Nonlinear Functional Responses Research Articles

Structure of positive solutions for a reaction-diffusion model with additional food and protection zone

The additional food and protection zone provided to predator and prey respectively are introduced into a reaction-diffusion predation model with nonlinear functional response in this paper. We mainly concentrate on the combined effects of additional food and protection zone on the existence and uniqueness of solution for the model. It is shown that, compared with the models with classical Beddington-DeAngelis functional response, the integration of additional food and protection zone into Beddington-DeAngelis functional response causes the model to produce new critical values for the existence of positive solutions. Moreover, we derive that the model permits a unique positive solution when the strength of mutual interference among predators or the quantity of additional food is large, where, the effect of quantity of additional food on the positive solution is a new finding. The results shed light on that the combination of additional food and protection zone is an effective and environmentally-friendly strategy in preserving biodiversity.

Steady-State Bifurcation and Spatial Patterns of a Chemical Reaction System

This paper studies the Lengyel–Epstein chemical reaction system with nonlinear functional response and no-flux boundary conditions. We first investigate the existence of steady-state bifurcation solutions of the system. Then, the stability of bifurcation solutions is analyzed. Meanwhile, some spatial patterns induced by steady-state bifurcation are simulated numerically and depicted graphically. It is well known that the classical bifurcation theory in nonlinear dynamical systems is based on simple eigenvalues. However, this is not always the case. Sometimes, the kernel of an objective operator is of two or more dimensions. For such cases, there is no existing theory to deal with them. In this paper, by using the space decomposition technique and implicit function theorem, we analyze the bifurcation phenomenon of the system in the case of a two-dimensional kernel of some certain objective operator. The results show that the chemical reaction between activator iodide and inhibitor chlorite can proceed stably under certain conditions.

Viral Infection Model with Diffusion and Distributed Delay: Finite-Dimensional Global Attractor.

We study a virus dynamics model with reaction-diffusion, logistic growth terms and a general non-linear infection rate functional response. The model has a distributed delay, including the case of state-selective delay. We construct a dynamical system in a Hilbert space and prove the existence of a finite-dimensional global attractor.

On the small time asymptotics of stochastic predator–prey model with nonlinear functional response

On the small time asymptotics of stochastic predator–prey model with nonlinear functional response

Stability and Hopf bifurcation of a diffusive plankton model with time-delay and mixed nonlinear functional responses

In this paper, we deal with a plankton reaction–diffusion model with time-delay and two different functional responses. Firstly, we consider the global stability of boundary equilibrium point. Secondly, we investigate the existence, uniqueness and stability of internal equilibrium point without time-delay. Then, we analyze the existence of Hopf bifurcation emitting from internal equilibrium point and give some characteristics on Hopf branch in detail. A new finding is presented, specifically, we find that there exist two critical values which have important effects on the occurrence of Hopf bifurcation. Finally, a few numerical examples are presented to check and illustrate the theoretical analysis, some simulation graphs, including the spatiotemporal graphs, trajectory graphs and phase portraits are depicted graphically.

Incorporating nonlinearity with generalized functional responses to simulate multiple predator effects.

Predicting the combined effects of predators on shared prey has long been a focus of community ecology, yet quantitative predictions often fail. Failure to account for nonlinearity is one reason for this. Moreover, prey depletion in multiple predator effects (MPE) studies generates biased predictions in applications of common experimental and quantitative frameworks. Here, we explore additional sources of bias stemming from nonlinearities in prey predation risk. We show that in order to avoid bias, predictions about the combined effects of independent predators must account for nonlinear size-dependent risk for prey as well as changes in prey risk driven by nonlinear predator functional responses and depletion. Historical failure to account for biases introduced by well-known nonlinear processes that affect predation risk suggest that we may need to reevaluate the general conclusions that have been drawn about the ubiquity of emergent MPEs over the past three decades.

Stability Analysis, Hopf Bifurcation and Drug Therapy Control of an HIV Viral Infection Model with Logistic Growth Rate and Cell-to-Cell and Cell-Free Transmissions

It is well known that dynamical systems are very useful tools to study viral diseases such as HIV, HBV, HCV, Ebola and Influenza. This paper focuses on a mathematical model of the cell-to-cell and the cell-free spread of HIV with both linear and nonlinear functional responses and logistic target cell growth. The reproduction number of each mode of transmission has been calculated and their sum has been considered as the basic reproduction number. Based on the values of the reproduction number, the local and global stabilities of the rest points are investigated. Choosing a suitable bifurcation parameter, some conditions for the occurrence of Hopf bifurcation are also obtained. Moreover, numerical simulations are presented to support the analytical results. Finally, to study the effect of the drug on the disease process, some control conditions are determined. Since two modes of transmission and both linear and nonlinear functional responses have been included in this manuscript, our obtained results are a generalization of those in the literature. Moreover, the results are obtained with weaker assumptions in comparison with the previous ones.

