On the impulsive controllability and bifurcation of a predator-pest model of IPM

From a practical point of view, the most efficient strategy for pest control is to combine an array of techniques to control the wide variety of potential pests that may threaten crops in an approach known as integrated pest management (IPM). In this paper, we propose a predator-prey (pest) model of IPM in which pests are impulsively controlled by means of spraying pesticides (the chemical control) and releasing natural predators (the biological control). It is assumed that the biological and chemical control are used with the same periodicity, but not simultaneously. The functional response of the predator is allowed to be predator-dependent, in the form of a Beddington-DeAngelis functional response, rather than to have a perhaps more classical prey-only dependence. The local and global stability of the pest-eradication periodic solution, as well as the permanence of the system, are obtained under integral conditions which are shown to have biological significance. In a certain limiting case, it is shown that a nontrivial periodic solution emerges via a supercritical bifurcation. Finally, our findings are confirmed by means of numerical simulations.     

一类具有两次脉冲和非线性传染率的害虫管理SI模型的灭绝性与持久性

讨论了一类在两个不同固定时刻分别释放染病害虫和喷洒农药且具有HollingⅡ类传染率的SI模型.通过脉冲微分方程的Floquet理论和小幅扰动技巧,证明了当释放的染病害虫数量超过某个临界值时,系统存在一个渐进稳定的易感害虫根除周期解,否则系统是持续生存的.通过数值模拟,验证了所得结论的正确性及系统动力学行为的复杂性,分析说明了所提出的脉冲控制策略的有效性.     

Modelling and analysis of impulsive releases of sterile mosquitoes

To study the impact of releasing sterile mosquitoes on mosquito-borne disease transmissions, we propose two mathematical models with impulsive releases of sterile mosquitoes. We consider periodic impulsive releases in the first model and obtain the existence, uniqueness, and globally stability of a wild-mosquito-eradication periodic solution. We also establish thresholds for the control of the wild mosquito population by selecting the release rate and the release period. In the second model, the impulsive releases are determined by the closely monitored wild mosquito density, or the state feedback. We prove the existence of an order one periodic solution and find a relatively small attraction region, which ensures the wild mosquito population is under control. We provide numerical analysis which shows that a smaller release rate and more frequent releases are more efficient in controlling the wild mosquito population for the periodic releases, but an early release of sterile mosquitoes is more effective for the state feedback releases.     

Increased control of thrips and aphids in greenhouses with two species of generalist predatory bugs involved in intraguild predation

The combined release of species of generalist predators can enhance multiple pest control when the predators feed on different prey, but, in theory, predators may be excluded through predation on each other. This study evaluated the co-occurrence of the generalist predators Macrolophus pygmaeus Rambur and Orius laevigatus (Fieber) and their control of two pests in a sweet pepper crop. Both predators consume pollen and nectar in sweet pepper flowers, prey on thrips and aphids, and O. laevigatus is an intraguild predator of M. pygmaeus. Observations in a commercial sweet pepper crop in a greenhouse with low densities of pests showed that the two predator species coexisted for 8 months. Moreover, their distributions in flowers suggested that they were neither attracted to each other, nor avoided or excluded each other. A greenhouse experiment showed that the predators together clearly controlled thrips and aphids better than each of them separately. Thrips control was significantly better in the presence of O. laevigatus and aphid control was significantly better in the presence of M. pygmaeus. Hence, combined inoculative releases of M. pygmaeus and O. laevigatus seem to be a good solution for controlling both thrips and aphids in greenhouse-grown sweet pepper. The predators are able to persist in one crop for a sufficiently long period and they complement each other in the control of both pests. This study also provides further evidence that intraguild predation does not necessarily have negative effects on biological control.     

Global stability and optimisation of a general impulsive biological control model

An impulsive model of augmentative biological control consisting of a general continuous predator-prey model in ordinary differential equations, i.e. a meta-model, augmented by a discrete part describing periodic introductions of predators is considered. The existence of an invariant periodic solution that corresponds to prey eradication is shown and a condition ensuring its global asymptotic stability is given. An optimisation problem related to the preemptive use of augmentative biological control is then considered. It is assumed that the per time unit budget of biological control (i.e. the number of predators to be released) is fixed and the best deployment of this budget is sought in terms of release frequency. The cost function to be minimised is the time needed to reduce an unforeseen prey (pest) invasion occurring at a worst time instant under some harmless level. The analysis shows that the optimisation problem admits a countable infinite number of solutions. An argumentation considering the required robustness of the optimisation result with respect to the invasive prey population level and to the model parameters is then conducted. It is shown that the cost function is decreasing in the predator release frequency so that the best deployment of the biocontrol agents is to carry out as frequent introductions as possible.     

