Modelling the effect of temperature on the range expansion of species by reaction-diffusion equations

The spatial dynamics of range expansion is studied in dependence of temperature. The main elements population dynamics, competition and dispersal are combined in a coherent approach based on a system of coupled partial differential equations of the reaction-diffusion type. The nonlinear reaction terms comprise population dynamic models with temperature dependent reproduction rates subject to an Allee effect and mutual competition. The effect of temperature on travelling wave solutions is investigated for a one dimensional model version. One main result is the importance of the Allee effect for the crossing of regions with unsuitable habitats. The nonlinearities of the interaction terms give rise to a richness of spatio-temporal dynamic patterns. In two dimensions, the resulting non-linear initial boundary value problems are solved over geometries of heterogeneous landscapes. Geo referenced model parameters such as mean temperature and elevation are imported into the finite element tool COMSOL Multiphysics from a geographical information system. The model is applied to the range expansion of species at the scale of middle Europe.     

Early development and quorum sensing in bacterial biofilms

 We develop mathematical models to examine the formation, growth and quorum sensing activity of bacterial biofilms. The growth aspects of the model are based on the assumption of a continuum of bacterial cells whose growth generates movement, within the developing biofilm, described by a velocity field. A model proposed in Ward et al. (2001) to describe quorum sensing, a process by which bacteria monitor their own population density by the use of quorum sensing molecules (QSMs), is coupled with the growth model. The resulting system of nonlinear partial differential equations is solved numerically, revealing results which are qualitatively consistent with experimental ones. Analytical solutions derived by assuming uniform initial conditions demonstrate that, for large time, a biofilm grows algebraically with time; criteria for linear growth of the biofilm biomass, consistent with experimental data, are established. The analysis reveals, for a biologically realistic limit, the existence of a bifurcation between non-active and active quorum sensing in the biofilm. The model also predicts that travelling waves of quorum sensing behaviour can occur within a certain time frame; while the travelling wave analysis reveals a range of possible travelling wave speeds, numerical solutions suggest that the minimum wave speed, determined by linearisation, is realised for a wide class of initial conditions. Received: 10 February 2002 / Revised version: 29 October 2002 / Published online: 19 March 2003 Key words or phrases: Bacterial biofilm – Quorum sensing – Mathematical modelling – Numerical solution – Asymptotic analysis – Travelling wave analysis     

Spreading speed, traveling waves, and minimal domain size in impulsive reaction-diffusion models

How growth, mortality, and dispersal in a species affect the species' spread and persistence constitutes a central problem in spatial ecology. We propose impulsive reaction-diffusion equation models for species with distinct reproductive and dispersal stages. These models can describe a seasonal birth pulse plus nonlinear mortality and dispersal throughout the year. Alternatively, they can describe seasonal harvesting, plus nonlinear birth and mortality as well as dispersal throughout the year. The population dynamics in the seasonal pulse is described by a discrete map that gives the density of the population at the end of a pulse as a possibly nonmonotone function of the density of the population at the beginning of the pulse. The dynamics in the dispersal stage is governed by a nonlinear reaction-diffusion equation in a bounded or unbounded domain. We develop a spatially explicit theoretical framework that links species vital rates (mortality or fecundity) and dispersal characteristics with species' spreading speeds, traveling wave speeds, as well as minimal domain size for species persistence. We provide an explicit formula for the spreading speed in terms of model parameters, and show that the spreading speed can be characterized as the slowest speed of a class of traveling wave solutions. We also give an explicit formula for the minimal domain size using model parameters. Our results show how the diffusion coefficient, and the combination of discrete- and continuous-time growth and mortality determine the spread and persistence dynamics of the population in a wide variety of ecological scenarios. Numerical simulations are presented to demonstrate the theoretical results.     

Dispersal and competition models for plants

New models for seed dispersal and competition between plant species are formulated and analyzed. The models are integrodifference equations, discrete in time and continuous in space, and have applications to annual and perennial species. The spread or invasion of a single plant species into a geographic region is investigated by studying the travelling wave solutions of these equations. Travelling wave solutions are shown to exist in the single-species models and are compared numerically. The asymptotic wave speed is calculated for various parameter values. The single-species integrodifference equations are extended to a model for two competing annual plants. Competition in the two-species model is based on a difference equation model developed by Pakes and Maller [26]. The two-species model with competition and dispersal yields a system of integrodifference equations. The effects of competition on the travelling wave solutions of invading plant species is investigated numerically.     

