest growth rate is the most likely to prevail. We demonstrate that the survival behavior and LOW of the zRPS model are critically affected by non-exponential waiting time distributions, and especially, by their coefficient of variation. Our findings provide key insight into the influence of long waiting times on non-Markovian evolutionary processes., Comment: 10 pages, 5 figures + appendices

Antimicrobial resistance to drugs (AMR), a global threat to human and animal health, is often regarded as resulting from cooperative behaviour. Moreover, microbes generally evolve in volatile environments that, together with demographic fluctuations (birth and death events), drastically alter population size and strain survival. Motivated by the need to better understand the evolution of AMR, we study a population of time-varying size consisting of two competing strains, one drug-resistant and one drug-sensitive, subject to demographic and environmental variability. This is modelled by a binary carrying capacity randomly switching between mild and harsh environmental conditions, and driving the fluctuating volume (total amount of nutrients and antimicrobials at fixed concentration), and thus the size of the community (number of resistant and sensitive cells). We assume that AMR is a shared public good when the concentration of resistant cells exceeds a fixed concentration cooperation threshold, above which the sensitive strain has a growth advantage, whereas resistant cells dominate below it. Using computational means, and devising an analytical treatment (built on suitable quenched and annealed averaging procedures), we fully characterise the influence of fluctuations on the eco-evolutionary dynamics of AMR, and notably obtain specific strain fixation and long-lasting coexistence probabilities as a function of the environmental variation rate and cooperation threshold. We find that microbial strains tend to coexistence, but demographic fluctuations eventually lead to the extinction of resistant or sensitive cells for small or large values of the concentration cooperation threshold, respectively. This also holds for dynamic environments, whose specific properties determine the extinction timescale., Comment: 34+5 pages, 5 figures. Simulation data and codes for all figures are electronically available from the University of Leeds Data Repository. DOI: https://doi.org/10.5518/1462. In this version: minor typos corrected

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 behavior 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, modeled by a carrying capacity randomly switching between states of abundance and scarcity. We assume that antimicrobial resistance 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 antimicrobial resistance eradication mechanism, where resistant microbes experience bottlenecks leading to extinction. We also discuss the possible applications of our findings to laboratory-controlled experiments., Comment: 21+7 pages, 4+1 figures. The revised version includes minor corrections to the original manuscript. Simulation data and codes for all figures are electronically available from the University of Leeds Data Repository. DOI: https://doi.org/10.5518/1360

Microbial populations generally evolve in volatile environments, under conditions fluctuating between harsh and mild, e.g. as the result of sudden changes in toxin concentration or nutrient abundance. Environmental variability thus shapes the long-time population dynamics, notably by influencing the ability of different strains of microorganisms to coexist. Inspired by the evolution of antimicrobial resistance, we study the dynamics of a community consisting of two competing strains subject to twofold environmental variability. The level of toxin varies in time, favouring the growth of one strain under low drug concentration and the other strain when the toxin level is high. We also model time-changing resource abundance by a randomly switching carrying capacity that drives the fluctuating size of the community. While one strain dominates in a static environment, we show that species coexistence is possible in the presence of environmental variability. By computational and analytical means, we determine the environmental conditions under which long-lived coexistence is possible and when it is almost certain. Notably, we study the circumstances under which environmental and demographic fluctuations promote, or hinder, the strains coexistence. We also determine how the make-up of the coexistence phase and the average abundance of each strain depend on the environmental variability., Comment: 15 pages + appendix of 10 pages (Supplementary Material); Figshare resources: https://doi.org/10.6084/m9.figshare.23553066

We study the effect of time-fluctuating social influences on the formation of polarization and consensus in a three-party community consisting of two types of voters ("leftists" and "rightists") holding extreme opinions, and moderate agents acting as "centrists". The former are incompatible and do not interact, while centrists hold an intermediate opinion and can interact with extreme voters. When a centrist and a leftist/rightist interact, they can become either both centrists or both leftists/rightists. The population eventually either reaches consensus with one of the three opinions, or a polarization state consisting of a frozen mixture of leftists and rightists. As a main novelty, here agents interact subject to time-fluctuating external influences favouring in turn the spread of leftist and rightist opinions, or the rise of centrism. We determine the fate of the population under various scenarios, and show how the rate of change of external influences can drastically affect the polarization and consensus probabilities, as well as the mean time to reach the final state., Comment: 18 pages, 6 figures, 3 appendices