A Theoretical Approach on Controlling Agricultural Pest by Biological Controls

In this paper we propose and analyze a prey-predator type dynamical system for pest control where prey population is treated as the pest. We consider two classes for the pest namely susceptible pest and infected pest and the predator population is the natural enemy of the pest. We also consider average delay for both the predation rate i.e. predation to the susceptible pest and infected pest. Considering a subsystem of original system in the absence of infection, we analyze the existence of all possible non-negative equilibria and their stability criteria for both the subsystem as well as the original system. We present the conditions for transcritical bifurcation and Hopf bifurcation in the disease free system. The theoretical evaluations are demonstrated through numerical simulations.     

State-dependent impulsive models of integrated pest management (IPM) strategies and their dynamic consequences

A state-dependent impulsive model is proposed for integrated pest management (IPM). IPM involves combining biological, mechanical, and chemical tactics to reduce pest numbers to tolerable levels after a pest population has reached its economic threshold (ET). The complete expression of an orbitally asymptotically stable periodic solution to the model with a maximum value no larger than the given ET is presented, the existence of which implies that pests can be controlled at or below their ET levels. We also prove that there is no periodic solution with order larger than or equal to three, except for one special case, by using the properties of the LambertW function and Poincare map. Moreover, we show that the existence of an order two periodic solution implies the existence of an order one periodic solution. Various positive invariant sets and attractors of this impulsive semi-dynamical system are described and discussed. In particular, several horseshoe-like attractors, whose interiors can simultaneously contain stable order 1 periodic solutions and order 2 periodic solutions, are found and the interior structure of the horseshoe-like attractors is discussed. Finally, the largest invariant set and the sufficient conditions which guarantee the global orbital and asymptotic stability of the order 1 periodic solution in the meaningful domain for the system are given using the Lyapunov function. Our results show that, in theory, a pest can be controlled such that its population size is no larger than its ET by applying effects impulsively once, twice, or at most, a finite number of times, or according to a periodic regime. Moreover, our theoretical work suggests how IPM strategies could be used to alter the levels of the ET in the farmers' favour.     

The prey-dependent consumption two-prey one-predator models with stage structure for the predator and impulsive effects

In this paper, we consider the prey-dependent consumption two-prey one-predator models with stage structure for the predator and impulsive effects. By applying the Floquet theory of linear periodic impulsive equation, we show that there exists a globally asymptotically stable pest-eradication periodic solution when the impulsive period is less than some critical value, that is, the pest population can be eradicated totally. But from the point of ecological balance and saving resources, we only need to control the pest population under the economic threshold level instead of eradicating it totally, and thus, we further prove that the system is uniformly permanent if the impulsive period is larger than some critical value, and meanwhile we also give the conditions for the extinction of one of the two preys and permanence of the remaining species. Thus, we can use the stability of the positive periodic solution and its period to control insect pests at acceptably low levels. Considering population communities always are imbedded in periodically varying environments, and the parameters in ecosystem models may oscillate simultaneously with the periodically varying environments, we add a forcing term into the prey population's intrinsic growth rate. The resulting bifurcation diagrams show that with the varying of parameters, the system experiences process of cycles, periodic windows, periodic-doubling cascade, symmetry breaking bifurcation as well as chaos.     

Global stability and periodic orbits for two-patch predator-prey diffusion-delay models

We consider a predator-prey system where the prey can diffuse between one patch with a low level of food and without predation and one patch with a higher level of food but with predation. We assume a Volterra within-patch dynamics, and we assume further that the benefit for the predator comes also from predation in the past through an exponential-delay memory function. By homotopy techniques we prove that, if the prey diffusion is weak enough, then a nonzero globally stable equilibrium exists. This result essentially depends upon the self-regulating coefficient of the predator. If we put this coefficient equal to zero, assuming that the predator density is regulated only by predation, then we can prove the existence of a Hopf bifurcating orbit from the positive equilibrium. The main cause of periodic orbits is the time delay in the predator response functional. We prove that diffusion, lack of delay in the predator response, and increase in the rate of the exponential decay of the memory play stabilizing roles.     

