Age structure effects in predator-prey interactions

A mathematical model of predator-prey interactions is proposed which incorporates both age structure in the predators and density dependence in the prey. The properties of the model are investigated by a linearized analysis, which enables the conditions for stability to be formulated. The analysis indicates that for a substantial domain of parameter space, a stable equilibrium is possible with the prey well below its carrying capacity. The effect of violating the stability conditions on the behaviour of the model was investigated by computer simulation. Two further types of behaviour are noted in which coexistence is possible. The first is a two point limit cycle in which young and old predators occur in alternate time periods. The second involves a limit cycle in which the component population trajectories lie on closed curves in phase space.     

具常数投放率功能反应为／x的食饵-捕食系统的定性分析

本文研究一类具有常数投放率的食饵-捕食系统的定性行为,得到了正平衡点全局渐近稳定以及在正平衡点周围存在唯一极限环的充分条件．利用数值模拟检验了结论．     

A new explicit stability criterion for human periodic breathing<--RID="*"--><--ID="*" This research was supported by grants from DRED (96-920).-->

The aim of this paper is to carry out a stability analysis for periodic breathing in humans that incorporates the dynamic characteristics of ventilation control. A simple CO₂ model that takes into account the main elements of the respiratory system, i.e. the lungs and the ventilatory controller with its dynamic properties, is presented. This model results in a three-dimensional non-linear delay differential system for which there exists a unique equilibrium point. Our stability analysis of this equilibrium point leads to the definition of a new explicit stability criterion and to the demonstration of the existence of a Hopf bifurcation. Numerical simulations illustrate the influence of physiological parameters on the stability of ventilation, and particularly the major role of the dynamic characteristics of the respiratory controller. Received: 2 February 1999 / Revised version: 18 June 1999 / Published online: 23 October 2000     

Temperature-mediated stability of the interaction between spider mites and predatory mites in orchards

The nonlinear behavior of the Holling-Tanner predatory-prey differential equation system, employed by R.M. May to illustrate the apparent robustness of Kolmogorov’s Theorem when applied to such exploitation systems, is re-examined by means of the numerical bifurcation code AUTO 86 with model parameters chosen appropriately for a temperature-dependent mite interaction on fruit trees. The most significant result of this analysis is that there exists a temperature range wherein multiple stable states can occur, in direct violation of May’s interpretation of this system’s satisfaction of Kolmogorov’s Theorem: namely, that linear stability predictions have global consequences. In particular these stable states consist of a focus (spiral point) and a limit cycle separated from each other in the phase plane by an unstable limit cycle, all of which are associated with the single community equilibrium point of the system. The ecological implications of such metastability, hysteresis, and threshold behavior for the occurrence of outbreaks, the persistence of oscillations, the resiliency of the system, and the biological control of mite populations are discussed.     

具有变消耗率微生物连续培养模型的定性分析

研究了一类具有变消耗率的微生物连续培养系统,当消耗率是线性函数时得到了正平衡点全局渐近稳定的充要条件,当消耗率是二次函数时得到了系统存在极限环的充分条件,同时利用分支理论研究系统存在Hopf分支的条件,判定了极限环的稳定性．     

具有捕食正效应的捕食-食饵系统

在经典的捕食食饵系统中考虑到由于捕食效应对食饵种群带来的正向调节作用后,提出了具有捕食正效应的捕食-食饵系统.通过对模型的动力学行为的分析,从理论上说明了正向调节作用对系统的影响,并就第一象限内平衡点存在时的相图解释了捕食正效应的作用.结果表明:(1)捕食系统中适当的正向调节作用会增加系统的稳定性;(2)当捕食正效应达到一定的程度后系统拥有一个不稳定的极限环;(3)当捕食正效应过大时会使系统的稳定性发生变化,使捕食者种群与食饵种群同时趋向无穷,出现了调节放纵现象.这些结果在保护生物学中具有重要的意义.     

The elephant problem–an alternative hypothesis

Published hypotheses to account for habitat changes wrought by elephants begin from the assumption that elephant-forest systems possess a stable equilibrium point. The ‘elephant problem’ is conceived as a displacement of this equilibrium by man. Controversy centres around which human activities caused the dislocation of equilibrium and by which mechanisms these activities resulted in local high densities of the elephant Loxodonta africana. A study on elephant-forest relationships in the Luangwa Valley of Zambia casts doubt upon the basic assumption of these hypotheses and an alternative hypothesis is therefore offered. It begins from the opposite assumption–that there is no attainable natural equilibrium between elephants and forests in eastern and southern Africa. The relationship is viewed instead as a stable limit cycle in which elephants increase while thinning the forest and decline until reaching a low density that allows resurgence of the forest. This in turn triggers an increase of elephants and the cycle repeats. The period of the cycle, if the hypothesis is correct, is in the order of 200 years in the Luangwa Valley. The activities of man can impose an artificial equilibrium on the system such that trees and elephants are trapped at the low density phase of the cycle. When interference is relaxed, as with the conversion of an area to a national park, the cyclic relationship reasserts. The parameters of a system possessing a stable limit cycle need not differ in kind or interrelationship from those of a system with a stable equilibrium. Whether one or other outcome manifests may depend only on the numerical values of the parameters. If the elephant-forest system is characterized by a stable limit cycle the period and amplitude should change along a climatic gradient and may contract to a stable equilibrium in some climatic zones. A set of predictions is offered to facilitate rejection of the hypothesis.     