Environmental variations can significantly influence how populations compete for resources, and hence shape their evolution. Here, we study population dynamics subject to a fluctuating environment modeled by a varying carrying capacity changing continuously in time according to either binary random switches, or by being driven by a noise of continuous range. We consider a prototypical example of two competing strains, one growing slightly slower than the other, and consider also the scenario where the slow strain is a public goods producer. By systematically comparing the effect of binary- versus continuously-varying environment, we study how different noise statistics (mean, variance) influence the population size and fixation properties. We show that the slow strain fixation probability can be greatly enhanced for a continuously-varying environment compared to binary switches, even when the first two moments of the carrying capacity coincide., Comment: 6 pages, 3 figures + Supplemental Material

In the evolutionary dynamics of a rock-paper-scissor (RPS) 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 towards 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 $2$-dimensional lattice having three cyclically-competing species. The spatio-temporal dynamics has been studied along with the two-site correlation function. Spatial distribution exhibits emergence of spiral patterns in the presence of diffusion. It reveals that the joint effect of death rate and diffusion (migration) leads to new coexistence and extinction scenarios., Comment: 7 pages, 5 figures

We investigate the long-time properties of a dynamic, out-of-equilibrium network of individuals holding one of two opinions in a population consisting of two communities of different sizes. Here, while the agents' opinions are fixed, they have a preferred degree which leads them to endlessly create and delete links. Our evolving network is shaped by homophily/heterophily, which is a form of social interaction by which individuals tend to establish links with others having similar/dissimilar opinions. Using Monte Carlo simulations and a detailed mean-field analysis, we study in detail how the sizes of the communities and the degree of homophily/heterophily affects the network structure. In particular, we show that when the network is subject to enough heterophily, an "overwhelming transition" occurs: individuals of the smaller community are overwhelmed by links from agents of the larger group, and their mean degree greatly exceeds the preferred degree. This and related phenomena are characterized by obtaining the network's total and joint degree distributions, as well as the fraction of links across both communities and that of agents having fewer edges than the preferred degree. We use our mean-field theory to discuss the network's polarization when the group sizes and level of homophily vary., Comment: 24 pages, 10 figures

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 \emph{homophily}, a form of social interaction. Characterized by the parameter $J \in [-1,1]$, 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 ($J= \pm 1$) 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., Comment: The paper includes 12 pages and 9 figures

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., Comment: 64 pages, 28 figures, 3 tables

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., Comment: 22 pages, 7 figures: main text (6 pages, 3 figures) followed by Supplementary Material (16 pages, 4 figures). Published in Physical Review Letters. Additional supporting resources available at https://figshare.com/articles/Supplementary_Material/12613370

There is mounting empirical evidence that many communities of living organisms display key features which closely resemble those of physical systems at criticality. We here introduce a minimal model framework for the dynamics of a community of individuals which undergoes local birth-death, immigration and local jumps on a regular lattice. We study its properties when the system is close to its critical point. Even if this model violates detailed balance, within a physically relevant regime dominated by fluctuations, it is possible to calculate analytically the probability density function of the number of individuals living in a given volume, which captures the close-to-critical behavior of the community across spatial scales. We find that the resulting distribution satisfies an equation where spatial effects are encoded in appropriate functions of space, which we calculate explicitly. The validity of the analytical formul{\ae} is confirmed by simulations in the expected regimes. We finally discuss how this model in the critical-like regime is in agreement with several biodiversity patterns observed in tropical rain forests., Comment: To appear in Physical Review X. Our manuscript contains a summary paragraph of 168 words and a main text 8,473 words long, 9 figures (comprising 26 panels in total) with legends (1,266 total words) and 53 references

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 ${\it coupling}$ 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., Comment: 31 pages, 15 figures: Main text (18 pages, 9 figures) followed by Supplementary Material (13 pages, 6 figures). Supplementary Information and resources available at https://doi.org/10.6084/m9.figshare.8858273.v1

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 that benefits also the fast ("freeriding") 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 public good production. We determine in what conditions it is best to produce a public good; when cooperating is beneficial but outcompeted by freeriding, and when the public good production is detrimental for cooperators. Within a linear noise approximation to populations of varying size, we also accurately analyze the coupled effects of demographic and environmental noise on the size distribution., Comment: 25 pages, 15 figures: Main text (13 pages, 8 figures) followed by Supplementary Material (12 pages, 7 figures). To appear in J. R. Soc. Interface. Supplementary information and resources are also available at https://doi.org/10.6084/m9.figshare.5683762