Integrated pest management models and their dynamical behaviour

Two impulsive models of integrated pest management (IPM) strategies are proposed, one with fixed intervention times and the other with these unfixed. The first model allows natural enemies to survive but under some conditions may lead to extinction of the pest. We use a simple prey-dependent consumption model with fixed impulsive effects and show that there exists a globally stable pesteradication periodic solution when the impulsive period is less than certain critical values. The effects of pest resistance to pesticides are also studied. The second model is constructed in the light of IPM practice such that when the pest population reaches the economic injury level (EIL), a combination of biological, cultural, and chemical tactics that reduce pests to tolerable levels is invoked. Using analytical methods, we show that there exists an orbitally asymptotically stable periodic solution with a maximum value no larger than the given Economic Threshold (ET). The complete expression for this periodic solution is given and the ET is evaluated for given parameters.We also show that in some cases control costs can be reduced by replacing IPM interventions at unfixed times with periodic interventions. Further, we show that small perturbations of the system do not affect the existence and stability of the periodic solution. Thus, we provide the first demonstration using mathematical models that an IPM strategy is more effective than classical control methods.     

Effect of scavenging on predation in a food web

Scavenging can have important consequences for food web dynamics, for example, it may support additional consumer species and affect predation on live prey. Still, few food web models include scavenging. We develop a dynamic model that includes two facultative scavenger species, which we refer to as the predator or scavenger species according to their natural scavenging propensity, as well as live prey, and a carrion pool to show ramifications of scavenging for predation in simple food webs. Our modeling suggests that the presence of scavengers can both increase and decrease predator kill rates and overall predation in model food webs and the impact varies (in magnitude and direction) with context. In particular, we explore the impact of the amount of dynamics (exploitative competition) allowed in the predator, scavenger, and prey populations as well as the direction and magnitude of interference competition between predators and scavengers. One fundamental prediction is that scavengers most likely increase predator kill rates, especially if there are exploitative feedback effects on the prey or carrion resources like is normally observed in natural systems. Scavengers only have minimal effects on predator kill rate when predator, scavenger, and prey abundances are kept constant by management. In such controlled systems, interference competition can greatly affect the interactions in contrast to more natural systems, with an increase in interference competition leading to a decrease in predator kill rate. Our study adds to studies that show that the presence of predators affects scavenger behavior, vital rates, and food web structure, by showing that scavengers impact predator kill rates through multiple mechanisms, and therefore indicating that scavenging and predation patterns are tightly intertwined. We provide a road map to the different theoretical outcomes and their support from different empirical studies on vertebrate guilds to provide guidance in wildlife management.     

Intraguild predation reduces redundancy of predator species in multiple predator assemblage

1. Interference between predator species frequently decreases predation rates, lowering the risk of predation for shared prey. However, such interference can also occur between conspecific predators. 2. Therefore, to understand the importance of predator biodiversity and the degree that predator species can be considered functionally interchangeable, we determined the degree of additivity and redundancy of predators in multiple- and single-species combinations. 3. We show that interference between two invasive species of predatory crabs, Carcinus maenas and Hemigrapsus sanguineus, reduced the risk of predation for shared amphipod prey, and had redundant per capita effects in most multiple- and single-species predator combinations. 4. However, when predator combinations with the potential for intraguild predation were examined, predator interference increased and predator redundancy decreased. 5. Our study indicates that trophic structure is important in determining how the effects of predator species combine and demonstrates the utility of determining the redundancy, as well as the additivity, of multiple predator species.     