Integrodifference equations in the presence of climate change: persistence criterion,travelling waves and inside dynamics

To understand the effects that the climate change has on the evolution of species as well as the genetic consequences, we analyze an integrodifference equation (IDE) models for a reproducing and dispersing population in a spatio-temporal heterogeneous environment described by a shifting climate envelope. Our analysis on the IDE focuses on the persistence criterion, travelling wave solutions, and the inside dynamics. First, the persistence criterion, characterizing the global dynamics of the IDE, is established in terms of the basic reproduction number. In the case of persistence, a unique travelling wave is found to govern the global dynamics. The effects of the size and the shifting speed of the climate envelope on the basic reproduction number, and hence, on the persistence criterion, are also investigated. In particular, the critical domain size and the critical shifting speed are found in certain cases. Numerical simulations are performed to complement the theoretical results. In the case of persistence, we separate the travelling wave and general solutions into spatially distinct neutral fractions to study the inside dynamics. It is shown that each neutral genetic fraction rearranges itself spatially so as to asymptotically achieve the profile of the travelling wave. To measure the genetic diversity of the population density we calculate the Shannon diversity index and related indices, and use these to illustrate how diversity changes with underlying parameters.

     

A mathematical model for weed dispersal and control

Mathematical models for weed dispersal and control are developed, analyzed and numerically simulated. A model incorporating periodic control, e.g. herbicide application, is derived for a plant population in a spatially homogeneous setting. The model is extended to a spatially heterogeneous population where plant dispersal is incorporated. The dispersal and control model involves integrodifference equations, discrete in time and continuous in space. The models are analyzed to determine values of the control parameter that prevent weed spread. The effects of the control on travelling wave solutions are investigated numerically.     

Breaking of a wake wave excited by a narrow laser pulse in a low-density plasma

The spatial structure of a wake wave excited in a low-density plasma by a laser pulse with a small focal spot radius is studied both analytically and numerically. Numerical study shows that, in a small-amplitude laser field, a wake wave breaks after the formation of an off-axis density maximum, which grows in height away from the pulse to become infinitely high after several periods. Analytical and numerical calculations show that the singularity in the density arises from the intersection of the trajectories of neighboring particles. Numerical simulations demonstrate that, as the laser field amplitude increases, the breaking point of the wake wave rapidly approaches the pulse trailing edge. For weakly nonlinear conditions, an analytic dependence of the coordinate of the breaking point on the amplitude and transverse size of the laser pulse is obtained.     

On the nonlinear theory of collective Cherenkov interaction of a high-density relativistic beam with a plasma

The nonlinear dynamics of the instability of a straight high-density relativistic electron beam under the conditions of the stimulated Cherenkov effect in a plasma waveguide is studied both analytically and numerically. It is shown that, for a beam of sufficiently high density such that the stabilizing factors are nonlinear frequency shifts and for a plasma described in a linear approximation, the basic equations have soliton-like solutions and the electron beam after saturation of the instability relaxes to its initial, weakly perturbed state, provided that only one harmonic of the plasma and the beam density is taken into account. The analytical solutions obtained here for this case correlate well with the numerical ones. A more general model that accounts for the generation of higher harmonics of the plasma and the beam density does not yield soliton-like solutions for the time evolution of the amplitudes of the plasma and beam waves. In such a model, the instability will be collective again: it can be described analytically (at least, up to the time at which it saturates) by using equations with cubic nonlinearities and the method of expansion of the electron trajectories and momenta.     