We study the influence of a randomly switching reproduction-predation rate on the survival behavior of the non-spatial 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/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 of 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 non-monotonically 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., Comment: 15 pages, 10 figures

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., Comment: 15 pages, 8 figures: Main text (5+ pages, 4 figures) followed by Supplemental Material (10 pages, 4 supplementary figures). To appear in Physical Review Letters. Supporting mp4 and ogv videos are available at https://doi.org/10.6084/m9.figshare.5082712 and http://www1.maths.leeds.ac.uk/~amtmmo/KEM_videos/index.html

Spatially extended population dynamics models that incorporate intrinsic noise serve as case studies for the role of fluctuations and correlations in biological systems. Including spatial structure and stochastic noise in predator-prey competition invalidates the deterministic Lotka-Volterra picture of neutral population cycles. Stochastic models yield long-lived erratic population oscillations stemming from a resonant amplification mechanism. In spatially extended predator-prey systems, one observes noise-stabilized activity and persistent correlations. Fluctuation-induced renormalizations of the oscillation parameters can be analyzed perturbatively. The critical dynamics and the non-equilibrium relaxation kinetics at the predator extinction threshold are characterized by the directed percolation universality class. Spatial or environmental variability results in more localized patches which enhances both species densities. Affixing variable rates to individual particles and allowing for trait inheritance subject to mutations induces fast evolutionary dynamics for the rate distributions. Stochastic spatial variants of cyclic competition with rock-paper-scissors interactions illustrate connections between population dynamics and evolutionary game theory, and demonstrate how space can help maintain diversity. In two dimensions, three-species cyclic competition models of the May-Leonard type are characterized by the emergence of spiral patterns whose properties are elucidated by a mapping onto a complex Ginzburg-Landau equation. Extensions to general food networks can be classified on the mean-field level, which provides both a fundamental understanding of ensuing cooperativity and emergence of alliances. Novel space-time patterns emerge as a result of the formation of competing alliances, such as coarsening domains that each incorporate rock-paper-scissors competition games., Comment: Topical review, to appear in: J. Phys. A: Math. Theor.; 53+ pages, 16 figures

We study the influence of the complex topology of scale-free graphs on the dynamics of anti-coordination games (e.g. snowdrift games). These reference models are characterized by the coexistence (evolutionary stable mixed strategy) of two competing species, say "cooperators" and "defectors", and, in finite systems, by metastability and large-fluctuation-driven fixation. In this work, we use extensive computer simulations and an effective diffusion approximation (in the weak selection limit) to determine under which circumstances, depending on the individual-based update rules, the topology drastically affects the long-time behavior of anti-coordination games. In particular, we compute the variance of the number of cooperators in the metastable state and the mean fixation time when the dynamics is implemented according to the voter model (death-first/birth-second process) and the link dynamics (birth/death or death/birth at random). For the voter update rule, we show that the scale-free topology effectively renormalizes the population size and as a result the statistics of observables depend on the network's degree distribution. In contrast, such a renormalization does not occur with the link dynamics update rule and we recover the same behavior as on complete graphs., Comment: 21 pages, 8 figures, to appear in JSTAT

We study the dynamics of the out-of-equilibrium nonlinear q-voter model with two types of susceptible voters and zealots, introduced in [EPL 113, 48001 (2016)]. 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 $q_1 \neq q_2$ and is characterized by two distinct regimes of low and high density of zealotry. Here, by combining analytical and numerical methods, we investigate the non-equilibrium stationary state of the system in terms of its probability distribution, non-vanishing currents and unequal-time two-point correlation functions. We also study the switching times properties of the model by exploiting an approximate mapping onto the model of [Phys. Rev. E 92, 012803 (2015)] that satisfies the detailed balance, and also outline some properties of the model near criticality., Comment: 17 pages, 12 figures

We consider a two-dimensional model of three species in rock-paper-scissors competition and study the self-organisation of the population into fascinating spiraling patterns. Within our individual-based metapopulation formulation, the population composition changes due to cyclic dominance (dominance-removal and dominance-replacement), mutations, and pair-exchange of neighboring individuals. Here, we study the influence of mobility on the emerging patterns and investigate when the pair-exchange rate is responsible for spiral waves to become elusive in stochastic lattice simulations. In particular, we show that the spiral waves predicted by the system's deterministic partial equations are found in lattice simulations only within a finite range of the mobility rate. We also report that in the absence of mutations and dominance-replacement, the resulting spiraling patterns are subject to convective instability and far-field breakup at low mobility rate. Possible applications of these resolution and far-field breakup phenomena are discussed., Comment: 20 pages, 5 figures; published in Games

