On the bifurcation analysis of a food web of four species

This paper is concerned with bifurcations of equilibria and the chaotic dynamics of a food web containing a bottom prey X, two competing predators Y and Z on X, and a super-predator W only on Y. Conditions for the existence of all equilibria and the stability properties of most equilibria are derived. A two-dimensional bifurcation diagram with the aid of a numerical method for identifying bifurcation curves is constructed to show the bifurcations of equilibria. We prove that the dynamical system possesses a line segment of degenerate steady states for the parameter values on a bifurcation line in the bifurcation diagram. Numerical simulations show that these degenerate steady states can help to switch the stabilities between two far away equilibria when the system crosses this bifurcation line. Some observations concerned with chaotic dynamics are also made via numerical simulations. Different routes to chaos are found in the system. Relevant calculations of Lyapunov exponents and power spectra are included to support the chaotic properties.     

NONLINEAR MATRIX MODELS AND POPULATION DYNAMICS

Nonlinear matrix difference equations are studied as models for the discrete time dynamics of a population whose individual members have been categorized into a finite number of classes. The equations are treated with sufficient generality so as to include virtually any type of structuring of the population (the sole constraint is that all newborns lie in the same class) and any types of nonlinearities which arise from the density dependence of fertility rates, survival rates and transition probabilities between classes. The existence and stability of equilibrium class distribution vectors are studied by means of bifurcation theory techniques using a single composite, biologically meaningful quantity as a bifurcation parameter, namely the inherent net reproductive rate r. It is shown that, just as in the case of linear matrix equations, a global continuum of positive equilibria exists which bifurcates as a function of r from the zero equilibrium state at and only at r = 1. Furthermore the zero equilibrium loses stability as r is increased through 1. Unlike the linear case however, for which the bifurcation is “vertical” (i.e., equilibria exist only for r = 1), the nonlinear equation in general has positive equilibria for an interval of r values. Methods for studying the geometry of the continuum based upon the density dependence of the net reproductive rate at equilibrium are developed. With regard to stability, it is shown that in general the positive equilibria near the bifurcation point are stable if the bifurcation is to the right and unstable if it is to the left. Some further results and conjectures concerning stability are also given. The methods are illustrated by several examples involving nonlinear models of various types taken from the literature.     

A delay differential equation model of HIV infection of CD4+ T-cells with cure rate

A delay differential equation as a mathematical model that described HIV infection of CD4⁺ T-cells is analyzed. When the constant death rate of infected but not yet virus-producing cells is equal to zero, the stability of the non-negative equilibria and the existence of Hopf bifurcation are investigated. A stability switch in the system due to variation of delay parameter has been observed, so is the phenomena of Hopf bifurcation and stable limit cycle. The estimation of the length of delay to preserve stability has been calculated. Further, when the constant death rate of infected but not yet virus-producing cells is not equal to zero, by using the geometric stability switch criterion in the delay differential system with delay dependent parameters, we present that stable equilibria become unstable as the time delay increases. Numerical simulations are carried out to explain the mathematical conclusions.     

Stability and Hopf bifurcation analysis of an eco-epidemic model with a stage structure

In this paper, an eco-epidemiological model with a stage structure is considered. The asymptotical stability of the five equilibria, the existence of stability switches about positive equilibrium, is investigated. It is found that Hopf bifurcation occurs when the delay τ passes though a critical value. Some explicit formulae determining the stability and the direction of the Hopf bifurcation periodic solutions bifurcating from Hopf bifurcations are obtained by using the normal form theory and center manifold theory. Some numerical simulations for justifying the theoretical analysis are also provided. Finally, biological explanations and main conclusions are given.     

Asymptotic behavior of an unstirred chemostat model with internal inhibitor

This paper deals with a chemostat model with an internal inhibitor. First, the elementary stability and asymptotic behavior of solutions of the system are determined. Second, the effects of the inhibitor are considered. It turns out that the parameter μ, which measures the effect of the inhibitor, plays a very important role in deciding the stability and longtime behavior of solutions of the system. The results show that if μ is sufficiently large, this model has no coexistence solution and one of the semitrivial equilibria is a global attractor when the maximal growth rate a of the species u lies in certain range; but when a belongs to another range, all positive solutions of this model are governed by a limit problem, and two semitrivial equilibria are bistable. The main tools used here include monotone system theory, degree theory, bifurcation theory and perturbation technique.     

Global Dynamics of a Tuberculosis Epidemic Model and the Influence of Backward Bifurcation

In this paper, we propose and analyze a tuberculosis (TB) model with exogenous re-infection. We assume that treated individual may be again infected by infectious individual. The model exhibits two bifurcations viz. transcritical bifurcation when the basic reproductive number R ₀?=?1 and backward bifurcation where the disease transmission rate β plays as control parameter. The persistent of the model and, the local and global stability criteria of disease-free and endemic equilibria are discussed. By carrying out bifurcation analysis, it is shown that the model exhibits the bistability and undergoes the Hopf bifurcation when immunological memory is everlasting i.e. when σ?=?0. Lastly, some simulations are given to verify our analytical results.     