Actin filament branching and protrusion velocity in a simple 1D model of a motile cell

We formulate and analyse a 1D model for the spatial distribution of actin density at the leading edge of a motile cell. The model incorporates nucleation, capping, growth and decay of actin filaments, as well as retrograde flow of the actin meshwork and known parameter values based on the literature. Using a simplified geometry, and reasonable assumptions about the biochemical processes, we derive PDEs for the density of actin filaments and their tips. Analytic travelling wave solutions are used to predict how the speed of the cell depends on rates of nucleation, capping, polymerization and membrane resistance. Analysis and simulations agree with experimental profiles for measured actin distributions. Extended versions of the model are studied numerically. We find that our model produces stable travelling wave solutions with reasonable cell speeds. Increasing the rate of nucleation of filaments (by the actin related protein Arp2/3) or the rate of actin polymerization leads to faster cell speed, whereas increasing the rate of capping or the membrane resistance reduces cell speed. We consider several variants of nucleation (spontaneous, tip, and side branching) and find best agreement with experimentally measured spatial profiles of filament and tip density in the side branching case.     

Longitudinal displacements of base pairs in DNA and effects on the dynamics of nonlinear excitations

A model of the DNA is proposed and studied analytically and numerically. The model is an extension of a well known model and describes the double helix as two chains of pendula (each pendulum representing a base). Each base (or pendulum) can rotate and translate along the helix axis. In the continuum limit the system is described by the perturbed Sine–Gordon equation describing the twist of the bases and by a nonlinear partial differential equation (PDE) describing the longitudinal displacements of the bases. This coupled system of PDEs was studied analytically using different approaches and the corresponding results were tested through numerical simulations. It was found that if the coupling parameters satisfy a well defined relationship, then there exist bounded travelling wave solutions.     

Travelling waves of a delayed SIR epidemic model with nonlinear incidence rate and spatial diffusion

This paper is concerned with the existence of travelling waves to a SIR epidemic model with nonlinear incidence rate, spatial diffusion and time delay. By analyzing the corresponding characteristic equations, the local stability of a disease-free steady state and an endemic steady state to this system under homogeneous Neumann boundary conditions is discussed. By using the cross iteration method and the Schauder's fixed point theorem, we reduce the existence of travelling waves to the existence of a pair of upper-lower solutions. By constructing a pair of upper-lower solutions, we derive the existence of a travelling wave connecting the disease-free steady state and the endemic steady state. Numerical simulations are carried out to illustrate the main results.     

Modelling the spatio-temporal dynamics of multi-species host-parasitoid interactions: heterogeneous patterns and ecological implications

A mathematical model of the spatio-temporal dynamics of a two host, two parasitoid system is presented. There is a coupling of the four species through parasitism of both hosts by one of the parasitoids. The model comprises a system of four reaction-diffusion equations. The underlying system of ordinary differential equations, modelling the host-parasitoid population dynamics, has a unique positive steady state and is shown to be capable of undergoing Hopf bifurcations, leading to limit cycle kinetics which give rise to oscillatory temporal dynamics. The stability of the positive steady state has a fundamental impact on the spatio-temporal dynamics: stable travelling waves of parasitoid invasion exhibit increasingly irregular periodic travelling wave behaviour when key parameter values are increased beyond their Hopf bifurcation point. These irregular periodic travelling waves give rise to heterogeneous spatio-temporal patterns of host and parasitoid abundance. The generation of heterogeneous patterns has ecological implications and the concepts of temporary host refuge and niche formation are considered.     

Computational modeling of stabilizing the instability of a relativistic electron beam in a dense plasma

The nonlinear dynamics of the instability developed upon the interaction between a relativistic electron beam and a dense plasma as a function of the beam density is numerically modeled. The appropriate solutions are obtained and analyzed.     

Nonlinear ion modes in a strongly coupled plasma in the presence of nonthermal ion fluids and polarization force

The nonlinear propagation of ion-acoustic (IA) waves in a strongly coupled plasma system containing Maxwellian electrons and nonthermal ions has been theoretically and numerically investigated. The well-known reductive perturbation technique is used to derive both the Burgers and Korteweg?de Vries (KdV) equations. Their shock and solitary wave solutions have also been numerically analyzed in understanding localized electrostatic disturbances. It has been observed that the basic features (viz. polarity, amplitude, width, etc.) of IA waves are significantly modified by the effect of polarization force and other plasma parameters (e.g., the electron-to-ion number density ratio and ion-to-electron temperature ratio). This is a unique finding among all theoretical investigations made before, whose probable implications are discussed in this investigation. The implications of the results obtained from this investigation may be useful in understanding the wave propagation in both space and laboratory plasmas.     