We introduce an heterogeneous nonlinear $q$-voter model with zealots and two types of susceptible voters, and study its non-equilibrium properties when the population is finite and well mixed. In this two-opinion model, each individual supports one of two parties and is either a zealot or a susceptible voter of type $q_1$ or $q_2$. While here zealots never change their opinion, a $q_i$-susceptible voter ($i=1,2$) consults a group of $q_i$ neighbors at each time step, and adopts their opinion if all group members agree. We show that this model violates the detailed balance whenever $q_1 \neq q_2$ and has surprisingly rich properties. Here, we focus on the characterization of the model's non-equilibrium stationary state (NESS) in terms of its probability distribution and currents in the distinct regimes of low and high density of zealotry. We unveil the NESS properties in each of these phases by computing the opinion distribution and the circulation of probability currents, as well as the two-point correlation functions at unequal times (formally related to a "probability angular momentum"). Our analytical calculations obtained in the realm of a linear Gaussian approximation are compared with numerical results., Comment: 6 pages, 2 figures, supplementary material and movie available at https://dx.doi.org/10.6084/m9.figshare.2060595

We study the dynamics of the nonlinear $q$-voter model with inflexible zealots in a finite well-mixed population. In this system, each individual supports one of two parties and is either a susceptible voter or an inflexible zealot. At each time step, a susceptible adopts the opinion of a neighbor if this belongs to a group of $q\geq 2$ neighbors all in the same state, whereas inflexible zealots never change their opinion. In the presence of zealots of both parties the model is characterized by a fluctuating stationary state and, below a zealotry density threshold, the distribution of opinions is bimodal. After a characteristic time, most susceptibles become supporters of the party having more zealots and the opinion distribution is asymmetric. When the number of zealots of both parties is the same, the opinion distribution is symmetric and, in the long run, susceptibles endlessly swing from the state where they all support one party to the opposite state. Above the zealotry density threshold, when there is an unequal number of zealots of each type, the probability distribution is single-peaked and non-Gaussian. These properties are investigated analytically and with stochastic simulations. We also study the mean time to reach a consensus when zealots support only one party., Comment: 11 pages, 10 figures. For Physical Review E (accepted, final version)

We generalize the classical Bass model of innovation diffusion to include a new class of agents --- Luddites --- that oppose the spread of innovation. Our model also incorporates ignorants, susceptibles, and adopters. When an ignorant and a susceptible meet, the former is converted to a susceptible at a given rate, while a susceptible spontaneously adopts the innovation at a constant rate. In response to the \emph{rate} of adoption, an ignorant may become a Luddite and permanently reject the innovation. Instead of reaching complete adoption, the final state generally consists of a population of Luddites, ignorants, and adopters. The evolution of this system is investigated analytically and by stochastic simulations. We determine the stationary distribution of adopters, the time needed to reach the final state, and the influence of the network topology on the innovation spread. Our model exhibits an important dichotomy: when the rate of adoption is low, an innovation spreads slowly but widely; in contrast, when the adoption rate is high, the innovation spreads rapidly but the extent of the adoption is severely limited by Luddites., Comment: 11 pages, 7 figures

Rock is wrapped by paper, paper is cut by scissors, and scissors are crushed by rock. This simple game is popular among children and adults to decide on trivial disputes that have no obvious winner, but cyclic dominance is also at the heart of predator-prey interactions, the mating strategy of side-blotched lizards, the overgrowth of marine sessile organisms, and the competition in microbial populations. Cyclical interactions also emerge spontaneously in evolutionary games entailing volunteering, reward, punishment, and in fact are common when the competing strategies are three or more regardless of the particularities of the game. Here we review recent advances on the rock-paper-scissors and related evolutionary games, focusing in particular on pattern formation, the impact of mobility, and the spontaneous emergence of cyclic dominance. We also review mean-field and zero-dimensional rock-paper-scissors models and the application of the complex Ginzburg-Landau equation, and we highlight the importance and usefulness of statistical physics for the successful study of large-scale ecological systems. Directions for future research, related for example to dynamical effects of coevolutionary rules and invasion reversals due to multi-point interactions, are outlined as well., Comment: 21 two-column pages, 16 figures; accepted for publication in Journal of the Royal Society Interface

