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Global bifurcation analysis of a rational Holling system
Computer Research and Modeling, 2017, v. 9, no. 4, pp. 537-545Views (last year): 11.In this paper, we consider a quartic family of planar vector fields corresponding to a rational Holling system which models the dynamics of the populations of predators and their prey in a given ecological or biomedical system and which is a variation on the classical Lotka–Volterra system. For the latter system, the change of the prey density per unit of time per predator called the response function is proportional to the prey density. This means that there is no saturation of the predator when the amount of available prey is large. However, it is more realistic to consider a nonlinear and bounded response function, and in fact different response functions have been used in the literature to model the predator response. After algebraic transformations, the rational Holling system can be written in the form of a quartic dynamical system. To investigate the character and distribution of the singular points in the phase plane of the quartic system, we use our method the sense of which is to obtain the simplest (well-known) system by vanishing some parameters (usually field rotation parameters) of the original system and then to input these parameters successively one by one studying the dynamics of the singular points (both finite and infinite) in the phase plane. Using the obtained information on singular points and applying our geometric approach to the qualitative analysis, we study the limit cycle bifurcations of the quartic system. To control all of the limit cycle bifurcations, especially, bifurcations of multiple limit cycles, it is necessary to know the properties and combine the effects of all of the rotation parameters. It can be done by means of the Wintner–Perko termination principle stating that the maximal one-parameter family of multiple limit cycles terminates either at a singular point which is typically of the same multiplicity (cyclicity) or on a separatrix cycle which is also typically of the same multiplicity (cyclicity). Applying this principle, we prove that the quartic system (and the corresponding rational Holling system) can have at most two limit cycles surrounding one singular point.
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Global bifurcation analysis of a quartic predator–prey model
Computer Research and Modeling, 2011, v. 3, no. 2, pp. 125-134Views (last year): 5. Citations: 3 (RSCI).We complete the global bifurcation analysis of a quartic predator–prey model. In particular, studying global bifurcations of singular points and limit cycles, we prove that the corresponding dynamical system has at most two limit cycles.
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Analysis of noise-induced destruction of coexistence regimes in «prey–predator» population model
Computer Research and Modeling, 2016, v. 8, no. 4, pp. 647-660Views (last year): 14. Citations: 4 (RSCI).The paper is devoted to the analysis of the proximity of the population system to dangerous boundaries. An intersection of these boundaries results in the collapse of the stable coexistence of interacting populations. As a reason of such destruction one can consider random perturbations inevitably presented in any living system. This study is carried out on the example of the well-known model of interaction between predator and prey populations, taking into account both a stabilizing factor of the competition of predators for another than prey resources, and also a destabilizing saturation factor for predators. To describe the saturation of predators, we use the second type Holling trophic function. The dynamics of the system is studied as a function of the predator saturation, and the coefficient of predator competition for resources other than prey. The paper presents a parametric description of the possible dynamic regimes of the deterministic model. Here, local and global bifurcations are studied, and areas of sustainable coexistence of populations in equilibrium and the oscillation modes are described. An interesting feature of this mathematical model, firstly considered by Bazykin, is a global bifurcation of the birth of limit cycle from the separatrix loop. We study the effects of noise on the equilibrium and oscillatory regimes of coexistence of predator and prey populations. It is shown that an increase of the intensity of random disturbances can lead to significant deformations of these regimes right up to their destruction. The aim of this work is to develop a constructive probabilistic criterion for the proximity of the population stochastic system to the dangerous boundaries. The proposed approach is based on the mathematical technique of stochastic sensitivity functions, and the method of confidence domains. In the case of a stable equilibrium, this confidence domain is an ellipse. For the stable cycle, this domain is a confidence band. The size of the confidence domain is proportional to the intensity of the noise and stochastic sensitivity of the initial deterministic attractor. A geometric criterion of the exit of the population system from sustainable coexistence mode is the intersection of the confidence domain and the corresponding separatrix of the unforced deterministic model. An effectiveness of this analytical approach is confirmed by the good agreement of theoretical estimates and results of direct numerical simulations.
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Mathematical model of predator – prey system with lower critical prey density
Computer Research and Modeling, 2009, v. 1, no. 1, pp. 51-56Views (last year): 23. Citations: 5 (RSCI).A mathematical model of predator – prey microecosystem with lower critical population number of prey is considered. The predator – prey system is assumed to be under harvesting. Harvesting intensity variations generate changes in two model parameters which are considered as controllable. Bifurcation diagram in control-lable parameters plane is constructed and corresponding phase portraits are represented.