Optimum timing for integrated pest management: Modelling rates of pesticide application and natural enemy releases

Many factors including pest natural enemy ratios, starting densities, timings of natural enemy releases, dosages and timings of insecticide applications and instantaneous killing rates of pesticides on both pests and natural enemies can affect the success of IPM control programmes. To address how such factors influence successful pest control, hybrid impulsive pest-natural enemy models with different frequencies of pesticide sprays and natural enemy releases were proposed and analyzed. With releasing both more or less frequent than the sprays, a stability threshold condition for a pest eradication periodic solution is provided. Moreover, the effects of times of spraying pesticides (or releasing natural enemies) and control tactics on the threshold condition were investigated with regard to the extent of depression or resurgence resulting from pulses of pesticide applications. Multiple attractors from which the pest population oscillates with different amplitudes can coexist for a wide range of parameters and the switch-like transitions among these attractors showed that varying dosages and frequencies of insecticide applications and the numbers of natural enemies released are crucial. To see how the pesticide applications could be reduced, we developed a model involving periodic releases of natural enemies with chemical control applied only when the densities of the pest reached the given Economic Threshold. The results indicate that the pest outbreak period or frequency largely depends on the initial densities and the control tactics.     

具有阶段结构和群体防卫的脉冲捕食系统

讨论了食饵具有群体防卫和捕食者具有阶段结构的脉冲控制捕食系统,根据Floquet乘子理论和脉冲比较定理,获得了食饵（害虫）灭绝周期解局部稳定与系统持续生存的充分条件．利用Matlab软件对害虫灭绝周期解和害虫周期爆发现象进行了数值模拟,并揭示了诸如高倍周期振荡,混沌,吸引子突变等复杂的动力学现象．得出的结论为害虫治理提供了可靠的策略依据．     

Indirect effects between shared prey: Predictions for

Suppression of a target prey by a predator can depend on its surrounding community, including the presence of nontarget, alternative prey. Basic theoretical models of two prey species that interact only via a shared predator predict that adding an alternative prey should increase predator numbers and ultimately lower target pest densities as compared to when the target pest is the only prey. While this is an alluring prediction, it does not explain the numerous responses empirically observed. To better understand and predict the indirect interactions produced by shared predation, we explore how additional prey species affect three broad ecological mechanisms, the predator's reproductive, movement, and functional responses. Specifically, we review current theoretical models of shared predation by focusing on these mechanisms, and make testable predictions about the effects of shared predation. We find that target predation is likely to be higher in the two prey system because of predator reproduction, especially when: predators are prey limited, alternative or total prey density is high, or alternative prey are available over time. Target predation may also be greater because of predator movement, but only under certain movement rules and spatial distributions. Predator foraging behavior is most likely to cause lower target predation in the two-prey system, when per capita predation is limited by something other than prey availability. It is clear from this review that no single theoretical generalization will accurately predict community-level effects for every system. However, we can provide testable hypotheses for future empirical and theoretical investigations of indirect interactions and help enhance their potential use in biological control.     

具有脉冲效应的多种群竞争-捕食系统的正周期解的存在性

本文研究一类具Beddington-DeAngeli类功能性反应和脉，中的多种群竞争一捕食系统，运用Gaines和Mawhin’s的重合度理论给出系统存在正周期解的一个充分条件．     

The sterile release method for population control with interspecific competition

A differential equations model of competing species with the release of sterile individuals of one of the species is examined. The system is found to have two positive steady states for certain parameter values; one of these is stable and the other is unstable. The system is quite resilient around the stable steady state. The release of steriles causes the nontarget species to increase in numbers. There exists a value of the release rate above which the pest species collapses to extinction. The existence of the competitor species assists the sterile release program since the pest equilibrium at any release rate is lower with the competitor species present than without it; in addition the release rate required to cause collapse of the pest species is lower with the competitor species present than without it. The effects of the parameters on the ease of eradication were examined. It was found that the ideal competitor species should have a high rate of increase, a large carrying capacity and exert strong competitive depression on the pests. The ideal pest would have a low rate of increase, a low carrying capacity and be a poor competitor.     

Non-interactive multiple predator effects on tadpole survival

Interactions among and within three species of predators were estimated in terms of their effects on prey survival using short-term predation experiments. The prey were tadpoles (Rana temporaria), and the predators were dragonfly larvae (Anax imperator), newts (Triturus alpestris), and backswimmers (Notonecta glauca). Mortality rate per predator imposed by Triturus and Notonecta did not decline with predator density, whereas the predation rate of Anax was strongly reduced when the number of predator individuals increased. Impacts of all three predators were not altered by the presence of other species in pairwise combinations. This system is therefore characterized by interference between individual dragonflies but relatively independent effects of predator species. These results were largely predictable based on the natural history of the predators and are encouraging for attempts to model communities as assemblages of interacting species.