Competition in size-structured populations: mechanisms inducing cohort formation and population cycles

In this paper we investigate the consequences of size-dependent competition among the individuals of a consumer population by analyzing the dynamic properties of a physiologically structured population model. Only 2 size-classes of individuals are distinguished: juveniles and adults. Juveniles and adults both feed on one and the same resource and hence interact by means of exploitative competition. Juvenile individuals allocate all assimilated energy into development and mature on reaching a fixed developmental threshold. The combination of this fixed threshold and the resource-dependent developmental rate, implies that the juvenile delay between birth and the onset of reproduction may vary in time. Adult individuals allocate all assimilated energy to reproduction. Mortality of both juveniles and adults is assumed to be inversely proportional to the amount of energy assimilated. In this setting we study how the dynamics of the population are influenced by the relative foraging capabilities of juveniles and adults.In line with results that we previously obtained in size-structured consumer-resource models with pulsed reproduction, population cycles primarily occur when either juveniles or adults have a distinct competitive advantage. When adults have a larger per capita feeding rate and are hence competitively superior to juveniles, population oscillations occur that are primarily induced by the fact that the duration of the juvenile period changes with changing food conditions. These cycles do not occur when the juvenile delay is a fixed parameter. When juveniles are competitively superior, two different types of population fluctuations can occur: (1) rapid, low-amplitude fluctuations having a period of half the juvenile delay and (2) slow, large-amplitude fluctuations characterized by a period, which is roughly equal to the juvenile delay. The analysis of simplified versions of the structured model indicates that these two types of oscillations also occur if mortality and/or development is independent of food density, i.e. in a situation with a constant juvenile developmental delay and a constant, food-independent background mortality. Thus, the oscillations that occur when juveniles are more competitive are induced by the juvenile delay per se. When juveniles exert a larger foraging pressure on the shared resource, maturation implies an increase not only in adult density, but also in food density and consequently fecundity. Our analysis suggests that this correlation in time between adult density and fecundity is crucial for the occurrence of population cycles when juveniles are competitively superior.     

A spatially discrete model for aggregating populations

 In this paper we introduce a spatially discrete model for aggregating populations described by a system of ODEs. We study the long time behavior of the solutions and we show that the model contains mechanisms by which individuals in the population aggregate at particular points in space. Received: 29 June 1996 / Revised version: 5 August 1997     

A deterministic size-structured population model for the worm Naidis elinguis

In this paper we model the population dynamics of the worm Nais elinguis, which reproduces by division into two unequal parts. By using renewal theory we derive the asymptotic behaviour of a Naidis elinguis population. In particular we prove a certain relation between the fraction of the population that was born small (respectively the fraction that was born large) and the inter-division times. Received 20 January 1999 / Revised version: 1 August 1999?Published online: 10 April 2001     

Periodic solutions: a robust numerical method for an S-I-R model of epidemics

 We describe and analyze a numerical method for an S-I-R type epidemic model. We prove that it is unconditionally convergent and that solutions it produces share many qualitative and quantitative properties of the solution of the differential problem being approximated. Finally, we establish explicit sufficient conditions for the unique endemic steady state of the system to be unstable and we use our numerical algorithm to approximate the solution in such cases and discover that it can be periodic, just as suggested by the instability of the endemic steady state. Received: 1 September 1995 / Revised version: 30 April 1997     

Interaction of maturation delay and nonlinear birth in population and epidemic models

 A population with birth rate function B(N) N and linear death rate for the adult stage is assumed to have a maturation delay T>0. Thus the growth equation N′(t)=B(N(t−T)) N(t−T) e⁻ d ₁ T−dN(t) governs the adult population, with the death rate in previous life stages d ₁≧0. Standard assumptions are made on B(N) so that a unique equilibrium N _e exists. When B(N) N is not monotone, the delay T can qualitatively change the dynamics. For some fixed values of the parameters with d ₁>0, as T increases the equilibrium N _e can switch from being stable to unstable (with numerically observed periodic solutions) and then back to stable. When disease that does not cause death is introduced into the population, a threshold parameter R ₀ is identified. When R ₀<1, the disease dies out; when R ₀>1, the disease remains endemic, either tending to an equilibrium value or oscillating about this value. Numerical simulations indicate that oscillations can also be induced by disease related death in a model with maturation delay. Received: 2 November 1998 / Revised version: 26 February 1999     

Multiparametric bifurcations for a model in epidemiology

 In the present paper we make a bifurcation analysis of an SIRS epidemiological model depending on all parameters. In particular we are interested in codimension-2 bifurcations. Received 8 April 1994; received in revised form 29 June 1995     

A model for dengue disease with variable human population

 A model for the transmission of dengue fever with variable human population size is analyzed. We find three threshold parameters which govern the existence of the endemic proportion equilibrium, the increase of the human population size, and the behaviour of the total number of human infectives. We prove the global asymptotic stability of the equilibrium points using the theory of competitive systems, compound matrices, and the center manifold theorem. Received: 3 November 1997 / Revised version: 3 July 1998     

Periodic solutions for a population dynamics problem with age-dependence and spatial structure

Using a linear model with age-dependence and spatial structure we show how a periodical supply of individuals will transform an exponentially decaying distribution of population into a non-trivial asymptotically stable periodic distribution. Next we give an application to an epidemic model.     

