Traveling wave solutions of a reaction diffusion model for competing pioneer and climax species

Presented is a reaction-diffusion model for the interaction of pioneer and climax species. For certain parameters the system exhibits bistability and traveling wave solutions. Specifically, we show that when the climax species diffuses at a slow rate there are traveling wave solutions which correspond to extinction waves of either the pioneer or climax species. A leading order analysis is used in the one-dimensional spatial case to estimate the wave speed sign that determines which species becomes extinct. Results of these analyses are then compared to numerical simulations of wave front propagation for the model on one and two-dimensional spatial domains. A simple mechanism for harvesting is also introduced.     

On mathematical modelling of complex ecosystems: Application to marine planktonic patchiness

Mathematical modeling of complex ecosystems is very difficult due to the very large number of components in the real ecosystem. Conceptual subdivision into interacting sub-systems is necessarily subjective and is made in view of explaining a particular aspect of the reality. In this paper, the North Sea planktonic ecosystem is reduced to a rather simple mathematical model with the purpose of showing the possibility of a spontaneous spatial emergence of plankton patches by diffusive instability. Due to the dependence of diffusion coefficients on the differential diameters of phytoplankton and herbivorous zooplankton patches, respectively, the spatially homogeneous steady state is unstable for spatial perturbations with wavelengths belonging to a certain range of values. As a consequence, these perturbations amplify leading to spatial heterogeneity.     

Stabilization through spatial pattern formation in metapopulations with long-range dispersal

Many studies of metapopulation models assume that spatially extended populations occupy a network of identical habitat patches, each coupled to its nearest neighbouring patches by density-independent dispersal. Much previous work has focused on the temporal stability of spatially homogeneous equilibrium states of the metapopulation, and one of the main predictions of such models is that the stability of equilibrium states in the local patches in the absence of migration determines the stability of spatially homogeneous equilibrium states of the whole metapopulation when migration is added. Here, we present classes of examples in which deviations from the usual assumptions lead to different predictions. In particular, heterogeneity in local habitat quality in combination with long-range dispersal can induce a stable equilibrium for the metapopulation dynamics, even when within-patch processes would produce very complex behaviour in each patch in the absence of migration. Thus, when spatially homogeneous equilibria become unstable, the system can often shift to a different, spatially inhomogeneous steady state. This new global equilibrium is characterized by a standing spatial wave of population abundances. Such standing spatial waves can also be observed in metapopulations consisting of identical habitat patches, i.e. without heterogeneity in patch quality, provided that dispersal is density dependent. Spatial pattern formation after destabilization of spatially homogeneous equilibrium states is well known in reaction–diffusion systems and has been observed in various ecological models. However, these models typically require the presence of at least two species, e.g. a predator and a prey. Our results imply that stabilization through spatial pattern formation can also occur in single-species models. However, the opposite effect of destabilization can also occur: if dispersal is short range, and if there is heterogeneity in patch quality, then the metapopulation dynamics can be chaotic despite the patches having stable equilibrium dynamics when isolated. We conclude that more general metapopulation models than those commonly studied are necessary to fully understand how spatial structure can affect spatial and temporal variation in population abundance.     

Restoration Ecology: Two-Sex Dynamics and Cost Minimization

We model a spatially detailed, two-sex population dynamics, to study the cost of ecological restoration. We assume that cost is proportional to the number of individuals introduced into a large habitat. We treat dispersal as homogeneous diffusion in a one-dimensional reaction-diffusion system. The local population dynamics depends on sex ratio at birth, and allows mortality rates to differ between sexes. Furthermore, local density dependence induces a strong Allee effect, implying that the initial population must be sufficiently large to avert rapid extinction. We address three different initial spatial distributions for the introduced individuals; for each we minimize the associated cost, constrained by the requirement that the species must be restored throughout the habitat. First, we consider spatially inhomogeneous, unstable stationary solutions of the model’s equations as plausible candidates for small restoration cost. Second, we use numerical simulations to find the smallest rectangular cluster, enclosing a spatially homogeneous population density, that minimizes the cost of assured restoration. Finally, by employing simulated annealing, we minimize restoration cost among all possible initial spatial distributions of females and males. For biased sex ratios, or for a significant between-sex difference in mortality, we find that sex-specific spatial distributions minimize the cost. But as long as the sex ratio maximizes the local equilibrium density for given mortality rates, a common homogeneous distribution for both sexes that spans a critical distance yields a similarly low cost.     

