Multiple limit cycles in the standard model of three species competition for three essential resources

We consider the dynamics of the standard model of 3 species competing for 3 essential (non-substitutable) resources in a chemostat using Liebig's law of the minimum functional response. A subset of these systems which possess cyclic symmetry such that its three single-population equilibria are part of a heteroclinic cycle bounding the two-dimensional carrying simplex is examined. We show that a subcritical Hopf bifurcation from the coexistence equilibrium together with a repelling heteroclinic cycle leads to the existence of at least two limit cycles enclosing the coexistence equilibrium on the carrying simplex- the ``inside' one is an unstable separatrix and the ``outside' one is at least semi-stable relative to the carrying simplex. Numerical simulations suggest that there are exactly two limit cycles and that almost every positive solution approaches either the stable limit cycle or the stable coexistence equilibrium, depending on initial conditions. Bifurcation diagrams confirm this picture and show additional features. In an alternative scenario, we show that the subcritical Hopf together with an attracting heteroclinic cycle leads to an unstable periodic orbit separatrix. This research was partially supported by NSF grant DMS 0211614. KY 40292, USA. This author's research was supported in part by NSF grant DMS 0107160     

Bifurcation structure of a chemostat model for an age-structured predator and its prey

We model a chemostat containing an age-structured predator and its prey using a linear function for the uptake of substrate by the prey and two different functional responses (linear and Monod) for the consumption of prey by the predator. Limit cycles (LCs) caused by the predator's age structure arise at Hopf bifurcations at low values of the chemostat dilution rate for both model cases. In addition, LCs caused by the predator–prey interaction arise for the case with the Monod functional response. At low dilution rates in the Monod case, the age structure causes cycling at lower values of the inflowing resource concentration and conversely prevents cycling at higher values of the inflowing resource concentration. The results shed light on a similar model by Fussmann et al. [G. Fussmann, S. Ellner, K. Shertzer, and N. Hairston, Crossing the Hopf bifurcation in a live predator–prey system, Science 290 (2000), pp. 1358–1360.], which correctly predicted conditions for the onset of cycling in a chemostat containing an age-structured rotifer population feeding on algal prey.     

具有抑制剂的恒化器竞争模型的分析（英文）

考虑了具有抑制剂的两种群竞争一种微生物的恒化器模型,其中吸收函数和功能反应函数都是营养的一般单调递增函数,并且一种微生物能够分泌一种对另一种微生物起致命影响的抑制剂.利用常微分方程定性理论,首先得到了平衡点的存在条件和局部渐进稳定性;然后讨论了全局渐进稳定性,以及极限环和Hopf分支的存在性.     

具有稀疏效应的Predator-Prey模型的分支问题

讨论了具有稀疏效应的捕食－食饵模型的分支问题,并利用Ｈｏｐｆ分支理论和分界线环分支理论,得到了有多个极限环的结果。     

Stability and Hopf bifurcation for a prey-predator model with prey-stage structure and diffusion

In this paper, we first propose a prey-predator model with prey-stage structure and diffusion. Then we discuss the following three problems: (1) stability of non-negative constant steady states for the reduced ODE system and the corresponding reaction diffusion system with homogeneous Neumann boundary conditions; (2) Hopf bifurcation for the ODE system; (3) Hopf bifurcation created by diffusion.     

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．     

Transient oscillations induced by delayed growth response in the chemostat

In this paper, in order to try to account for the transient oscillations observed in chemostat experiments, we consider a model of single species growth in a chemostat that involves delayed growth response. The time delay models the lag involved in the nutrient conversion process. Both monotone response functions and nonmonotone response functions are considered. The nonmonotone response function models the inhibitory effects of growth response of certain nutrients when concentrations are too high. By applying local and global Hopf bifurcation theorems, we prove that the model has unstable periodic solutions that bifurcate from unstable nonnegative equilibria as the parameter measuring the delay passes through certain critical values and that these local periodic solutions can persist, even if the delay parameter moves far from the critical (local) bifurcation values.When there are two positive equilibria, then positive periodic solutions can exist. When there is a unique positive equilibrium, the model does not have positive periodic oscillations and the unique positive equilibrium is globally asymptotically stable. However, the model can have periodic solutions that change sign. Although these solutions are not biologically meaningful, provided the initial data starts close enough to the unstable manifold of one of these periodic solutions they may still help to account for the transient oscillations that have been frequently observed in chemostat experiments. Numerical simulations are provided to illustrate that the model has varying degrees of transient oscillatory behaviour that can be controlled by the choice of the initial data.Mathematics Subject Classification: 34D20, 34K20, 92D25Research was partially supported by NSERC of Canada.This work was partly done while this author was a postdoc at McMaster.     