The spatio-temporal arrangement of interacting populations often influences the maintenance of species diversity and is a subject of intense research. Here, we study the spatio-temporal patterns arising from the cyclic competition between three species in two dimensions. Inspired by recent experiments, we consider a generic metapopulation model comprising "rock-paper-scissors" interactions via dominance removal and replacement, reproduction, mutations, pair-exchange and hopping of individuals. By combining analytical and numerical methods, we obtain the model's phase diagram near its Hopf bifurcation and quantitatively characterize the properties of the spiraling patterns arising in each phase. The phases characterizing the cyclic competition away far from the Hopf bifurcation (at low mutation rate) are also investigated. Our analytical approach relies on the careful analysis of the properties of the complex Ginzburg-Landau equation derived through a controlled (perturbative) multiscale expansion around the model's Hopf bifurcation. Our results allows us to clarify when spatial "rock-paper-scissors" competition leads to stable spiral waves and under which circumstances they are influenced by nonlinear mobility., Comment: 16 two-column pages, 16 figures

Reciprocity is firmly established as an important mechanism that promotes cooperation. An efficient information exchange is likewise important, especially on structured populations, where interactions between players are limited. Motivated by these two facts, we explore the role of facilitators in social dilemmas on networks. Facilitators are here mirrors to their neighbors -- they cooperate with cooperators and defect with defectors -- but they do not participate in the exchange of strategies. As such, in addition to introducing direct reciprocity, they also obstruct information exchange. In well-mixed populations, facilitators favor the replacement and invasion of defection by cooperation as long as their number exceeds a critical value. In structured populations, on the other hand, there exists a delicate balance between the benefits of reciprocity and the deterioration of information exchange. Extensive Monte Carlo simulations of social dilemmas on various interaction networks reveal that there exists an optimal interplay between reciprocity and information exchange, which sets in only when a small number of facilitators occupies the main hubs of the scale-free network. The drawbacks of missing cooperative hubs are more than compensated by reciprocity and, at the same time, the compromised information exchange is routed via the auxiliary hubs with only marginal losses in effectivity. These results indicate that it is not always optimal for the main hubs to become ''leaders of the masses'', but rather to exploit their highly connected state to promote tit-for-tat-like behavior., Comment: 8 two-column pages, 6 figures; accepted for publication in Physical Review E

We study the combined influence of selection and random fluctuations on the evolutionary dynamics of two-strategy ("cooperation" and "defection") games in populations comprising cooperation facilitators. The latter are individuals that support cooperation by enhancing the reproductive potential of cooperators relative to the fitness of defectors. By computing the fixation probability of a single cooperator in finite and well-mixed populations that include a fixed number of facilitators, and by using mean field analysis, we determine when selection promotes cooperation in the important classes of prisoner's dilemma, snowdrift and stag-hunt games. In particular, we identify the circumstances under which selection favors the replacement and invasion of defection by cooperation. Our findings, corroborated by stochastic simulations, show that the spread of cooperation can be promoted through various scenarios when the density of facilitators exceeds a critical value whose dependence on the population size and selection strength is analyzed. We also determine under which conditions cooperation is more likely to replace defection than vice versa. Keywords: Evolutionary games; dynamics of cooperation; social dilemmas; fixation; population dynamics., Comment: 12 pages, 5 figures. Version to be published (special issue on "collective behavior and evolutionary games")

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., Comment: 7 pages, 5 figures - final version

Species diversity in ecosystems is often accompanied by the self-organisation of the population into fascinating spatio-temporal patterns. Here, we consider a two-dimensional three-species population model and study the spiralling patterns arising from the combined effects of generic cyclic dominance, mutation, pair-exchange and hopping of the individuals. The dynamics is characterised by nonlinear mobility and a Hopf bifurcation around which the system's phase diagram is inferred from the underlying complex Ginzburg-Landau equation derived using a perturbative multiscale expansion. While the dynamics is generally characterised by spiralling patterns, we show that spiral waves are stable in only one of the four phases. Furthermore, we characterise a phase where nonlinearity leads to the annihilation of spirals and to the spatially uniform dominance of each species in turn. Away from the Hopf bifurcation, when the coexistence fixed point is unstable, the spiralling patterns are also affected by nonlinear diffusion., Comment: 6 pages, 5 figures, supplementary material and movies available at http://dx.doi.org/10.6084/m9.figshare.96949, accepted by the EPL (Europhysics Letters)