A Turing model with correlated random walk

 If in the classical Turing model the diffusion process (Brownian motion) is replaced by a more general correlated random walk, then the parameters describing spatial spread are the particle speeds and the rates of change in direction. As in the Turing model, a spatially constant equilibrium can become unstable if the different species have different turning rates and different speeds. Furthermore, a Hopf bifurcation can be found if the reproduction rate of the activator is greater than its rate of change of direction, and oscillating patterns are possible. Received 24 February 1995; received in revised form 6 September 1995     

Sustainability without coexistence state in Durrett–Levin hawk–dove model

Models that explain the sustainability of an exploiter–victim ecosystem admit, generally, a coexistence state of both species in the well-mixed limit. Even if this state is unstable, the extinction-prone system may acquire stability on spatial domains where different patches oscillate incoherently around the coexistence state. New experiments, however, suggest that a spatially segregated system may be stable even in the absence of such a coexistence state. Here we revisit the hawk–dove (case 3) model of Durrett and Levin, which has been shown to support persistent population for system of interacting particles. It turns out that this model does not admit a (stable or unstable) coexistence state on a single habitat. We analyze the peculiar mechanism that leads to persistence in this case and the role of demographic stochasticity with and without self-interaction, using numerical simulations and exact solutions in the infinite diffusion limit.     

Spatial patterns in population conflicts

We consider a dynamical model for evolutionary games, and enquire how the introduction of diffusion may lead to the formation of stationary spatially inhomogeneous solutions, that is patterns. For the model equations being used it is already known that if there is an evolutionarily stable strategy (ESS), then it is stable. Equilibrium solutions which are not ESS's and which are stable with respect to spatially constant perturbations may be unstable for certain choices of the dispersal rates. We prove by a bifurcation technique that under appropriate conditions the instability leads to patterns. Computations using a curve-following technique show that the bifurcations exhibit a rich structure with loops joined by symmetry-breaking branches.     

An analysis of an ordinary differential equation model for a two-species predator-prey system with harvesting and stocking

An analysis is presented for a model of a two-species predator-prey system where each species can be harvested or stocked. Using methods from bifurcation theory the qualitative nature of the steady-state solutions is examined. The effect of harvesting and stocking rates and the prey carrying capacity is examined in detail.     

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.     

Predator-prey models with delay and prey harvesting

It is known that predator-prey systems with constant rate harvesting exhibit very rich dynamics. On the other hand, incorporating time delays into predator-prey models could induce instability and bifurcation. In this paper we are interested in studying the combined effects of the harvesting rate and the time delay on the dynamics of the generalized Gause-type predator-prey models and the Wangersky-Cunningham model. It is shown that in these models the time delay can cause a stable equilibrium to become unstable and even a switching of stabilities, while the harvesting rate has a stabilizing effect on the equilibrium if it is under the critical harvesting level. In particular, one of these models loses stability when the delay varies and then regains its stability when the harvesting rate is increased. Computer simulations are carried to explain the mathematical conclusions. Received: 1 March 2000 / Revised version: 7 September 2000 /?Published online: 21 August 2001     

Effects of convective and dispersive interactions on the stability of two species

The evolution and local stability of a system of two interacting species in a finite two-dimensional habitat is investigated by taking into account the effects of self- and cross-dispersion and convection of the species. In absence of cross-dispersion, an equilibrium state which is stable without dispersion is always stable with dispersion provided that the dispersion coefficients of the two species are equal. However, when the dispersion coefficients of the two species are different, the possibility of self-dispersive instability arises. It is also pointed out that the cross-dispersion of species may lead to stability or instability depending upon the nature and the magnitude of the cross-dispersive interactions in comparison to the self-dispersive interactions. The self-convective movement of species increases the stability of the equilibrium state and can stabilize an otherwise unstable equilibrium state. The effect of cross-convection (in absence of self-dispersion and self-convection) is to stabilize the equilibrium state in a prey-predator model with positive cross-dispersion coefficients for the prey species. Finally, it is shown that if the system is stable under homogeneous boundary conditions it remains so under non-homogeneous boundary conditions.     