Separation of arterial pressure into a nonlinear superposition of solitary waves and a windkessel flow

A simplified model of arterial blood pressure intended for use in model-based signal processing applications is presented. The main idea is to decompose the pressure into two components: a travelling wave which describes the fast propagation phenomena predominating during the systolic phase and a windkessel flow that represents the slow phenomena during the diastolic phase. Instead of decomposing the blood pressure pulse into a linear superposition of forward and backward harmonic waves, as in the linear wave theory, a nonlinear superposition of travelling waves matched to a reduced physical model of the pressure, is proposed. Very satisfactory experimental results are obtained by using forward waves, the N-soliton solutions of a Korteweg–de Vries equation in conjunction with a two-element windkessel model. The parameter identifiability in the practically important 3-soliton case is also studied. The proposed approach is briefly compared with the linear one and its possible clinical relevance is discussed.     

A numerical study of the Belousov-Zhabotinskii reaction using Galerkin finite element methods

The Belousov-Zhabotinskii reaction has been modelled by Field and Noyes [5] as a pair of nonlinear parabolic equations. Previous studies of these, both theoretical and numerical, have assumed wave solutions travelling with constant velocity leading to a simplification of the mathematical model in the form of a system of ordinary differential equations. In the present study a finite element Galerkin method is used directly on the original parabolic system for a range of parameter values.     

Are time delays always destabilizing? Revisiting the role of time delays and the Allee effect

One of the main challenges in ecology is to determine the cause of population fluctuations. Both theoretical and empirical studies suggest that delayed density dependence instigates cyclic behavior in many populations; however, underlying mechanisms through which this occurs are often difficult to determine and may vary within species. In this paper, we consider single species population dynamics affected by the Allee effect coupled with discrete time delay. We use two different mathematical formulations of the Allee effect and analyze (both analytically and numerically) the role of time delay in different feedback mechanisms such as competition and cooperation. The bifurcation value of the delay (that results in the Hopf bifurcation) as a function of the strength of the Allee effect is obtained analytically. Interestingly, depending on the chosen delayed mechanism, even a large time delay may not necessarily lead to instability. We also show that, in case the time delay affects positive feedback (such as cooperation), the population dynamics can lead to self-organized formation of intermediate quasi-stationary states. Finally, we discuss ecological implications of our findings.     

Travelling Waves in Hyperbolic Chemotaxis Equations

Mathematical models of bacterial populations are often written as systems of partial differential equations for the densities of bacteria and concentrations of extracellular (signal) chemicals. This approach has been employed since the seminal work of Keller and Segel in the 1970s (Keller and Segel, J. Theor. Biol. 30:235–248, 1971). The system has been shown to permit travelling wave solutions which correspond to travelling band formation in bacterial colonies, yet only under specific criteria, such as a singularity in the chemotactic sensitivity function as the signal approaches zero. Such a singularity generates infinite macroscopic velocities which are biologically unrealistic. In this paper, we formulate a model that takes into consideration relevant details of the intracellular processes while avoiding the singularity in the chemotactic sensitivity. We prove the global existence of solutions and then show the existence of travelling wave solutions both numerically and analytically.     

Understanding the interplay between density dependent birth function and maturation time delay using a reaction-diffusion population model

The present work employs a nonlocal delay reaction-diffusion model to study the impacts of the density dependent birth function, maturation time delay and population dispersal on single species dynamics (i.e., extinction, survival, extinction-survival). It is shown that the maturation time and the birth function are two major factors determining the fate of single species. Whereas the dispersal acts as a subsidiary factor that only affects the spatial patterns of population densities. When the birth function has a compensating density dependence, maturation time delay cannot destabilize the population survival at the positive equilibrium. Nevertheless, when the birth function has an over-compensating density dependence, the population densities of single species fluctuate in the spatial domain due to the increased maturation time delay. With the Allee effect and over-compensating density dependence, the increases in the maturation time may cause extinction of the single species in the entire spatial domain. The numerical simulations suggest that the solutions of the general model may temporarily remain nearby a stationary wave pulse or a stationary wavefront of the reduced model. The former indicates the survival of single species in a narrow region of the spatial domain. Whereas the latter represents the survival in the entire left-half or right-half of the spatial domain.