Analysis of SIR epidemic models with nonlinear incidence rate and treatment

This paper deals with the nonlinear dynamics of a susceptible-infectious-recovered (SIR) epidemic model with nonlinear incidence rate, vertical transmission, vaccination for the newborns of susceptible and recovered individuals, and the capacity of treatment. It is assumed that the treatment rate is proportional to the number of infectives when it is below the capacity and constant when the number of infectives reaches the capacity. Under some conditions, it is shown that there exists a backward bifurcation from an endemic equilibrium, which implies that the disease-free equilibrium coexists with an endemic equilibrium. In such a case, reducing the basic reproduction number less than unity is not enough to control and eradicate the disease, extra measures are needed to ensure that the solutions approach the disease-free equilibrium. When the basic reproduction number is greater than unity, the model can have multiple endemic equilibria due to the effect of treatment, vaccination and other parameters. The existence and stability of the endemic equilibria of the model are analyzed and sufficient conditions on the existence and stability of a limit cycle are obtained. Numerical simulations are presented to illustrate the analytical results.     

时滞传染病模型的指数稳定性

利用分析技巧研究了一类SEIRS传染病模型的动力学行为.结论表明如果再生数小于1,则带变时滞的传染病模型的无病平衡点是全局指数渐近稳定的,如果再生数大于1,得到传染病平衡点局部指数稳定的充分条件,同时给出了例子说明结论的有效性.     

Multiple Limit Cycles in a Gause Type Predator–Prey Model with Holling Type III Functional Response and Allee Effect on Prey

This work aims to examine the global behavior of a Gause type predator–prey model considering two aspects: (i) the functional response is Holling type III and, (ii) the prey growth is affected by the Allee effect. We prove the origin of the system is an attractor equilibrium point for all parameter values. It has also been shown that it is the ω-limit of a wide set of trajectories of the system, due to the existence of a separatrix curve determined by the stable manifold of the equilibrium point (m,0), which is associated to the Allee effect on prey. When a weak Allee effect on the prey is assumed, an important result is obtained, involving the existence of two limit cycles surrounding a unique positive equilibrium point: the innermost cycle is unstable and the outermost stable. This property, not yet reported in models considering a sigmoid functional response, is an important aspect for ecologists to acknowledge as regards the kind of tristability shown here: (1) the origin; (2) an interior equilibrium; and (3) a limit cycle of large amplitude. These models have undoubtedly been rather sensitive to disturbances and require careful management in applied conservation and renewable resource contexts.     

The control of ventilation: a theoretical analysis of the response to transient disturbances

Serial measurements of respiration in infants from birth to 7 months have revealed a changing pattern of response with age to spontaneous transient disturbances (e.g. deep sighs). These responses may be interpreted in terms of changing stability of the respiratory control system. The simplest possible model of respiratory control is analysed here. The control system is described by a feedback loop comprising a lung compartment, central and peripheral CO2 detectors and a circulatory delay. The differential equations representing this model are analysed in terms of the phase portrait. First we define the bounds of the solutions in the phase space and the equilibrium points. Next we discuss the local stability and damping of the system under small displacements from these equilibrium points. This local stability can be described in terms of the solution of a linear approximation about the equilibrium points. The trajectories of the solution vector under larger displacements from the equilibrium points are discussed in terms of Taylor expansions. It is shown by this analyses that this simple model can account qualitatively for the observed patterns of respiration following a deep sigh and the changes in that response from birth to 7 months.     

Global behaviour of a predator–prey like model with piecewise constant arguments

The present study deals with the analysis of a predator–prey like model consisting of system of differential equations with piecewise constant arguments. A solution of the system with piecewise constant arguments leads to a system of difference equations which is examined to study boundedness, local and global asymptotic behaviour of the positive solutions. Using Schur–Cohn criterion and a Lyapunov function, we derive sufficient conditions under which the positive equilibrium point is local and global asymptotically stable. Moreover, we show numerically that periodic solutions arise as a consequence of Neimark-Sacker bifurcation of a limit cycle.     

Global Hopf bifurcation analysis of an susceptible-infective-removed epidemic model incorporating media coverage with time delay

An susceptible-infective-removed epidemic model incorporating media coverage with time delay is proposed. The stability of the disease-free equilibrium and endemic equilibrium is studied. And then, the conditions which guarantee the existence of local Hopf bifurcation are given. Furthermore, we show that the local Hopf bifurcation implies the global Hopf bifurcation after the second critical value of delay. The obtained results show that the time delay in media coverage can not affect the stability of the disease-free equilibrium when the basic reproduction number is less than unity. However, the time delay affects the stability of the endemic equilibrium and produces limit cycle oscillations while the basic reproduction number is greater than unity. Finally, some examples for numerical simulations are included to support the theoretical prediction.     