A predator prey model with age structure

A general predator-prey model is considered in which the predator population is assumed to have an age structure which significantly affects its fecundity. The model equations are derived from the general McKendrick equations for age structured populations. The existence, stability and destabilization of equilibria are studied as they depend on the prey's natural carrying capacity and the maturation periodm of the predator. The main result of the paper is that for a broad class of maturation functions positive equilibria are either unstable for smallm or are destabilized asm decreases to zero. This is in contrast to the usual rule of thumb that increasing (not decreasing) delays in growth rate responses cause instabilities.Research supported by National Science Foundation Grant No. MCS-7901307-01Research supported by National Scholarship for Study Abroad No. EDN/S-59/80 from the government of India     

The periodically forced Droop model for phytoplankton growth in a chemostat

 It is proved that the periodically forced Droop model for phytoplankton growth in a chemostat has precisely two dynamic regimes depending on a threshold condition involving the dilution rate. If the dilution rate is such that the sub-threshold condition holds, the phytoplankton population is washed out of the chemostat. If the super-threshold condition holds, then there is a unique periodic solution, having the same period as the forcing, characterized by the presence of the phytoplankton population, to which all solutions approach asymptotically. Furthermore, this result holds for a general class of models with monotone growth rate and monotone uptake rate, the latter possibly depending on the cell quota. Received 10 October 1995; received in revised form 26 March 1996     

具Holling-Ⅳ型功能反应函数的捕食者-食饵系统的定性分析(英文)

本文研究一类具Holling-Ⅳ型功能反应函数的捕食者-食饵模型.对模型进行定性分析得知系统正解都是有界的;因此,当平衡点不稳定,系统至少存在一稳定的极限环.本文还运用Poincare形式级数法,得到了正平衡点至多为二阶稳定细焦点的结论.并基于Hopf分支理论得知系统在一定条件下至少存在两个极限环.     

Long run coexistence in the chemostat with multiple species

In this work we analyze the transient behavior of the dynamics of multiple species competing in a chemostat for a single resource, presenting slow/fast characteristics. We prove that coexistence among a subset of species, with growth functions close to each other, can last for a substantially long time. For these cases, we also show that the proportion of non-dominant species can be increasing before decreasing, under certain conditions on the initial distribution.     

Competing predators for a prey in a chemostat model with periodic nutrient input

A system of ordinary differential equations is used to model the interactions of n competing predators on a single prey population in a chemostat environment with a periodic nutrient input. In the case of one or no predators, criteria for the existence of periodic solutions are given. In the general case, conditions for all populations to persist are derived.Research is in part based on a Ph.D. thesis submitted to the Faculty of Graduate Studies, University of AlbertaResearch is partly supported by the Natural Sciences and Engineering Research Council of Canada, Grant No. NSERC A4823     

Oscillatory coexistence in a food chain model with competing predators

A food chain model with two predators feeding on a single prey in a chemostat is studied. Using a multiparameter bifurcation analysis, we find parameters values for which there is stable oscillatory coexistence of the predators. It is also shown how these coexistent states provide a transition between two possible states of competitive exclusion. It is shown that the competitive exclusion principle need not hold if one or more of the predators has oscillatory behavior in the absence of other predators.This work was partially supported by National Science Foundation Grant MCS 83-01881     

Global analysis of a model of plasmid-bearing,plasmid-free competition in a chemostat

A model of competition between plasmid-bearing and plasmid-free organisms in a chemostat was proposed in a paper of Stephanopoulis and Lapidus. The model was in the form of a system of nonlinear ordinary differential equations. Such models are relevant to commercial production by genetically altered organisms in continuous culture. The analysis there was local (using index arguments). This paper provides a mathematically rigorous analysis of the global asymptotic behavior of the governing equations in the case of uninhibited specific growth rate.Research supported by the National Council of Science, Republic of ChinaResearch supported by National Science Foundation Grant, DMS-9204490Research supported by the Natural Science and Engineering Council of Canada. This author's contribution was made while on research leave visiting the Department of Ecology and Evolutionary Biology at Princeton University. She would especially like to thank Simon Levin for his guidance as well as for providing an exceptional working environment     