Effects of quarantine in six endemic models for infectious diseases

Thresholds, equilibria, and their stability are found for SIQS and SIQR epidemiology models with three forms of the incidence. For most of these models, the endemic equilibrium is asymptotically stable, but for the SIQR model with the quarantine-adjusted incidence, the endemic equilibrium is an unstable spiral for some parameter values and periodic solutions arise by Hopf bifurcation. The Hopf bifurcation surface and stable periodic solutions are found numerically.     

The management of pesticide resistance.

We solve the two-species Volterra type non-linear reactive-diffusive growth equations in one spatial dimension which is taken to be infinite. We find periodic solutions which oscillate about the constant (space independent) equilibrium solutions and are stable with respect to the non-linear perturbations for small values of the diffusion constants.     

Biological Control Through Intraguild Predation: Case Studies in Pest Control,Invasive Species and Range Expansion

Intraguild predation (IGP), the interaction between species that eat each other and compete for shared resources, is ubiquitous in nature. We document its occurrence across a wide range of taxonomic groups and ecosystems with particular reference to non-indigenous species and agricultural pests. The consequences of IGP are complex and difficult to interpret. The purpose of this paper is to provide a modelling framework for the analysis of IGP in a spatial context. We start by considering a spatially homogeneous system and find the conditions for predator and prey to exclude each other, to coexist and for alternative stable states. Management alternatives for the control of invasive or pest species through IGP are presented for the spatially homogeneous system. We extend the model to include movement of predator and prey. In this spatial context, it is possible to switch between alternative stable steady states through local perturbations that give rise to travelling waves of extinction or control. The direction of the travelling wave depends on the details of the nonlinear intraguild interactions, but can be calculated explicitly. This spatial phenomenon suggests means by which invasions succeed or fail, and yields new methods for spatial biological control. Freshwater case studies are used to illustrate the outcomes.     

Spatial stability analysis of Eigen's quasispecies model and the less than five membered hypercycle under global population regulation

A generalization of the “constant overall organization” constraint of Eigen's quasispecies and hypercycle models, called herein “global population regulation”, is shown to lead to mathematically tractable spatial generalizations of these two models. The spatially uniform steady state of Eigen's quasispecies model is shown to be stable and globally attracting for all possible values of the mutation and replication rates. In contrast, the spatially and temporally uniform solutions to the hypercycle with fewer than five members, the only ones insensitive to stochastic perturbations, are shown to be unstable, and a lower bound to the spatial inhomogeneities is obtained. The prospect that the spatially localized hypercycle might be immune to various instabilities cited in the literature is then briefly considered. Although spatial localization makes possible a much richer dynamical repertoire than previously considered, it is also more difficult to understand how Darwinian selection of hypercycles could result in a unique genetic code.     

Nutrient flows between ecosystems can destabilize simple food chains

Dispersal of organisms has large effects on the dynamics and stability of populations and communities. However, current metacommunity theory largely ignores how the flows of limiting nutrients across ecosystems can influence communities. We studied a meta-ecosystem model where two autotroph-consumer communities are spatially coupled through the diffusion of the limiting nutrient. We analyzed regional and local stability, as well as spatial and temporal synchrony to elucidate the impacts of nutrient recycling and diffusion on trophic dynamics. We show that nutrient diffusion is capable of inducing asynchronous local destabilization of biotic compartments through a diffusion-induced spatiotemporal bifurcation. Nutrient recycling interacts with nutrient diffusion and influences the susceptibility of the meta-ecosystem to diffusion-induced instabilities. This interaction between nutrient recycling and transport is further shown to depend on ecosystem enrichment. It more generally emphasizes the importance of meta-ecosystem theory for predicting species persistence and distribution in managed ecosystems.     

