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Connectivity is the safety network for biodiversity conservation because connected habitats are more effective for saving the species and ecological functions. The nature of coupling for connectivity also plays an important role in the co-existence of species in cyclic-dominance. The rock-paper-scissors game is one of the paradigmatic mathematical model in evolutionary game theory to understand the mechanism of biodiversity in cyclic-dominance. In this paper, the metapopulation model for rock-paper-scissors with mutations is presented in which the total population is divided into patches and the patches form a network of complete graph. The migration among patches is allowed through simple random walk. The replicator-mutator equations are used with the migration term. When migration is allowed then the population of the patches will synchronized and attain stable state through Hopf bifurcation. Apart form this, two phases are observed when the strategies of one of the species mutate to other two species: co-existence of all the species phase and existence of one kind of species phase. The transition from one phase to another phase is taking place due to transcritical bifurcation. The dynamics of the population of species of rock, paper, scissors is studied in the environment of homogeneous and heterogeneous mutation. Numerical simulations have been performed when mutation is allowed in all the patches (homogeneous mutation) and some of the patches (heterogeneous mutation). It has been observed that when the number of patches is increased in the case of heterogeneous mutation then the population of any of the species will not extinct and all the species will co-exist.

One of the simplest examples of a robust heteroclinic cycle involves three saddle equilibria: each one is unstable to the next in turn, and connections from one to the next occur within invariant subspaces. Such a situation can be described by a third-order ordinary differential equation (ODE), and typical trajectories approach each equilibrium point in turn, spending progressively longer to cycle around the three points but never stopping. This cycle has been invoked as a model of cyclic competition between populations adopting three strategies, characterised as Rock, Paper and Scissors. When spatial distribution and mobility of the populations is taken into account, waves of Rock can invade regions of Scissors, only to be invaded by Paper in turn. The dynamics is described by a set of partial differential equations (PDEs) that has travelling wave (in one dimension) and spiral (in two dimensions) solutions. In this paper, we explore how the robust heteroclinic cycle in the ODE manifests itself in the PDEs. Taking the wavespeed as a parameter, and moving into a travelling frame, the PDEs reduce to a sixth-order set of ODEs, in which travelling waves are created in a Hopf bifurcation and are destroyed in three different heteroclinic bifurcations, depending on parameters, as the travelling wave approaches the heteroclinic cycle. We explore the three different heteroclinic bifurcations, none of which have been observed in the context of robust heteroclinic cycles previously. These results are an important step towards a full understanding of the spiral patterns found in two dimensions, with possible application to travelling waves and spirals in other population dynamics models., Comment: 36 pages, 8 figures

Classic frameworks of rock-paper-scissors game have been assumed in a closed community that a density of each group is only affected by internal factors such as competition interplay among groups and reproduction itself. In real systems in ecological and social sciences, however, the survival and a change of a density of a group can be also affected by various external factors. One of common features in real population systems in ecological and social sciences is population flow that is characterized by population inflow and outflow in a group or a society, which has been usually overlooked in previous works on models of rock-paper-scissors game. In this paper, we suggest the rock-paper-scissors system by implementing population flow and investigate its effect on biodiversity. For two scenarios of either balanced or imbalanced population flow, we found that the population flow can strongly affect group diversity by exhibiting rich phenomena. In particular, while the balanced flow can only lead the persistent coexistence of all groups which accompanies a phase transition through supercritical Hopf bifurcation on different carrying simplices, the imbalanced flow strongly facilitates rich dynamics such as alternative stable survival states by exhibiting various group survival states and multistability of sole group survivals by showing not fully covered but spirally entangled basins of initial densities due to local stabilities of associated fixed points. In addition, we found that, the system can exhibit oscillatory dynamics for coexistence by relativistic interplay of population flows which can capture the robustness of the coexistence state. Applying population flow in the rock-paper-scissors system can ultimately change a community paradigm from closed to open one, and our foundation can eventually reveal that population flow can be also a significant factor on a group density which is independent to fundamental interactions among groups.

Two limit cycles for the continuous model of the rockscissorspaper (RSP) game is constructed.The Hopf bifurcation method is used for the construction of limit cycles.The results give a partial answer to an open question posed by Neumann and Schuster. In this paper we construct two limit cycles with a heteroclinic polycycle for the three-dimensional continuous model of the rockscissorspaper (RSP) game between bacteriocin producing bacteria. Our construction gives a partial answer to an open question posed by Neumann and Schuster (2007) concerning how many limit cycles can coexist for the RSP game.