Bridging Theories for Ecosystem Stability Through Structural Sensitivity Analysis of Ecological Models in Equilibrium

Ecologists are challenged by the need to bridge and synthesize different approaches and theories to obtain a coherent understanding of ecosystems in a changing world. Both food web theory and regime shift theory shine light on mechanisms that confer stability to ecosystems, but from different angles. Empirical food web models are developed to analyze how equilibria in real multi-trophic ecosystems are shaped by species interactions, and often include linear functional response terms for simple estimation of interaction strengths from observations. Models of regime shifts focus on qualitative changes of equilibrium points in a slowly changing environment, and typically include non-linear functional response terms. Currently, it is unclear how the stability of an empirical food web model, expressed as the rate of system recovery after a small perturbation, relates to the vulnerability of the ecosystem to collapse. Here, we conduct structural sensitivity analyses of classical consumer-resource models in equilibrium along an environmental gradient. Specifically, we change non-proportional interaction terms into proportional ones, while maintaining the equilibrium biomass densities and material flux rates, to analyze how alternative model formulations shape the stability properties of the equilibria. The results reveal no consistent relationship between the stability of the original models and the proportionalized versions, even though they describe the same biomass values and material flows. We use these findings to critically discuss whether stability analysis of observed equilibria by empirical food web models can provide insight into regime shift dynamics, and highlight the challenge of bridging alternative modelling approaches in ecology and beyond.

Existence of positive periodic solutions of a delayed periodic Microcystins degradation model with nonlinear functional responses

This paper proposes a delayed periodic Microcystins degradation model with nonlinear functional responses. By using the continuation theorem in the coincidence degree, we obtain two classes of explicit sufficient conditions for the existence of positive periodic solutions of the model.

Accounting for a nonlinear functional response when estimating prey dynamics using predator diet data

Abstract Forage fish species are key in the transfer of energy from lower to upper trophic levels in marine ecosystems. Therefore, understanding their population dynamics, including population levels, is crucial for understanding productivity and the regulation of marine food webs. However, many forage fishes are poorly sampled by bottom trawl surveys, leading to poor estimates of their abundance. These estimates can be improved by using predator stomach contents as an additional sampling strategy; however, nonlinear relationships between prey abundance and predator consumption (i.e. the functional response) may bias stomach data as well. Using predator stomach contents and bottom trawl survey data, this study aimed to minimize this bias by developing a model to estimate prey dynamics and account for the predator functional response. This model was tested using a series of simulations and applied to a case study of northern sand lance Ammodytes dubius on the Grand Bank, Newfoundland, Canada. The simulations revealed that when predators consumed prey following a nonlinear functional response, our model outperformed a classical model (the model adopted by most studies) that assumed a linear functional response. In the case study, we estimated the relative abundance of sand lance from 1995 to 2018, which exhibited oscillatory dynamics with a period of approximately 7 years. Our results demonstrate that our model is capable of more accurately estimating the abundance of data‐limited prey populations, which contributes to a better understanding of food web dynamics.

Non-additive biotic interactions improve predictions of tropical tree growth and impact community size structure.

Growth in individual size or biomass is a key demographic component in population models, with wide-ranging applications from quantifying species performance across abiotic or biotic conditions to assessing landscape-level dynamics under global change. In forest ecology, the responses of tree growth to biotic interactions are widely held to be crucial for understanding forest diversity, function, and structure. To date, most studies on plant-plant interactions only examine the additive competitive or facilitative interactions between species pairs; however, there is increasing evidence of non-additive, higher-order interactions (HOIs) impacting species demographic rates. When HOIs are present, the dynamics of a multispecies community cannot be fully understood or accurately predicted solely from pairwise outcomes because of how additional species "interfere" with the direct, pairwise interactions. Such HOIs should be particularly prevalent when species show non-linear functional responses to resource availability and resource-acquisition traits themselves are density dependent. With this in mind, we used data from a tropical secondary forest-a system that fulfills both of these conditions-to build an ontogenetic diameter growth model for individuals across 10 woody-plant species. We allowed both direct and indirect interactions within communities to influence the species-specific growth parameters in a generalized Lotka-Volterra model. Specifically, indirect interactions entered the model as higher-order quadratic terms, i.e., non-additive effects of conspecific and heterospecific neighbor size on the focal individual's growth. For the whole community and for four out of 10 focal species, the model that included HOIs had more statistical support than the model that included only direct interactions, despite the former containing a far greater number of parameters. HOIs had comparable effect sizes to direct interactions, and tended to further reduce the diameter growth rates of most species beyond what direct interactions had already reduced. In a simulation of successional stand dynamics, the inclusion of HOIs led to rank swaps in species' diameter hierarchies, even when community-level size distributions remained qualitatively similar. Our study highlights the implications, and discusses possible mechanisms, of non-additive density dependence in highly diverse and light-competitive tropical forests.