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Diffusion–reaction–advection equations for the predator–prey system in a heterogeneous environment
Computer Research and Modeling, 2021, v. 13, no. 6, pp. 1161-1176We analyze variants of considering the inhomogeneity of the environment in computer modeling of the dynamics of a predator and prey based on a system of reaction-diffusion–advection equations. The local interaction of species (reaction terms) is described by the logistic law for the prey and the Beddington –DeAngelis functional response, special cases of which are the Holling type II functional response and the Arditi – Ginzburg model. We consider a one-dimensional problem in space for a heterogeneous resource (carrying capacity) and three types of taxis (the prey to resource and from the predator, the predator to the prey). An analytical approach is used to study the stability of stationary solutions in the case of local interaction (diffusionless approach). We employ the method of lines to study diffusion and advective processes. A comparison of the critical values of the mortality parameter of predators is given. Analysis showed that at constant coefficients in the Beddington –DeAngelis model, critical values are variable along the spatial coordinate, while we do not observe this effect for the Arditi –Ginzburg model. We propose a modification of the reaction terms, which makes it possible to take into account the heterogeneity of the resource. Numerical results on the dynamics of species for large and small migration coefficients are presented, demonstrating a decrease in the influence of the species of local members on the emerging spatio-temporal distributions of populations. Bifurcation transitions are analyzed when changing the parameters of diffusion–advection and reaction terms.
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Modeling the dynamics of plankton community considering phytoplankton toxicity
Computer Research and Modeling, 2022, v. 14, no. 6, pp. 1301-1323We propose a three-component discrete-time model of the phytoplankton-zooplankton community, in which toxic and non-toxic species of phytoplankton compete for resources. The use of the Holling functional response of type II allows us to describe an interaction between zooplankton and phytoplankton. With the Ricker competition model, we describe the restriction of phytoplankton biomass growth by the availability of external resources (mineral nutrition, oxygen, light, etc.). Many phytoplankton species, including diatom algae, are known not to release toxins if they are not damaged. Zooplankton pressure on phytoplankton decreases in the presence of toxic substances. For example, Copepods are selective in their food choices and avoid consuming toxin-producing phytoplankton. Therefore, in our model, zooplankton (predator) consumes only non-toxic phytoplankton species being prey, and toxic species phytoplankton only competes with non-toxic for resources.
We study analytically and numerically the proposed model. Dynamic mode maps allow us to investigate stability domains of fixed points, bifurcations, and the evolution of the community. Stability loss of fixed points is shown to occur only through a cascade of period-doubling bifurcations. The Neimark – Sacker scenario leading to the appearance of quasiperiodic oscillations is found to realize as well. Changes in intrapopulation parameters of phytoplankton or zooplankton can lead to abrupt transitions from regular to quasi-periodic dynamics (according to the Neimark – Sacker scenario) and further to cycles with a short period or even stationary dynamics. In the multistability areas, an initial condition variation with the unchanged values of all model parameters can shift the current dynamic mode or/and community composition.
The proposed discrete-time model of community is quite simple and reveals dynamics of interacting species that coincide with features of experimental dynamics. In particular, the system shows behavior like in prey-predator models without evolution: the predator fluctuations lag behind those of prey by about a quarter of the period. Considering the phytoplankton genetic heterogeneity, in the simplest case of two genetically different forms: toxic and non-toxic ones, allows the model to demonstrate both long-period antiphase oscillations of predator and prey and cryptic cycles. During the cryptic cycle, the prey density remains almost constant with fluctuating predators, which corresponds to the influence of rapid evolution masking the trophic interaction.
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The key approaches and review of current researches on dynamics of structured and interacting populations
Computer Research and Modeling, 2019, v. 11, no. 1, pp. 119-151Views (last year): 40. Citations: 2 (RSCI).The review and systematization of current papers on the mathematical modeling of population dynamics allow us to conclude the key interests of authors are two or three main research lines related to the description and analysis of the dynamics of both local structured populations and systems of interacting homogeneous populations as ecological community in physical space. The paper reviews and systematizes scientific studies and results obtained within the framework of dynamics of structured and interacting populations to date. The paper describes the scientific idea progress in the direction of complicating models from the classical Malthus model to the modern models with various factors affecting population dynamics in the issues dealing with modeling the local population size dynamics. In particular, they consider the dynamic effects that arise as a result of taking into account the environmental capacity, density-dependent regulation, the Allee effect, complexity of an age and a stage structures. Particular attention is paid to the multistability of population dynamics. In addition, studies analyzing harvest effect on structured population dynamics and an appearance of the hydra effect are presented. The studies dealing with an appearance and development of spatial dissipative structures in both spatially separated populations and communities with migrations are discussed. Here, special attention is also paid to the frequency and phase multistability of population dynamics, as well as to an appearance of spatial clusters. During the systematization and review of articles on modeling the interacting population dynamics, the focus is on the “prey–predator” community. The key idea and approaches used in current mathematical biology to model a “prey–predator” system with community structure and harvesting are presented. The problems of an appearance and stability of the mosaic structure in communities distributed spatially and coupled by migration are also briefly discussed.