We investigate the dynamics of three-strategy (rock–paper–scissors) replicator equations in which the fitness of each strategy is a function of the population frequencies delayed by a time interval $$T$$ . Taking $$T$$ as a bifurcation parameter, we demonstrate the existence of (non-degenerate) Hopf bifurcations in these systems and present an analysis of the resulting limit cycles using Lindstedt’s method.

The role of mutation, which is an error process in gene evolution, in systems of cyclically competing species has been studied from various perspectives, and it is regarded as one of the key factors for promoting coexistence of all species. In addition to naturally occurring mutations, many experiments in genetic engineering have involved targeted mutation techniques such as recombination between DNA and somatic cell sequences and have studied genetic modifications through loss or augmentation of cell functions. In this paper, we investigate nonlinear dynamics with targeted mutation in cyclically competing species. In different ways to classic approaches of mutation in cyclic games, we assume that mutation may occur in targeted individuals who have been removed from intraspecific competition. By investigating each scenario depending on the number of objects for targeted mutation analytically and numerically, we found that targeted mutation can lead to persistent coexistence of all species. In addition, under the specific condition of targeted mutation, we found that targeted mutation can lead to emergences of bistable states for species survival. Through the linear stability analysis of rate equations, we found that those phenomena are accompanied by Hopf bifurcation which is supercritical. Our findings may provide more global perspectives on understanding underlying mechanisms to control biodiversity in ecological/biological sciences, and evidences with mathematical foundations to resolve social dilemmas such as a turnover of group members by resigning with intragroup conflicts in social sciences.

We investigate a population of individuals who play the Rock–Paper–Scissors (RPS) game. The players choose strategies not only by optimizing their payoffs, but also taking into account the popularity of the strategies. For the standard RPS game, we find an asymptotically stable polymorphism with coexistence of all strategies. For the general RPS game we find the limit cycles. Their stability depends exclusively on two model parameters: the sum of the entries of the RPS payoff matrix, and a sensitivity parameter which characterizes the personality of the players. Apart from the supercritical Hopf bifurcation, we found the subcritical bifurcation numerically for some intervals of the parameters of the model.

In this work, important aspects of bacteriocin producing bacteria and their interplay are elucidated. Various attempts to model the resistant, producer and sensitive Escherichia coli strains in the so-called rock-scissors-paper (RSP) game had been made in the literature. The question arose whether there is a continuous model with a cyclic structure and admitting an oscillatory dynamics as observed in various experiments. The May-Leonard system admits a Hopf bifurcation, which is, however, degenerate and hence inadequate. The traditional differential equation model of the RSP-game cannot be applied either to the bacteriocin system because it involves positive interaction terms. In this paper, a plausible competitive Lotka-Volterra system model of the RSP game is presented and the dynamics generated by that model is analyzed. For the first time, a continuous, spatially homogeneous model that describes the competitive interaction between bacteriocin-producing, resistant and sensitive bacteria is established. The interaction terms have negative coefficients. In some experiments, for example, in mice cultures, migration seemed to be essential for the reinfection in the RSP cycle. Often statistical and spatial effects such as migration and mutation are regarded to be essential for periodicity. Our model gives rise to oscillatory dynamics in the RSP game without such effects. Here, a normal form description of the limit cycle and conditions for its stability are derived. The toxicity of the bacteriocin is used as a bifurcation parameter. Exact parameter ranges are obtained for which a stable (robust) limit cycle and a stable heteroclinic cycle exist in the three-species game. These parameters are in good accordance with the observed relations for the E. coli strains. The roles of growth rate and growth yield of the three strains are discussed. Numerical calculations show that the sensitive, which might be regarded as the weakest, can have the longest sojourn times.