A bifurcation theorem for nonlinear matrix models of population dynamics

ABSTRACT

We prove a general theorem for nonlinear matrix models of the type used in structured population dynamics that describes the bifurcation that occurs when the extinction equilibrium destabilizes as a model parameter is varied. The existence of a bifurcating continuum of positive equilibria is established, and their local stability is related to the direction of bifurcation. Our theorem generalizes existing theorems found in the literature in two ways. First, it allows for a general appearance of the bifurcation parameter (existing theorems require the parameter to appear linearly). This significantly widens the applicability of the theorem to population models. Second, our theorem describes circumstances in which a backward bifurcation can produce stable positive equilibria (existing theorems allow for stability only when the bifurcation is forward). The signs of two diagnostic quantities determine the stability of the bifurcating equilibrium and the direction of bifurcation. We give examples that illustrate these features.     

Bifurcation analysis in an age-structured model of a single species living in two identical patches

In this paper, we consider the age-structured model of a single species living in two identical patches derived in So et al. [J.W.-H. So, J. Wu, X. Zou, Structured population on two patches: modeling dispersal and delay, J. Math. Biol. 43 (2001) 37–51]. We chose a birth function that is frequently used but different from the one used in So et al. which leads to a different structure of the homogeneous equilibria. We investigate the stability of these equilibria and Hopf bifurcations by analyzing the distribution of the roots of associated characteristic equation. By the theory of normal form and center manifold, an explicit algorithm for determining the direction of the Hopf bifurcation and stability of the bifurcating periodic solutions are derived. Finally, some numerical simulations are carried out for supporting the analytic results.     

Qualitative analysis of a SIR epidemic model with saturated treatment rate

On account of the effect of limited treatment resources on the control of epidemic disease, a saturated removal rate is incorporated into Hethcote’s SIR epidemiological model (Hethcote, SIAM Rev. 42:599–653, 2000). Unlike the original model, the model has two endemic equilibria when R ₀<1. Furthermore, under some conditions, both the disease free equilibrium and one of the two endemic equilibria are asymptotically stable, i.e., the model has bistable equilibria. Therefore, disease eradication not only depends on R ₀ but also on the initial sizes of all sub-populations. By the Poincaré-Bendixson theorem, Poincaré index, center manifold theorem, Hopf bifurcation theorem and Lyapunov-Lasalle theorem, etc., the existence and asymptotical stability of the equilibria, the existence, stability and direction of Hopf bifurcation are discussed, respectively.     

A METAPOPULATION MODEL WITH DISCRETE SIZE STRUCTURE

ABSTRACT. We consider a discrete size‐structured meta‐population model with the proportions of patches occupied by n individuals as dependent variables. Adults are territorial and stay on a certain patch. The juveniles may emigrate to enter a dispersers' pool from which they can settle on another patch and become adults. Absence of colonization and absence of emigration lead to extinction of the metapopula‐tion. We define the basic reproduction number R₀ of the metapopulation as a measure for its strength of persistence. The metapopulation is uniformly weakly persistent if R₀> 1. We identify subcritical bifurcation of persistence equilibria from the extinction equilibrium as a source of multiple persistence equilibria: it occurs, e.g., when the immigration rate, into occupied patches, exceeds the colonization rate (of empty patches). We determine that the persistence‐optimal dispersal strategy which maximizes the basic reproduction number is of bang‐bang type: If the number of adults on a patch is below carrying capacity all the juveniles should stay, if it is above the carrying capacity all the juveniles should leave.     

Bifurcation analysis in a model of cytotoxic T-lymphocyte response to viral infections

In this paper, we study the dynamics of a mathematical model on primary and secondary cytotoxic T-lymphocyte (CTL) response to viral infections by Wodarz et al. This model has three equilibria and their stability criteria are discussed. The system transitions from one equilibrium to the next as the basic reproductive number, R₀, increases. When R₀ increases even further, we analytically show that periodic solutions may arise from the third equilibrium via Hopf bifurcation. Numerical simulations of the model agree with the theoretical results and these dynamics occur within biologically realistic parameter range. The normal form theory is also applied to find the amplitude, phase and stability information on the limit cycles. Biological implications of the results are discussed.     

Degenerate relative equilibria, curvature of the momentum map, and homoclinic bifurcation

A fundamental class of solutions of symmetric Hamiltonian systems is relative equilibria. In this paper the nonlinear problem near a degenerate relative equilibrium is considered. The degeneracy creates a saddle-center and attendant homoclinic bifurcation in the reduced system transverse to the group orbit. The surprising result is that the curvature of the pullback of the momentum map to the Lie algebra determines the normal form for the homoclinic bifurcation. There is also an induced directional geometric phase in the homoclinic bifurcation. The backbone of the analysis is the use of singularity theory for smooth mappings between manifolds applied to the pullback of the momentum map. The theory is constructive and generalities are given for symmetric Hamiltonian systems on a vector space of dimension (2n+2) with an n-dimensional abelian symmetry group. Examples for n=1,2,3 are presented to illustrate application of the theory.     