In the framework of the three-party constrained voter model, where voters of two radical parties (A and B) interact with "centrists" (C and Cz), we study the competition between a persuasive majority and a committed minority. In this model, A's and B's are incompatible voters that can convince centrists or be swayed by them. Here, radical voters are more persuasive than centrists, whose sub-population consists of susceptible agents C and a fraction zeta of centrist zealots Cz. Whereas C's may adopt the opinions A and B with respective rates 1+delta_A and 1+delta_B (with delta_A>=delta_B>0), Cz's are committed individuals that always remain centrists. Furthermore, A and B voters can become (susceptible) centrists C with a rate 1. The resulting competition between commitment and persuasion is studied in the mean field limit and for a finite population on a complete graph. At mean field level, there is a continuous transition from a coexistence phase when zeta= Delta_c. In a finite population of size N, demographic fluctuations lead to centrism consensus and the dynamics is characterized by the mean consensus time tau. Because of the competition between commitment and persuasion, here consensus is reached much slower (zeta=Delta_c) than in the absence of zealots (when tau\simN). In fact, when zetadelta_B, whereas all species coexist when delta_A=delta_B. When zeta>=Delta_c and the initial density of centrists is low, one finds tau\simln N (when N>>1). Our analytical findings are corroborated by stochastic simulations., Comment: 25 pages, 6 figures. Final version for the Journal of Statistical Physics (special issue on the "applications of statistical mechanics to social phenomena")

In the framework of the paradigmatic prisoner's dilemma, we investigate the evolutionary dynamics of social dilemmas in the presence of "cooperation facilitators". In our model, cooperators and defectors interact as in the classical prisoner's dilemma game, where selection favors defection. However, here the presence of a small number of cooperation facilitators enhances the fitness (reproductive potential) of cooperators, while it does not alter that of defectors. In a finite population of size N, the dynamics of the prisoner's dilemma with facilitators is characterized by the probability that cooperation takes over (fixation probability) and by the mean times to reach the absorbing states. These quantities are computed exactly and using Fokker-Planck equations. Our findings, corroborated by stochastic simulations, demonstrate that the influence of facilitators crucially depends on the difference between their density z and the game's cost-to-benefit ratio r. When z>r, the fixation of cooperators is likely in a large population and, under weak selection pressure, invasion and replacement of defection by cooperation is favored by selection if b(z-r)(1-z)>1/N, where 0

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., Comment: 5 pages, 4 figures, to appear in Physical Review Letters

Motivated by the dynamics of cultural change and diversity, we generalize the three-species constrained voter model on a complete graph introduced in [J. Phys. A 37, 8479 (2004)]. In this opinion dynamics model, a population of size N is composed of "leftists" and "rightists" that interact with "centrists": a leftist and centrist can both become leftists with rate (1+q)/2 or centrists with rate (1-q)/2 (and similarly for rightists and centrists), where q denotes the bias towards extremism (q>0) or centrism (q<0). This system admits three absorbing fixed points and a "polarization" line along which a frozen mixture of leftists and rightists coexist. In the realm of Fokker-Planck equation, and using a mapping onto a population genetics model, we compute the fixation probability of ending in every absorbing state and the mean times for these events. We therefore show, especially in the limit of weak bias and large population size when |q|~1/N and N>>1, how fluctuations alter the mean field predictions: polarization is likely when q>0, but there is always a finite probability to reach a consensus; the opposite happens when q<0. Our findings are corroborated by stochastic simulations., Comment: 6 pages in EPL format, 3 color figures (6 panels). Minor modifications. To appear in EPL (Europhysics Letters)

We employ Monte Carlo simulations to numerically study the temporal evolution and transient oscillations of the population densities, the associated frequency power spectra, and the spatial correlation functions in the (quasi-)steady state in two-dimensional stochastic May--Leonard models of mobile individuals, allowing for particle exchanges with nearest-neighbors and hopping onto empty sites. We therefore consider a class of four-state three-species cyclic predator-prey models whose total particle number is not conserved. We demonstrate that quenched disorder in either the reaction or in the mobility rates hardly impacts the dynamical evolution, the emergence and structure of spiral patterns, or the mean extinction time in this system. We also show that direct particle pair exchange processes promote the formation of regular spiral structures. Moreover, upon increasing the rates of mobility, we observe a remarkable change in the extinction properties in the May--Leonard system (for small system sizes): (1) As the mobility rate exceeds a threshold that separates a species coexistence (quasi-)steady state from an absorbing state, the mean extinction time as function of system size N crosses over from a functional form ~ e^{cN} / N (where c is a constant) to a linear dependence; (2) the measured histogram of extinction times displays a corresponding crossover from an (approximately) exponential to a Gaussian distribution. The latter results are found to hold true also when the mobility rates are randomly distributed., Comment: 9 pages, 4 figures; to appear in Eur. Phys. J. B (2011)