Floquet theory: a useful tool for understanding nonequilibrium dynamics

Many ecological systems experience periodic variability. Theoretical investigation of population and community dynamics in periodic environments has been hampered by the lack of mathematical tools relative to equilibrium systems. Here, I describe one such mathematical tool that has been rarely used in the ecological literature but has widespread use: Floquet theory. Floquet theory is the study of the stability of linear periodic systems in continuous time. Floquet exponents/multipliers are analogous to the eigenvalues of Jacobian matrices of equilibrium points. In this paper, I describe the general theory, then give examples to illustrate some of its uses: it defines fitness of structured populations, it can be used for invasion criteria in models of competition, and it can test the stability of limit cycle solutions. I also provide computer code to calculate Floquet exponents and multipliers. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Costly dispersal can destabilize the homogeneous equilibrium of a metapopulation

I investigate the stability of the homogeneous equilibrium of a discrete-time metapopulation assuming costly dispersal with arbitrary (but fixed) spatial pattern of connectivity between the local populations. First, I link the stability of the metapopulation to the stability of a single isolated population by proving that the homogeneous metapopulation equilibrium, provided that it exists, is stable if and only if a single population, which is subject to extra mortality matching the average dispersal-induced mortality of the metapopulation, has a stable fixed point. Second, I demonstrate that extra mortality may destabilize the fixed point of a single population. Taken together, the two results imply that costly dispersal can destabilize the homogeneous equilibrium of a metapopulation. I illustrate this by simulations and discuss why earlier work, arriving at the opposite conclusion, was flawed.     

Chaotic dynamics of a three species prey-predator competition model with bionomic harvesting due to delayed environmental noise as external driving force

We consider a biological economic model based on prey-predator interactions to study the dynamical behaviour of a fishery resource system consisting of one prey and two predators surviving on the same prey. The mathematical model is a set of first order non-linear differential equations in three variables with the population densities of one prey and the two predators. All the possible equilibrium points of the model are identified, where the local and global stabilities are investigated. Biological and bionomical equilibriums of the system are also derived. We have analysed the population intensities of fluctuations i.e., variances around the positive equilibrium due to noise with incorporation of a constant delay leading to chaos, and lastly have investigated the stability and chaotic phenomena with a computer simulation.     

A delayed prey-predator system with parasitic infection

This paper analyzes a prey-predator system in which some members of the prey population and all predators are subjected to infection by a parasite. The predator functional response is a function of a weighted sum of prey abundances. Persistence and extinction criteria are derived. The stability of the interior equilibrium point is discussed. The role of delay is also addressed. Lastly the results are verified through computer simulation. Numerical simulation suggests that the delay has a destabilizing effect.     

基于生态位构建的2-物种集合种群关系动力学分析

结合生态位构建对两局域种群的生存竞争关系的影响,建立了基于生态位构建的2-物种集合种群关系模型,得到了共存平衡点的唯一性和全局稳定性条件,分析了两种群共存过程中空间混沌现象的存在性,并作了实例仿真加以验证．     

Multiple attractors and boundary crises in a tri-trophic food chain

The asymptotic behaviour of a model of a tri-trophic food chain in the chemostat is analysed in detail. The Monod growth model is used for all trophic levels, yielding a non-linear dynamical system of four ordinary differential equations. Mass conservation makes it possible to reduce the dimension by 1 for the study of the asymptotic dynamic behaviour. The intersections of the orbits with a Poincaré plane, after the transient has died out, yield a two-dimensional Poincaré next-return map. When chaotic behaviour occurs, all image points of this next-return map appear to lie close to a single curve in the intersection plane. This motivated the study of a one-dimensional bi-modal, non-invertible map of which the graph resembles this curve. We will show that the bifurcation structure of the food chain model can be understood in terms of the local and global bifurcations of this one-dimensional map. Homoclinic and heteroclinic connecting orbits and their global bifurcations are discussed also by relating them to their counterparts for a two-dimensional map which is invertible like the next-return map. In the global bifurcations two homoclinic or two heteroclinic orbits collide and disappear. In the food chain model two attractors coexist; a stable limit cycle where the top-predator is absent and an interior attractor. In addition there is a saddle cycle. The stable manifold of this limit cycle forms the basin boundary of the interior attractor. We will show that this boundary has a complicated structure when there are heteroclinic orbits from a saddle equilibrium to this saddle limit cycle. A homoclinic bifurcation to a saddle limit cycle will be associated with a boundary crisis where the chaotic attractor disappears suddenly when a bifurcation parameter is varied. Thus, similar to a tangent local bifurcation for equilibria or limit cycles, this homoclinic global bifurcation marks a region in the parameter space where the top-predator goes extinct. The 'Paradox of Enrichment' says that increasing the concentration of nutrient input can cause destabilization of the otherwise stable interior equilibrium of a bi-trophic food chain. For a tri-trophic food chain enrichment of the environment can even lead to extinction of the highest trophic level.