In this paper we analyze the population dynamics of bacteria competing by anti-bacterial toxins (bacteriocins). Three types of bacteria involved in these dynamics can be distinguished: toxin producers, resistant bacteria and sensitive bacteria. Their interplay can be regarded as a R ock- S cissors- P aper - game (RSP). Here, this is modeled by a reasonable three-dimensional Lotka- Volterra ($L$V) type differential equation system. In contrast to earlier approaches to modeling the RSP game such as replicator equations, all interaction terms have negative signs because the interaction between the three different types of bacteria is purely competitive, either by toxification or by competition for nutrients. The model allows one to choose asymmetric parameter values. Depending on parameter values, our model gives rise to a stable steady state, a stable limit cycle or a heteroclinic orbit with three fixed points, each fixed point corresponding to the existence of only one bacteria type. An alternative model, the May - Leonard model, leads to coexistence only under very restricted conditions. We carry out a comprehensive analysis of the generic stability conditions of our model, using, among other tools, the Volterra-Lyapunov method. We also give biological interpretations of our theoretical results, in particular, of the predicted dynamics and of the ranges for parameter values where different dynamic behavior occurs. For example, one result is that the intrinsic growth rate of the producer is lower than that of the resistant while its growth yield is higher. This is in agreement with experimental results for the bacterium Listeria monocytogenes.

When playing games in groups, it is an advantage for individuals to have accurate statistical information on the strategies of their opponents. Such information may be obtained by remembering previous interactions. We consider a rock-paper-scissors game in which agents are able to recall their last m interactions, used to estimate the behavior of their opponents. At critical memory length, a Hopf bifurcation leads to the formation of stable limit cycles. In a mixed population, agents with longer memories have an advantage, provided the system has a stable fixed point, and there is some asymmetry in the payoffs of the pure strategies. However, at a critical concentration of long memory agents, the appearance of limit cycles destroys their advantage. By introducing population dynamics that favors successful agents, we show that the system evolves toward the bifurcation point.

We analyze the replicator-mutator equations for the Rock-Paper-Scissors game. Various graph-theoretic patterns of mutation are considered, ranging from a single unidirectional mutation pathway between two of the species, to global bidirectional mutation among all the species. Our main result is that the coexistence state, in which all three species exist in equilibrium, can be destabilized by arbitrarily small mutation rates. After it loses stability, the coexistence state gives birth to a stable limit cycle solution created in a supercritical Hopf bifurcation. This attracting periodic solution exists for all the mutation patterns considered, and persists arbitrarily close to the limit of zero mutation rate and a zero-sum game., 6 pages, 5 figures

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., 16 two-column pages, 16 figures

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

This paper proposes a novel fractional-order delayed Ross-Macdonald model for malaria transmission. This paper aims to systematically investigate the effect of both the incubation periods of Plasmodium and the order on the dynamic behavior of diseases. Utilizing inequality techniques, contraction mapping theory, fractional linear stability theorem, and bifurcation theory, several sufficient conditions for the existence and uniqueness of solutions, the local stability of the positive equilibrium point, and the existence of fractional-order Hopf bifurcation are obtained under different time delays cases. The results show that time delay can change the stability of system. System becomes unstable and generates a Hopf bifurcation when the delay increases to a certain value. Besides, the value of order influences the stability interval size. Thus, incubation periods and the order have a major effect on the dynamic behavior of the model. The effectiveness of the theoretical results is shown through numerical simulations.

In the theoretical research of chaotic dynamical system, the different type of bifurcations is a very interesting powerful tool for analyzing the qualitative behavior of chaotic dynamical system; this short paper is devoted to analysis of a simple 2-D symmetry discrete chaotic map with quadratic and cubic nonlinearities. The dynamical behaviors of the map are investigated by mathematical analysis and simulated numerically using package of Matlab . We compute numerically the bifurcation diagram and largest Lyapunov exponent and phase portraits. The research results indicate that there are interesting nonlinear physical phenomena in this simple 2-D symmetry discrete cubic map, such as symmetry bifurcation, Hopf bifurcation, symmetry breaking bifurcation and identical symmetric attractors. The important nonlinear physical phenomena obtained in this paper would benefit the study of the cubic chaotic map and the development of the theory of chaotic discrete dynamical systems. En la investigación teórica de los sistemas dinámicos caóticos, los diferentes tipos de bifurcaciones son una herramienta poderosa muy interesante para analizar el comportamiento cualitativo de los sistemas dinámicos caóticos; este breve artículo está dedicado al análisis de un mapa caótico discreto de simetría bidimensional simple con no linealidades cuadráticas y cúbicas. Los comportamientos dinámicos del mapa se investigan mediante análisis matemático y se simulan numéricamente utilizando el paquete de Matlab . Calculamos numéricamente el diagrama de bifurcación y el mayor exponente de Lyapunov y los retratos de fase. Los resultados de la investigación indican que existen interesantes fenómenos físicos no lineales en este sencillo mapa cúbico discreto de simetría 2-D, como la bifurcación de simetría, la bifurcación de Hopf, la bifurcación de ruptura de simetría y los atractores simétricos idénticos. Los importantes fenómenos físicos no lineales obtenidos en este trabajo beneficiarían el estudio del mapa cúbico caótico y el desarrollo de la teoría de los sistemas dinámicos discretos caóticos.