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. U.S.A. Vol.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<< s. Here, we adopt an approach based on the direct treatment of the underlying Markov chain (birth-death process) obeyed by the allele frequency (whose dynamics is prescribed by the Moran model), which allows to accurately account for the effects of large fluctuations. After a general description of the theory, we focus on the case of a deleterious mutant allele (i.e. s>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 that is shown to be an accurate approximation only when N_e s^2 << 1, where N_e is the effective population size., Comment: 26 pages, 5 figures. Accepted by the Journal of Theoretical Biology

We study large fluctuations in evolutionary games belonging to the coordination and anti-coordination classes. The dynamics of these games, modeling cooperation dilemmas, is characterized by a coexistence fixed point separating two absorbing states. We are particularly interested in the problem of fixation that refers to the possibility that a few mutants take over the entire population. Here, the fixation phenomenon is induced by large fluctuations and is investigated by a semi-classical WKB (Wentzel-Kramers-Brillouin) theory generalized to treat stochastic systems possessing multiple absorbing states. Importantly, this method allows us to analyze the combined influence of selection and random fluctuations on the evolutionary dynamics \textit{beyond} the weak selection limit often considered in previous works. We accurately compute, including pre-exponential factors, the probability distribution function in the long-lived coexistence state and the mean fixation time necessary for a few mutants to take over the entire population in anti-coordination games, and also the fixation probability in the coordination class. Our analytical results compare excellently with extensive numerical simulations. Furthermore, we demonstrate that our treatment is superior to the Fokker-Planck approximation when the selection intensity is finite., Comment: 17 pages, 10 figures, to appear in JSTAT

We study several variants of the stochastic four-state rock-paper-scissors game or, equivalently, cyclic three-species predator-prey models with conserved total particle density, by means of Monte Carlo simulations on one- and two-dimensional lattices. Specifically, we investigate the influence of spatial variability of the reaction rates and site occupancy restrictions on the transient oscillations of the species densities and on spatial correlation functions in the quasi-stationary coexistence state. For small systems, we also numerically determine the dependence of typical extinction times on the number of lattice sites. In stark contrast with two-species stochastic Lotka-Volterra systems, we find that for our three-species models with cyclic competition quenched disorder in the reaction rates has very little effect on the dynamics and the long-time properties of the coexistence state. Similarly, we observe that site restriction only has a minor influence on the system's dynamical properties. Our results therefore demonstrate that the features of the spatial rock-paper-scissors system are remarkably robust with respect to model variations, and stochastic fluctuations as well as spatial correlations play a comparatively minor role., Comment: 11 pages, revtex, 17 figures, to appear in Phys. Rev. E (2010)

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., Comment: 25 pages, 9 figures. To appear in the Journal of Theoretical Biology

One of the most striking effect of fluctuations in evolutionary game theory is the possibility for mutants to fixate (take over) an entire population. Here, we generalize a recent WKB-based theory to study fixation in evolutionary games under non-vanishing selection, and investigate the relation between selection intensity w and demographic (random) fluctuations. This allows the accurate treatment of large fluctuations and yields the probability and mean times of fixation beyond the weak selection limit. The power of the theory is demonstrated on prototypical models of cooperation dilemmas with multiple absorbing states. Our predictions compare excellently with numerical simulations and, for finite w, significantly improve over those of the Fokker-Planck approximation., Comment: 4 figures, to appear in EPL (Europhysics Letters)

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., Comment: 21 pages, 10 figures. To appear in J. Theor. Biol

Noise and spatial degrees of freedom characterize most ecosystems. Some aspects of their influence on the coevolution of populations with cyclic interspecies competition have been demonstrated in recent experiments [e.g. B. Kerr et al., Nature {\bf 418}, 171 (2002)]. To reach a better theoretical understanding of these phenomena, we consider a paradigmatic spatial model where three species exhibit cyclic dominance. Using an individual-based description, as well as stochastic partial differential and deterministic reaction-diffusion equations, we account for stochastic fluctuations and spatial diffusion at different levels, and show how fascinating patterns of entangled spirals emerge. We rationalize our analysis by computing the spatio-temporal correlation functions and provide analytical expressions for the front velocity and the wavelength of the propagating spiral waves., Comment: 4 pages of main text, 3 color figures + 2 pages of supplementary material (EPAPS Document). Final version for Physical Review Letters