Stability and bifurcation in a harmonic oscillator with delays

We consider a harmonic oscillator with delays. Linear stability is investigated by analyzing the associated characteristic transcendental equation. The bifurcation analysis of the equation shows that Hopf bifurcation can occur as the delay τ (taken as a parameter) crosses some critical values. The direction and stability of the Hopf bifurcation are considered by using the normal form theory due to Faria and Magalhães. An example is given to explain the results. Numerical simulations support our results.     

Hopf bifurcation analysis of a density predator-prey model with crowley-martin functional response and two time delays

In this paper, a delayed density dependent predator-prey model with Crowley-Martin functional response and two time delays for the predator is considered. By analyzing the corresponding characteristic equations, the local stability of each of the feasible equilibria of the system is addressed and the existence of Hopf bifurcation at the coexistence equilibrium is established. With the help of normal form method and center manifold theorem, some explicit formulas determining the direction of Hopf bifurcation and the stability of bifurcating period solutions are derived. Finally, numerical simulations are given to illustrate the theoretical results.     

Analysis of a spatially extended nonlinear SEIS epidemic model with distinct incidence for exposed and infectives

We present a nonlinear SEIS epidemic model which incorporates distinct incidence rates for the exposed and the infected populations. The model is analyzed for stability and bifurcation behavior. To account for the realistic phenomenon of non-homogeneous mixing, the effect of diffusion on different population subclasses is considered. The diffusive model is analyzed using matrix stability theory and conditions for Turing bifurcation derived. Numerical simulations are performed to justify analytical findings.     

Synchronous Dynamics and Bifurcation Analysis in Two Delay Coupled Oscillators with Recurrent Inhibitory Loops

In this study, the dynamics and low-codimension bifurcation of the two delay coupled oscillators with recurrent inhibitory loops are investigated. We discuss the absolute synchronization character of the coupled oscillators. Then the characteristic equation of the linear system is examined, and the possible low-codimension bifurcations of the coupled oscillator system are studied by regarding eigenvalues of the connection matrix as bifurcation parameter, and the bifurcation diagram in the γ–ρ plane is obtained. Applying normal form theory and the center manifold theorem, the stability and direction of the codimension bifurcations are determined. Moreover, the symmetric bifurcation theory and representation theory of Lie groups are used to investigate the spatio-temporal patterns of the periodic oscillations. Finally, numerical results are applied to illustrate the results obtained.     

一类三维神经元模型的分支研究

考虑一类三维神经元模型的分支问题.利用常微分方程的定性与分支理论的知识,讨论了模型的平衡点个数及其稳定性,主要分析了平衡点的Hopf分支和Bogdanov-Takens分支,并得到了相应的鞍结点分支曲线,Hopf分支曲线与同宿分支曲线.     

Dynamics of a delayed predator–prey model with predator migration

Based on the availability of prey and a simple predator–prey model, we propose a delayed predator–prey model with predator migration to describe biological control. We first study the existence and stability of equilibria. It turns out that backward bifurcation occurs with the migration rate as bifurcation parameter. The stability of the trivial equilibrium and the boundary equilibrium is delay-independent. However, the stability of the positive equilibrium may be delay-dependent. Moreover, delay can switch the stability of the positive equilibrium. When the positive equilibrium loses stability, Hopf bifurcation can occur. The direction and stability of Hopf bifurcation is derived by applying the center manifold method and the normal form theory. The main theoretical results are illustrated with numerical simulations.     

The effect of population dispersal on the spread of a disease

The effect of population dispersal among n patches on the spread of a disease is investigated. Population dispersal does not destroy the uniqueness of a disease free equilibrium and its attractivity when the basic reproduction number of a disease R₀<1. When R₀>1, the uniqueness and global attractivity of the endemic equilibrium can be obtained if dispersal rates of susceptible individuals and infective individuals are the same or very close in each patch. However, numerical calculations show that population dispersal may result in multiple endemic equilibria and even multi-stable equilibria among patches, and also may result in the extinction of a disease, even though it cannot be eradicated in each isolated patch, provided the basic reproduction numbers of isolated patches are not very large.     

Stability and Hopf bifurcation analysis in a three-level food chain system with delay

A class of three level food chain system is studied. With the theory of delay equations and Hopf bifurcation, the conditions of the positive equilibrium undergoing Hopf bifurcation is given regarding τ as the parameter. The stability and direction of Hopf bifurcation are determined by applying the normal form theory and the center manifold argument, and numerical simulations are performed to illustrate the analytical results.