Global analysis of a model of plasmid-bearing,plasmid-free competition in a chemostat with inhibitions

A model of competition between plasmid-bearing and plasmid-free organisms in a chemostat was proposed in a paper of Stephanopoulis and Lapidus. The model was in the form of a system of nonlinear ordinary differential equations. Such models were relevant to commercial production by genetically altered organisms in continuous culture. The analysis there was local. The rigorous global analysis was done in a paper of Hsu, Waltman and Wolkowicz in the case of the uninhibited specific growth rates. This paper provides a mathematically rigorous analysis of the global asymptotic behavior of the governing equations in the cases of combinations of inhibited and uninhibited specific growth rates.Research Supported by the National Council of Science, Republic of China     

Global dynamics of a chemostat competition model with distributed delay

 We study the global dynamics of n-species competition in a chemostat with distributed delay describing the time-lag involved in the conversion of nutrient to viable biomass. The delay phenomenon is modelled by the gamma distribution. The linear chain trick and a fluctuation lemma are applied to obtain the global limiting behavior of the model. When each population can survive if it is cultured alone, we prove that at most one competitor survives. The winner is the population that has the smallest delayed break-even concentration, provided that the orders of the delay kernels are large and the mean delays modified to include the washout rate (which we call the virtual mean delays) are bounded and close to each other, or the delay kernels modified to include the washout factor (which we call the virtual delay kernels) are close in L ¹-norm. Also, when the virtual mean delays are relatively small, it is shown that the predictions of the distributed delay model are identical with the predictions of the corresponding ODEs model without delay. However, since the delayed break-even concentrations are functions of the parameters appearing in the delay kernels, if the delays are sufficiently large, the prediction of which competitor survives, given by the ODEs model, can differ from that given by the delay model. Received: 9 August 1997 / Revised version: 2 July 1998     

An epidemiological model with a delay and a nonlinear incidence rate

An epidemiological model with both a time delay in the removed class and a nonlinear incidence rate is analysed to determine the equilibria and their stability. This model is for diseases where individuals are first susceptible, then infected, then removed with temporary immunity and then susceptible again when they lose their immunity. There are multiple equilibria for some parameter values, and, for certain of these, periodic solutions arise by Hopf bifurcation from the large nontrivial equilibrium state.Research supported in parts by Centers for Disease Control Contract 200-87-0515Research supported in part by NSERC A-8965     

The asymptotic behavior of a chemostat model with the Beddington-DeAngelis functional response

In this paper, the global asymptotic behavior of a chemostat model with Beddington-DeAngelis functional response is studied. The conditions for the global asymptotical stability of the model with time delays are obtained via monotone dynamical systems. Our results demonstrate that those time delays affect the competitive outcome of the organisms.     

Allee dynamics generated by protection mutualisms can drive oscillations in trophic cascades

Understanding the relative effect of top predators and primary producers on intermediate trophic levels is a key question in ecology. Most previous work, however, has not considered either realistic nonlinearities in feedback between trophic levels or the effect of mutualists on trophic cascades. Here, we develop a realistic model for a protection mutualism that explicitly includes interactions between a protected herbivore and both its food plant and generalist predators. In the absence of protection, herbivores and plant resources approach a stable equilibrium, provided that predation is not so high as to cause herbivore extinction. In contrast, adding protection by mutualists increases the range of dynamical outcomes to include unstable equilibria, stable and unstable limit cycles, and heteroclinic orbits. By reducing the impact of predators, protection by mutualists can allow herbivores to exert strong negative effects on their host plants, which in turn can lead to repeated cycles of overexploitation and recovery. Our results indicate that it may be essential to consider protection mutualisms to understand the dynamics of trophic cascades. Conversely, it may be essential to explicitly include dynamical feedback between plants and herbivores to fully understand the population and community dynamical consequences of protection mutualism.     

Nonlinear control for algae growth models in the chemostat

This paper deals with output feedback control of phytoplanktonic algae growth models in the chemostat. The considered class of model is of variable yield type, meaning that the ratio between the environmental nutrient absorption rate and the cells’ growth rate varies, which is different from classical bioprocesses assumptions. On the basis of weak qualitative hypotheses on the analytical expressions of the involved biological phenomena (which guarantee robustness of the procedure toward modeling uncertainties) we propose a nonlinear controller and prove its ability to globally stabilize such processes. Finally, we illustrate our approach with numerical simulations and show its benefits for biological laboratory experiments, especially for ensuring persistence of the culture facing classical experimental problems.