Two Generalized Predator-Prey Models for Integrated Pest Management with Stage Structure and Disease in the Prey Population
Abstract
Stage-structured predator-prey models with disease in the prey are constructed. For the purpose of integrated pest management, two types of impulsive control strategies (impulsive release of infective prey and impulsive release of predator) are used. For Case 1, infective prey applications are more frequent than releases of predator (natural enemies). For Case 2, predator (natural enemies) releases are more frequent than infective prey applications. In both cases, we get the sufficient conditions for the global attractivity of the susceptible prey-eradication periodic solution. In addition, the persistence of the systems is also discussed. At last, the results are discussed and some possible future work is put forward.
1. Introduction
Pests, such as insects, mice and other animals, unwanted plants (weeds), fungi, microorganisms, and so forth, are living organisms that occur where they are not wanted or that cause damage to crops or other animals. How to minimize the loss caused by the injurious insects and injurious germ carrier to the important plants, animals, and human being is always the common problem concerned by the entomologists and society. Human has adopted some advanced and modern weapons such as chemical pesticides, biological pesticides, remote sensing and measuring, and so on to deal with pests, and some great achievements have been obtained [1–7].
The traditional chemical control only care about the current effect, but seldom take the influence on the ecosystem into consideration. And it caused many problems such as environment pollution, pest resistance to the pesticide and pest reemergence, and the like. In this regard, it has been observed that beneficial insects are often more susceptible to chemical pesticides than the target pests are. In the same time, the concentration of the pesticides in use tends to increase with time and usage, since many pests develop resistance to these chemicals. This kind of pest management strategy was considered by many authors [8–12]. At present more and more people are concerned about the effects of pesticide residues on human health and on the environment.
Compared to chemical treatment, nonchemical methods are safer to man and are generally effective for longer periods of time. One example of nonchemical pest control methods is biological treatment [13–17], including microbial control with pathogens, as disease can be important natural controls of some pests. Insects, like humans and other animals, can be infected by disease-causing organisms such as bacteria, viruses, and fungi.
People also use natural enemy to control pest or regulate it to densities below the threshold for economic damage. Often with augmentation or release, the natural enemy is applied like a pesticide after the pest has reached or exceeded the economic threshold. There are many literatures concerning natural enemy for pest control [18–25].
Many kinds of predator-prey models have been studied extensively [1, 2, 6, 20, 26–28]. In the natural world there are many species whose individual members have a life history that takes them through two stages: immature and mature. In particular, we have in mind mammalian populations and some amphibious animals, which exhibit these two stages. In recent years, stage-structured models, with or without delays, have been studied by several authors [26–33]. In addition, there are many control methods and results for complex dynamical network model [34–38], from which we can learn for the proof of our main results.
Motivated by [1, 14, 24, 25, 33, 34], in this paper, we will consider predator-prey models with stage structure in the prey. The prey stands for the pest population and the predator stands for the natural enemy population. That is, we call the pest and natural enemy as prey and predator, respectively. Here, the pest population will be controlled by releasing natural enemy and infective pests together. The infective pests can be cultivated in the laboratory and the natural enemy can be migrated from other regions. Once the susceptible pest meets with the infective pest, there is a chance to be infected. The infective pests have more possibility of death due to the disease and have less damage to the crops and environment. In fact, there is such example: salt cedar leaf beetle is a pest, and it is hatched from eggs. We call the egg stage as immature pest, and mature pest after it is hatched. In view of its eggshell, pathogens may not be effective against pest eggs. That is, the disease only attacks the mature susceptible pest. Birds are the natural enemy of the beetle, and we call them predator.
The organization of this paper is as follows. In the next section, the main biological assumptions on which the models rely are formulated and the models are constructed. In Section 3, to prove our main results we give several definitions, notations, and lemmas. In Section 4, we analyze the first case and determine the sufficient conditions for the global attractivity of the susceptible pest-eradication periodic solution and permanence of the system (5). In Section 5, we analyze the second case by similar method and obtain the sufficient conditions for the global attractivity of the susceptible pest-eradication periodic solution and permanence of the system (6). In the last section, a brief discussion and some possible future work for pest management are provided.
2. Model Formulation
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(H1) We suppose that the infective prey can neither produce offspring nor attack crops due to the disease, and only the susceptible prey can reproduce. At any time t > 0, birth into the immature prey population is proportional to the existing susceptible mature prey population with proportionality r. The immature prey population will transfer to the mature prey class after its birth with a maturity period of τ. The term represents the immature prey that were born at time t − τ (i.e., rS(t − τ)) and still survive at time t (with the immature prey death rate d1), and therefore represent the transformation from immature prey to mature prey.
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(H2) The immature prey population has the natural death rate d1. The parameters d2, d3 are the death rate for the susceptible and infective mature prey, respectively. The predator population has the natural death rate d4.
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(H3) We also suppose the natural enemy only capture the susceptible mature prey population, since the immature and infective prey population are hidden in the sanctuary, and the predation functional response is type Holling II, δ (0 < δ < 1) is the conversion rate for predation. The mature prey is divided into two classes, susceptible and infective. The incidence rate is classic bilinear βS(t)I(t), and β is the contact number per unit time for every infective prey with susceptible prey.
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(H4) We assume τn (n = 1,2, …) and λm (m = 1,2, …) are impulsive point series at which the infective prey and natural enemies are released, with the releasing amounts q1 and q2, respectively. According to the above assumptions, we have two different cases as follows.