Prey Selection,Vertical Migrations and the Impacts of Harvesting Upon the Population Dynamics of a Predator-Prey System

A model is developed to describe the interaction between a predator and two prey types located in different regions. Conditions for stability and persistence are analysed. The effects of harvesting the predators are investigated by making the predator mortality rate habitat dependent. Results demonstrate that for any given set of parameter values there is a value of the intrinsic preference of the predator for each prey type at which the system undergoes a Hopf bifurcation. Above this critical value the system evolves towards a stable equilibrium, whereas below it, stable limit cycles arise by Hopf bifurcations. Simulations demonstrate that the presence of demographic stochasticity may destabilise oscillatory populations, thereby causing population extinctions. An application of the model to the foraging behaviour of North Sea cod is described. It is shown that if the preferred prey is more productive, it is likely that the equilibrium will be stable, whereas if the less preferred prey is more productive, populations are likely to display cycles and in the stochastic case become extinct. As cod fishing mortality is increased, the point of bifurcation and region of parameter space for which the system is unstable decreases. An increased understanding of how cod behave may enable fish stocks to be managed more successfully, for example by indicating where marine reserves should be placed.     

Bifurcations and phase transitions in spatially extended two-member hypercycles

Mounting theoretical and experimental evidence indicates that the success of molecular replicators is strongly tied to the local nature of their interactions. Local dispersal in a given spatial domain, particularly on surfaces, might strongly enhance the growth and selection of fit molecules and their resistance to parasites. In this work the spatial dynamics of a simple hypercycle model consisting of two molecular species is analysed. In order to characterize it, both mean field models and stochastic, spatially explicit approaches are considered. The mean field approach predicts the presence of a saddle-node bifurcation separating a phase involving stable hypercycles from extinction, consistently with spatially explicit models, where an absorbing first-order phase transition is shown to exist and diffusion is explicitly introduced. The saddle-node bifurcation is shown to leave a ghost in the phase plane. A metapopulation-based model is also developed in order to account for the observed phases when both diffusion and reaction are considered. The role of information and diffusion as well as the relevance of these phases and the underlying spatial structures are discussed, and their potential implications for the evolution of early replicators are outlined.     

A new mathematical model for the homeostatic effects of sleep loss on neurobehavioral performance

The two-process model of sleep regulation makes accurate predictions of sleep timing and duration for a variety of experimental sleep deprivation and nap sleep scenarios. Upon extending its application to waking neurobehavioral performance, however, the model fails to predict the effects of chronic sleep restriction. Here we show that the two-process model belongs to a broader class of models formulated in terms of coupled non-homogeneous first-order ordinary differential equations, which have a dynamic repertoire capturing waking neurobehavioral functions across a wide range of wake/sleep schedules. We examine a specific case of this new model class, and demonstrate the existence of a bifurcation: for daily amounts of wakefulness less than a critical threshold, neurobehavioral performance is predicted to converge to an asymptotically stable state of equilibrium; whereas for daily wakefulness extended beyond the critical threshold, neurobehavioral performance is predicted to diverge from an unstable state of equilibrium. Comparison of model simulations to laboratory observations of lapses of attention on a psychomotor vigilance test (PVT), in experiments on the effects of chronic sleep restriction and acute total sleep deprivation, suggests that this bifurcation is an essential feature of performance impairment due to sleep loss. We present three new predictions that may be experimentally verified to validate the model. These predictions, if confirmed, challenge conventional notions about the effects of sleep and sleep loss on neurobehavioral performance. The new model class implicates a biological system analogous to two connected compartments containing interacting compounds with time-varying concentrations as being a key mechanism for the regulation of psychomotor vigilance as a function of sleep loss. We suggest that the adenosinergic neuromodulator/receptor system may provide the underlying neurobiology.