During the outbreak of emerging infectious diseases, media coverage and medical resource play important roles in affecting the disease transmission. To investigate the effects of the saturation of media coverage and limited medical resources, we proposed a mathematical model with extra compartment of media coverage and two nonlinear functions. We theoretically and numerically investigate the dynamics of the proposed model. Given great difficulties caused by high nonlinearity in theoretical analysis, we separately considered subsystems with only nonlinear recovery or with only saturated media impact. For the model with only nonlinear recovery, we theoretically showed that backward bifurcation can occur and multiple equilibria may coexist under certain conditions in this case. Numerical simulations reveal the rich dynamic behaviors, including forward-backward bifurcation, Hopf bifurcation, saddle-node bifurcation, homoclinic bifurcation and unstable limit cycle. So the limitation of medical resources induces rich dynamics and causes much difficulties in eliminating the infectious diseases. We then investigated the dynamics of the system with only saturated media impact and concluded that saturated media impact hardly induces the complicated dynamics. Further, we parameterized the proposed model on the basis of the COVID-19 case data in mainland China and data related to news items, and estimated the basic reproduction number to be 2.86. Sensitivity analyses were carried out to quantify the relative importance of parameters in determining the cumulative number of infected individuals at the end of the first month of the outbreak. Combining with numerical analyses, we suggested that providing adequate medical resources and improving media response to infection or individuals' response to mass media may reduce the cumulative number of the infected individuals, which mitigates the transmission dynamics during the early stage of the COVID-19 pandemic.

The ecological theory of species interactions rests largely on the competition, interference, and predator–prey models. In this paper, we propose and investigate a three-species predator–prey model to inspect the mutual interference between predators. We analyze boundedness and Kolmogorov conditions for the non-spatial model. The dynamical behavior of the system is analyzed by stability and Hopf bifurcation analysis. The Turing instability criteria for the Spatio-temporal system is estimated. In the numerical simulation, phase portrait with time evolution diagrams shows periodic and chaotic oscillations. Bifurcation diagrams show the very rich and complex dynamical behavior of the non-spatial model. We calculate the Lyapunov exponent to justify the dynamics of the non-spatial model. A variety of patterns like interference, spot, and stripe are observed with special emphasis on Beddington–DeAngelis function response. These complex patterns explore the beauty of the spatio-temporal model and it can be easily related to real-world biological systems.

In this paper, we analyze the complexity of an eco-epidemiological model for phytoplankton–zooplankton system in presence of toxicity and time delay. Holling type II function response is incorporated to address the predation rate as well as toxic substance distribution in zooplankton. It is also presumed that infected phytoplankton does recover from the viral infection. In the absence of time delay, stability and Hopf-bifurcation conditions are investigated to explore the system dynamics around all the possible equilibrium points. Further, in the presence of time delay, conditions for local stability are derived around the interior equilibria and the properties of the periodic solution are obtained by applying normal form theory and central manifold arguments. Computational simulation is performed to illustrate our theoretical findings. It is explored that system dynamics is very sensitive corresponding to carrying capacity and toxin liberation rate and able to generate chaos. Further, it is observed that time delay in the viral infection process can destabilize the phytoplankton density whereas zooplankton density remains in its old state. Incorporation of time delay also gives the scenario of double Hopf-bifurcation. Some control parameters are discussed to stabilize system dynamics. The effect of time delay on (i) growth rate of susceptible phytoplankton shows the extinction and double Hopf-bifurcation in the zooplankton population, (ii) a sufficiently large value of carrying capacity stabilizes the chaotic dynamics or makes the whole system chaotic with further increment.