Biodiversity is essential to the viability of ecological systems. Species diversity in ecosystems is promoted by cyclic, non-hierarchical interactions among competing populations. Such non-transitive relations lead to an evolution with central features represented by the `rock-paper-scissors' game, where 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 temporal evolution. 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 evolution of ecological systems and are relevant for the formation and propagation of patterns in excitable media, such as chemical kinetics or epidemic outbreaks., Comment: Final submitted version; the printed version can be found at http://dx.doi.org/10.1038/nature06095 Supplementary movies are available at http://www.theorie.physik.uni-muenchen.de/lsfrey/images_content/movie1.AVI and http://www.theorie.physik.uni-muenchen.de/lsfrey/images_content/movie2.AVI

Nonequilibrium collective motion is ubiquitous in nature and often results in a rich collection of intringuing phenomena, such as the formation of shocks or patterns, subdiffusive kinetics, traffic jams, and nonequilibrium phase transitions. These stochastic many-body features characterize transport processes in biology, soft condensed matter and, possibly, also in nanoscience. Inspired by these applications, a wide class of lattice-gas models has recently been considered. Building on the celebrated {\it totally asymmetric simple exclusion process} (TASEP) and a generalization accounting for the exchanges with a reservoir, we discuss the qualitative and quantitative nonequilibrium properties of these model systems. We specifically analyze the case of a dimeric lattice gas, the transport in the presence of pointwise disorder and along coupled tracks., Comment: 21 pages, 10 figures. Pedagogical paper based on a lecture delivered at the conference on "Stochastic models in biological sciences" (May 29 - June 2, 2006 in Warsaw). For the Banach Center Publications

Finite-size fluctuations arising in the dynamics of competing populations may have dramatic influence on their fate. As an example, in this article, we investigate a model of three species which dominate each other in a cyclic manner. Although the deterministic approach predicts (neutrally) stable coexistence of all species, for any finite population size, the intrinsic stochasticity unavoidably causes the eventual extinction of two of them., Comment: 5 pages, 2 figures. Proceedings paper of the workshop "Stochastic models in biological sciences" (May 29 - June 2, 2006 in Warsaw) for the Banach Center Publications

We consider a broad class of stochastic lattice predator-prey models, whose main features are overviewed. In particular, this article aims at drawing a picture of the influence of spatial fluctuations, which are not accounted for by the deterministic rate equations, on the properties of the stochastic models. Here, we outline the robust scenario obeyed by most of the lattice predator-prey models with an interaction "a' la Lotka-Volterra". We also show how a drastically different behavior can emerge as the result of a subtle interplay between long-range interactions and a nearest-neighbor exchange process., Comment: 5 pages, 2 figures. Proceedings paper of the workshop "Stochastic models in biological sciences" (May 29 - June 2, 2006 in Warsaw) for the Banach Center Publications

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 {\bf 418}, 171 (2002)] and B. Kirkup and M. A. Riley [Nature {\bf 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 \emph{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., Comment: 12 pages, 9 figures, minor corrections

We study the general properties of stochastic two-species models for predator-prey competition and coexistence with Lotka-Volterra type interactions defined on a $d$-dimensional lattice. Introducing spatial degrees of freedom and allowing for stochastic fluctuations generically invalidates the classical, deterministic mean-field picture. Already within mean-field theory, however, spatial constraints, modeling locally limited resources, lead to the emergence of a continuous active-to-absorbing state phase transition. Field-theoretic arguments, supported by Monte Carlo simulation results, indicate that this transition, which represents an extinction threshold for the predator population, is governed by the directed percolation universality class. In the active state, where predators and prey coexist, the classical center singularities with associated population cycles are replaced by either nodes or foci. In the vicinity of the stable nodes, the system is characterized by essentially stationary localized clusters of predators in a sea of prey. Near the stable foci, however, the stochastic lattice Lotka-Volterra system displays complex, correlated spatio-temporal patterns of competing activity fronts. Correspondingly, the population densities in our numerical simulations turn out to oscillate irregularly in time, with amplitudes that tend to zero in the thermodynamic limit. Yet in finite systems these oscillatory fluctuations are quite persistent, and their features are determined by the intrinsic interaction rates rather than the initial conditions. We emphasize the robustness of this scenario with respect to various model perturbations., Comment: 19 pages, 11 figures, 2-column revtex4 format. Minor modifications. Accepted in the Journal of Statistical Physics. Movies corresponding to Figures 2 and 3 are available at http://www.phys.vt.edu/~tauber/PredatorPrey/movies/