Holling meets habitat selection: functional response of large herbivores revisited

BackgroundHolling (Can Entomol 91(5):293–320, 1959) was the first to describe a functional response between a predator’s consumption-rate and the density of its prey. The same concept can be applied to the habitat selection of herbivores, specifically, the change in relative habitat use with the change in habitat availability. Functional responses in habitat selection at a home-range scale have been reported for several large herbivores. However, a link to Holling’s original functional response types has never been drawn, although it could replace the current phenomenological view with a more mechanistically based understanding of functional responses.MethodsIn this study, discrete choice models were implemented as mixed-effects baseline-category logit models to analyze the variation in habitat selection of a large herbivore at seasonal and diurnal scales. Thus, changes in the use of land cover types with respect to their availability were investigated by monitoring 11 land cover types commonly used by roe deer (Capreolus capreolus) in the Bavarian Forest National Park, Germany. Functional response curves were then fitted using Holling’s formulas.ResultsStrong evidence of non-linear functional responses was obtained for almost all of the examined land cover types. The shape of the functional response curves varied depending on the season, the time of day, and in some cases between sexes. These responses could be referenced to Holling’s types, with a predominance of type II.ConclusionsOur results indicate that Holling’s types can be applied to describe general patterns of the habitat selection behavior of herbivores. Functional responses in habitat selection may occur in situations requiring a trade-off in the selection of land cover types offering different resources, such as due to the temporally varying physiological needs of herbivores. Moreover, two associated parameters defining the curves (prey density and predation rate) can aid in the identification of temporal variations and in determinations of the strength of the cost-benefit ratio for a specific land cover type. Application of our novel approach, using Holling’s equations to describe functional responses in the habitat selection of herbivores, will allow the assignment of general land cover attraction values, independent of availability, thus facilitating the identification of suitable habitats.

Stochasticity in host-parasitoid models informs mechanisms regulating population dynamics

Population dynamics of host-parasitoid interactions have been traditionally studied using a discrete-time formalism starting from the classical work of Nicholson and Bailey. It is well known that differences in parasitism risk among individual hosts can stabilize the otherwise unstable equilibrium of the Nicholson-Bailey model. Here, we consider a stochastic formulation of these discrete-time models, where the host reproduction is a random variable that varies from year to year and drives fluctuations in population densities. Interestingly, our analysis reveals that there exists an optimal level of heterogeneity in parasitism risk that minimizes the extent of fluctuations in the host population density. Intuitively, low variation in parasitism risk drives large fluctuations in the host population density as the system is on the edge of stability. In contrast, high variation in parasitism risk makes the host equilibrium sensitive to the host reproduction rate, also leading to large fluctuations in the population density. Further results show that the correlation between the adult host and parasitoid densities is high for the same year, and gradually decays to zero as one considers cross-species correlations across different years. We next consider an alternative mechanism of stabilizing host-parasitoid population dynamics based on a Type III functional response, where the parasitoid attack rate accelerates with increasing host density. Intriguingly, this nonlinear functional response makes qualitatively different correlation signatures than those seen with heterogeneity in parasitism risk. In particular, a Type III functional response leads to uncorrelated adult and parasitoid densities in the same year, but high cross-species correlation across successive years. In summary, these results argue that the cross-correlation function between population densities contains signatures for uncovering mechanisms that stabilize consumer-resource population dynamics.

Dynamic analysis of a predator-prey system with nonlinear prey harvesting and square root functional response

Abstract In this work, the dynamics of a predator-prey system considering square root type functional response for prey herd behaviour and nonlinear prey harvesting has been analyzed. The conditions under which all equilibria exist as well as the stability of every equilibrium point of the system have been investigated. The proposed model conditionally posses two types of bifurcations, Hopf bifurcation, and saddle-node bifurcation. The saddle-node bifurcation has been analyzed, where the bifurcation parameter is harvesting rate. The existence of a maximum sustainable yield to ensure both populations coexist has been discussed. The results give a clear idea that, if the harvesting rate is chosen at a proper value lesser than the maximum sustainable yield then both populations will coexist and the ecological balance will be maintained. The calculation of the first Lyapunov number provides the Hopf bifurcation direction. To verify our analytical results, several numerical simulations have been carried out.

A mathematical analysis of Hopf-bifurcation in a prey-predator model with nonlinear functional response

In this paper, our aim is mathematical analysis and numerical simulation of a prey-predator model to describe the effect of predation between prey and predator with nonlinear functional response. First, we develop results concerning the boundedness, the existence and uniqueness of the solution. Furthermore, the Lyapunov principle and the Routh–Hurwitz criterion are applied to study respectively the local and global stability results. We also establish the Hopf-bifurcation to show the existence of a branch of nontrivial periodic solutions. Finally, numerical simulations have been accomplished to validate our analytical findings.