Periodic solutions in a model of competition between plasmid-bearing and plasmid-free organisms in a chemostat with an inhibitor

We obtain necessary and sufficient conditions on the existence of a unique positive equilibrium point and a set of sufficient conditions on the existence of periodic solutions for a 3-dimensional system which arises from a model of competition between plasmid-bearing and plasmid-free organisms in a chemostat with an inhibitor. Our results improve the corresponding results obtained by Hsu, Luo, and Waltman [1]. Received: 20 November 1997 / Revised version: 12 February 1999 / Published online: 20 December 2000     

Asymptotic behaviour of a model of hierarchically structured population dynamics

 A hierarchically structured population model with a dependence of the vital rates on a function of the population density (environment) is considered. The existence, uniqueness and the asymptotic behaviour of the solutions is obtained transforming the original non-local PDE of the model into a local one. Under natural conditions, the global asymptotical stability of a nontrivial equilibrium is proved. Finally, if the environment is a function of the biomass distribution, the existence of a positive total biomass equilibrium without a nontrivial population equilibrium is shown. Received 16 February 1996; received in revised form 16 September 1996     

A weakly nonlinear analysis of a model of avascular solid tumour growth

 In this paper we study a mathematical model that describes the growth of an avascular solid tumour. Our analysis concentrates on the stability of steady, radially-symmetric model solutions with respect to perturbations taken from the class of spherical harmonics. Using weakly nonlinear analysis, previous results are extended to show how the amplitudes of the asymmetric modes interact. Attention focuses on a special case for which the model equations simplify. Analysis of the simplified model equations leads to the identification of a two-parameter family of asymmetric steady solutions, the dimensions of whose stable and unstable manifolds depend on the system parameters. The asymmetric steady solutions limit the basin of attraction of the radially-symmetric steady state when it is linearly stable. On the basis of these numerical and analytical results we postulate the existence of fully nonlinear steady solutions which are stable with respect to time-dependent perturbations. Received: 25 October 1998 / Revised version: 20 June 1998     

On a two-phase size-structured population model with infinite states-at-birth and distributed delay in birth process

In this paper we study the two-phase size-structured population model with infinite states-at-birth and distributed delay in birth process. The model distinguishes individuals by two different status: the ‘reproductive’ stage and the ‘nonreproductive’ stage. We establish the well-posedness for this model and show that the solution of this model exhibits asynchronous exponential growth by means of semigroups. We also consider a special case in which the individuals in the ‘reproductive’ stage and the ‘nonreproductive’ stage have the same growth rates and give a comparison between this two-phase model with the classical one-phase model.     

Analytical solutions of a minimal model of species migration in a bounded domain

 A minimal model of species migration is presented which takes the form of a parabolic equation with boundary conditions and initial data. Solutions to the differential problem are obtained that can be used to describe the small- and large-time evolution of a species distribution within a bounded domain. These expressions are compared with the results of numerical simulations and are found to be satisfactory within appropriate temporal regimes. The solutions presented can be used to describe existing observations of nematode distributions, can be used as the basis for further work on nematode migration, and may also be interpreted more generally. Received: 15 August 1999     

The dynamics of infectious diseases in orchards with roguing and replanting as control strategy

 Roguing and replanting is a widely adopted control strategy of infectious diseases in orchards. Little is known about the effect of this type of management on the dynamics of the infectious disease. In this paper we analyze a structured population model for the dynamics of an S-I-R type epidemic under roguing and replanting management. The model is structured with respect to the total number of infections and the number of post-infectious infections on a tree. Trees are assumed to be rogued, and replaced by uninfected trees, when the total number of infections on the tree reaches a threshold value. Stability analysis and numerical exploration of the model show that for specific parameter combinations the internal equilibrium can become unstable and large amplitude periodic fluctuations arise. Several hypothesis on the mechanism causing the destabilisation of the steady-state are considered. The mechanism leading to the large amplitude fluctuations is identified and biologically interpreted. Received 2 September 1994     

A mathematical model for drug administration by using the phagocytosis of red blood cells