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Stochastic sensitivity analysis of dynamic transformations in the “two prey – predator” model
Computer Research and Modeling, 2022, v. 14, no. 6, pp. 1343-1356This work is devoted to the study of the problem of modeling and analyzing complex oscillatory modes, both regular and chaotic, in systems of interacting populations in the presence of random perturbations. As an initial conceptual deterministic model, a Volterra system of three differential equations is considered, which describes the dynamics of prey populations of two competing species and a predator. This model takes into account the following key biological factors: the natural increase in prey, their intraspecific and interspecific competition, the extinction of predators in the absence of prey, the rate of predation by predators, the growth of the predator population due to predation, and the intensity of intraspecific competition in the predator population. The growth rate of the second prey population is used as a bifurcation parameter. At a certain interval of variation of this parameter, the system demonstrates a wide variety of dynamic modes: equilibrium, oscillatory, and chaotic. An important feature of this model is multistability. In this paper, we focus on the study of the parametric zone of tristability, when a stable equilibrium and two limit cycles coexist in the system. Such birhythmicity in the presence of random perturbations generates new dynamic modes that have no analogues in the deterministic case. The aim of the paper is a detailed study of stochastic phenomena caused by random fluctuations in the growth rate of the second population of prey. As a mathematical model of such fluctuations, we consider white Gaussian noise. Using methods of direct numerical modeling of solutions of the corresponding system of stochastic differential equations, the following phenomena have been identified and described: unidirectional stochastic transitions from one cycle to another, trigger mode caused by transitions between cycles, noise-induced transitions from cycles to the equilibrium, corresponding to the extinction of the predator and the second prey population. The paper presents the results of the analysis of these phenomena using the Lyapunov exponents, and identifies the parametric conditions for transitions from order to chaos and from chaos to order. For the analytical study of such noise-induced multi-stage transitions, the technique of stochastic sensitivity functions and the method of confidence regions were applied. The paper shows how this mathematical apparatus allows predicting the intensity of noise, leading to qualitative transformations of the modes of stochastic population dynamics.
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Modeling of plankton community state with density-dependent death and spatial activity of zooplankton
Computer Research and Modeling, 2016, v. 8, no. 3, pp. 549-560Views (last year): 6.A vertically distributed three-component model of marine ecosystem is considered. State of the plankton community with nutrients is analyzed under the active movement of zooplankton in a vertical column of water. The necessary conditions of the Turing instability in the vicinity of the spatially homogeneous equilibrium are obtained. Stability of the spatially homogeneous equilibrium, the Turing instability and the oscillatory instability are examined depending on the biological characteristics of zooplankton and spatial movement of plankton. It is shown that at low values of zooplankton grazing rate and intratrophic interaction rate the system is Turing instable when the taxis rate is low. Stabilization occurs either through increased decline of zooplankton either by increasing the phytoplankton diffusion. With the increasing rate of consumption of phytoplankton range of parameters that determine the stability is reduced. A type of instability depends on the phytoplankton diffusion. For large values of diffusion oscillatory instability is observed, with a decrease in the phytoplankton diffusion zone of Turing instability is increases. In general, if zooplankton grazing rate is faster than phytoplankton growth rate the spatially homogeneous equilibrium is Turing instable or oscillatory instable. Stability is observed only at high speeds of zooplankton departure or its active movements. With the increase in zooplankton search activity spatial distribution of populations becomes more uniform, increasing the rate of diffusion leads to non-uniform spatial distribution. However, under diffusion the total number of the population is stabilized when the zooplankton grazing rate above the rate of phytoplankton growth. In general, at low rate of phytoplankton consumption the spatial structures formation is possible at low rates of zooplankton decline and diffusion of all the plankton community. With the increase in phytoplankton predation rate the phytoplankton diffusion and zooplankton spatial movement has essential effect on the spatial instability.
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Models of population process with delay and the scenario for adaptive resistance to invasion
Computer Research and Modeling, 2022, v. 14, no. 1, pp. 147-161Changes in abundance for emerging populations can develop according to several dynamic scenarios. After rapid biological invasions, the time factor for the development of a reaction from the biotic environment will become important. There are two classic experiments known in history with different endings of the confrontation of biological species. In Gause’s experiments with ciliates, the infused predator, after brief oscillations, completely destroyed its resource, so its $r$-parameter became excessive for new conditions. Its own reproductive activity was not regulated by additional factors and, as a result, became critical for the invader. In the experiments of the entomologist Uchida with parasitic wasps and their prey beetles, all species coexisted. In a situation where a population with a high reproductive potential is regulated by several natural enemies, interesting dynamic effects can occur that have been observed in phytophages in an evergreen forest in Australia. The competing parasitic hymenoptera create a delayed regulation system for rapidly multiplying psyllid pests, where a rapid increase in the psyllid population is allowed until the pest reaches its maximum number. A short maximum is followed by a rapid decline in numbers, but minimization does not become critical for the population. The paper proposes a phenomenological model based on a differential equation with a delay, which describes a scenario of adaptive regulation for a population with a high reproductive potential with an active, but with a delayed reaction with a threshold regulation of exposure. It is shown that the complication of the regulation function of biotic resistance in the model leads to the stabilization of the dynamics after the passage of the minimum number by the rapidly breeding species. For a flexible system, transitional regimes of growth and crisis lead to the search for a new equilibrium in the evolutionary confrontation.
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