Quantifying the effects of intraspecific variation on predator feeding rates through nonlinear averaging

Abstract Theory suggests that intraspecific trait variation will alter species interaction strengths through nonlinear averaging when interaction strengths are nonlinear functions of individuals' traits. This effect is expected to be widespread, yet what factors mediate its magnitude in nature and hence its potential effects on ecosystems and communities are unclear. We sought to quantify how nonlinear predator functional responses, variation in prey densities and counteracting variation in attack rates and handling times among predator individuals of similar body size alter their population‐level feeding rates through nonlinear averaging in a natural system, and to determine the processes influencing the net magnitude of this effect. We used a field caging experiment in the rocky intertidal of Oregon, USA to quantify attack‐rate variation and feeding rates of the whelk Nucella ostrina on its barnacle and mussel prey. We also used empirically parameterized simulations to examine the effects of handling‐time variation among individuals on population‐level feeding rates. Within cages, individual attack‐rate variation reduced population‐level whelk feeding rates. However, the magnitude of this reduction differed among prey species and cages depending on cage‐specific magnitudes of attack‐rate variation and functional‐response nonlinearity. The inferred effects of handling‐time variation among individuals were of smaller magnitude than those of attack‐rate variation, yet counteracted them to cause a net weakening of the effect of individual attack‐rate variation on population‐level feeding rates. Across cages, attack‐rate and prey‐density variation had non‐additive effects that produced greater feeding‐rate reductions at the experiment scale relative to the cage scale. Our results indicate that the effects of trait variation via nonlinear averaging depend critically on the features of systems that determine the magnitudes of nonlinearities and trait variation. Because of counteracting trait variation, nonlinear‐averaging effects may be quite complex, involving both the variances and covariances of all traits and environmental variables influencing the ecological process of interest. A free Plain Language Summary can be found within the Supporting Information of this article.

Analysis and simulation on dynamics of a partial differential system with nonlinear functional responses

We introduce a reaction–diffusion system with modified nonlinear functional responses. We first discuss the large-time behavior of positive solutions for the system. And then, for the corresponding steady-state system, we are concerned with the priori estimate, the existence of the nonconstant positive solutions as well as the bifurcations emitting from the positive constant equilibrium solution. Finally, we present some numerical examples to test the theoretical and computational analysis results. Meanwhile, we depict the trajectory graphs and spatiotemporal patterns to simulate the dynamics for the system. The numerical computations and simulated graphs imply that the available food resource for consumer is very likely not single.

The Existence and Uniqueness of the Solution of a Diusive Predator-Prey Model with Omnivory and General Nonlinear Functional Response

In this work, we consider a three species modified Lesie-Gower food web model with general nonlinear functional response and omnivory which is defined as feeding on more than one trophic level. The carrying capacity of the model is proportional to the population size of the biotic resource plus a const. The main objective of this paper is to investigate the existence and uniqueness of the solution of this model. It is shown that the omnivory has important influence on the existence and uniqueness of the solution of the model.

Analysis on Steady States of a Competition System with Nonlinear Diffusion Terms

Competition is a fundamental force shaping population size and structure as a result of limited availability of resources. In biomathematics, the biological models with competitive interactions exist widely. Furthermore, the nonlinear-diffusion (including self- and cross-diffusions) terms are incorporated to the biological models to better simulate the actual movement of species. Therefore, better compatibility with reality can be achieved by introducing nonlinear-diffusion into biological models with competitive interactions. As a result, a competition system with nonlinear-diffusion and nonlinear functional response is proposed and analyzed in this paper. We first briefly discuss the stability of trivial and semi-trivial solutions by spectrum analysis. Then the boundedness and the non-existence of steady states are studied. Based on the boundedness of the solutions, the existence of the steady states is also investigated by the fixed point index theory in a positive cone. The result shows that the two species can coexist when their diffusion and inter-specific competition pressures are controlled in a certain range.

Modelling and analysis of a stochastic nonautonomous predator-prey model with impulsive effects and nonlinear functional response.

In this paper, a new stochastic predator-prey model with impulsive perturbation and Crowley-Martin functional response is proposed. The dynamical properties of the model are systematically investigated. The existence and stochastically ultimate boundedness of a global positive solution are derived using the theory of impulsive stochastic differential equations. Some sufficient criteria are obtained to guarantee the extinction and a series of persistence in the mean of the system. Moreover, we provide conditions for the stochastic permanence and global attractivity of the model. Numerical simulations are performed to support our qualitative results.