 A mathematical model for the delivery of drug directly to the macrophages by using the phagocytosis of senescent red blood cells is proposed. The model is based on the following assumption: At time t=0 a preassigned red blood cell population n(0, a)=φ(a), a>0, loaded by the drug, is injected in the blood circulation. Among the cells of that population only those with an age a≧ā (ā=120 days) will be phagocytosed by macrophages. Of course, the lifetime of the drug must be higher than ā. Within the red blood cells it cannot be metabolized, neither can it diffuse through their membranes. The emphasis of the paper is on the mathematical properties and on the formulation of the control problem. Received 15 December 1994; received in revised form 20 July 1995     

Modelling the depletion and conservation of forestry resources: effects of population and pollution

 In this paper, a mathematical model is proposed to study the depletion of resources in a forest habitat due to the increase of both population and pollution. It is shown that if the rate of pollutant emission into the environment is either population dependent, constant, or periodic, the equilibrium biomass density of the resource settles down to a lower equilibrium than its original carrying capacity, the magnitude of which decreases as the equilibrium levels of the density of population and the concentration of pollutant increase. However, in the case of an instantaneous spill of pollutant into the environment, the equilibrium biomass density decreases with the increase of the equilibrium density of population only. It is found that if the population density and the emission rate of pollutant increase without control, the forestry resource may become extinct. A conservation model is also proposed, the analysis of which shows that the resource biomass can be maintained at a desired level by conserving the forestry resource and by controlling the growth of population and the emission rate of pollutant in the habitat. Received 1 June 1993; received in revised form 1 January 1997     

The diffusion model for migration and selection in a plant population

 The diffusion approximation is derived for migration and selection at a multiallelic locus in a partially selfing plant population subdivided into a lattice of colonies. Generations are discrete and nonoverlapping; both pollen and seeds disperse. In the diffusion limit, the genotypic frequencies at each point are those determined at equilibrium by the local rate of selfing and allelic frequencies. If the drift and diffusion coefficients are taken as the appropriate linear combination of the corresponding coefficients for pollen and seeds, then the migration terms in the partial differential equation for the allelic frequencies have the standard form for a monoecious animal population. The selection term describes selection on the local genotypic frequencies. The boundary conditions and the unidimensional transition conditions for a geographical barrier and for coincident discontinuities in the carrying capacity and migration rate have the standard form. In the diallelic case, reparametrization renders the entire theory of clines and of the wave of advance of favorable alleles directly applicable to plant populations. Received 30 August 1995; received in revised form 23 February 1996     

A non-local model for a swarm

 This paper describes continuum models for swarming behavior based on non-local interactions. The interactions are assumed to influence the velocity of the organisms. The model consists of integro-differential advection-diffusion equations, with convolution terms that describe long range attraction and repulsion. We find that if density dependence in the repulsion term is of a higher order than in the attraction term, then the swarm profile is realistic: i.e. the swarm has a constant interior density, with sharp edges, as observed in biological examples. This is our main result. Linear stability analysis, singular perturbation theory, and numerical experiments reveal that weak, density-independent diffusion leads to disintegration of the swarm, but only on an exponentially large time scale. When density dependence is put into the diffusion term, we find that true, locally stable traveling band solutions occur. We further explore the effects of local and non-local density dependent drift and unequal ranges of attraction and repulsion. We compare our results with results of some local models, and find that such models cannot account for cohesive, finite swarms with realistic density profiles. Received: 17 September 1997 / Revised version: 17 March 1998     

Properties of solutions for a chemotaxis system

 We investigate mathematically the system of equations proposed by Chaplain and Stuart [2], to describe the chemotactic response of endothelial cells under the angiogenesis stimulus. In particular, we characterize the steady state endothelial cell density function, and give conditions on the chemotactic parameter k and cell proliferation parameter b that ensure that migration/ proliferation either does or does not occur in steady state. The time dependent problem is also treated. Received 12 September 1995; received in revised form 6 August 1996     

On the role of body size in a tri-trophic metapopulation model

 A particular tri-trophic (resource, prey, predator) metapopulation model with dispersal of preys and predators is considered in this paper. The analysis is carried out numerically, by finding the bifurcations of the equilibria and of the limit cycles with respect to prey and predator body sizes. Two routes to chaos are identified. One is characterized by an intriguing cascade of flip and tangent bifurcations of limit cycles, while the other corresponds to the crisis of a strange attractor. The results are summarized by partitioning the space of body sizes in eight subregions, each one of which is associated to a different asymptotic behavior of the system. Emphasis is put on the possibility of having different modes of coexistence (stationary, cyclic, and chaotic) and/or extinction of the predator population. Received 1 August 1995; received in revised form 8 January