Case 1. The releases of infective pests are more frequent than releases of natural enemies.
Case 2. Natural enemies releases are more frequent than the release of the infective pests.
3. Preliminary
We give some definitions and lemmas which will be useful for stating and proving our main results.
Definition 1. System (5) and (6) are said to be permanent if there are constants m, M > 0 (independent of the initial values) and a finite time T0 such that for all solution (S(t), I(t), y(t)) with initial conditions (7), m ≤ S(t) ≤ M, m ≤ I(t) ≤ M, m ≤ y(t) ≤ M hold for all t ≥ T0. Here T0 may depend on the initial values.
Lemma 2 (see [39].)Consider the following equation:
- (i)
If a > b, then lim t→∞x(t) = (a − b)/c.
- (ii)
If a ≤ b, then lim t→∞x(t) = 0.
Lemma 3 (see [40].)Let V : R+ × Rn → R and V ∈ V0. Assume that
Note that if one has some smoothness conditions of g to guarantee the existence and uniqueness of solutions for (11), then R(t) is exactly the unique solution of (11).
Lemma 4 (see [1].)System
Lemma 5. There exists a positive constant M such that S(t) ≤ M, I(t) ≤ M, y(t) ≤ M, for each solution (S(t), I(t), y(t)) of system (5) with positive initial values (7), where t is large enough.
Proof. Define a function V such that
Consider system
According to Lemmas 3 and 4, we derive
4. Analysis of System (5)
In this section, we determine the global attractive condition for the susceptible pest-eradication periodic solution and permanence of system (5).
Theorem 7. The susceptible pest-eradication periodic solution of system (5) is globally attractive provided that
Proof. Since (22) holds, we can choose a sufficiently small ϵ > 0 such that
Thus, for t > n1TN + τ, we have
From system (5), for all t > n2TN we have,
Corollary 8. (1) If q1 = 0, then the condition (22) of Theorem 7 becomes
(2) If q2 = 0, then the condition (22) of Theorem 7 becomes
Through Theorem 7 and Corollary 8, we can get the sufficient conditions for global attractivity of the susceptible pests-eradication periodic solution. That is, the susceptible pest population is eradicated totally. But in practice, from the view point of keeping ecosystem balance and preserving biological resources, it is not necessary to eradicate the susceptible pest population completely. In fact we hope the susceptible pests and natural enemies can coexist, and at the same time the susceptible pests do not cause immense economic loss. Thus, it is meaningful to study the permanence of system (5).
Theorem 9. The system (5) is permanent provided that
Proof. Since (34) holds, we can choose sufficiently small positive number ϵ, η4 (0 < η4 < min ((d3/β), (d4/(δa − bd4)))) such that
We claim that for any t0 > 0, it is impossible that S(t) < η4 for all t ≥ t0. Suppose that the claim is not true. Then there is a t0 > 0 such that S(t) < η4 for all t ≥ t0.
Then, from system (5) we have
We will show that S(t) ≥ S0 for all t > T1. Otherwise there exists a nonnegative constant T2 such that S(t) ≥ S0 for t ∈ [T1, T1 + τ + T2], S(T1 + τ + T2) = S0 and . Thus from the first equation of system (5) and (35) we easily see that
From (37) we have for all t > T1
By the claim, we need to consider two cases.
Case 1. (S(t) > η4) for all large t, then our aim is obtained.
Case 2. (S(t)) oscillates about η4 for all large t. Denote
If θ ≤ T3, our aim is obtained.
If T3 < θ < τ, from the first equation of system (5) we have, for
It is clear that S(t) ≥ η3 for .
If θ > τ, then we have that S(t) ≥ η3 for . Then, proceeding exactly as the proof for above claim, we see that S(t) ≥ η3 for . Since this kind of interval is arbitrarily chosen (we only need to be large), we can get that S(t) ≥ η3 for all t large enough. In view of our arguments above, the choice of η3 is independent of the positive solution of system (5) which satisfies that S(t) ≥ η3 for sufficiently large t. This completes the proof of the theorem.
Corollary 10. (1) If q1 = 0, then condition (34) of Theorem 9 becomes
(2) If q2 = 0, then condition (34) of Theorem 9 becomes
5. Analysis of System (6)
In this section we will discuss system (6), the condition for the global attractive of the susceptible pest-eradication periodic solution, and the permanence of system (6) will be obtained.
Thus, system (6) has a unique nonnegative periodic solution (0, I*(t), y*(t)), which is called as the susceptible pest-eradication periodic solution. Next, we will discuss the global attractivity of this periodic solution and the permanence of system (6).
Theorem 11. The susceptible pest-eradication periodic solution of system (6) is attractive provided that
Proof. Since (53) holds, we can choose a sufficiently small ϵ > 0 such that
Thus, from system (6), for all t > n2Tk we have
Corollary 12. (1) If q1 = 0, then condition (53) of Theorem 11 becomes
(2) If q2 = 0, then condition (53) of Theorem 11 becomes
Through Theorem 11 and Corollary 12, we can get the sufficient conditions for the susceptible pests-eradication periodic solution; that is, the susceptible pest population is eradicated totally. But in practice, from the view point of keeping ecosystem balance and preserving biological resources, it is not necessary to eradicate the susceptible pest population. In fact we hope the susceptible pests and enemies can coexist when the susceptible pests do not cause immense economic loss. Thus, it is necessary to discuss the permanence of system (6).
Theorem 13. System (6) is permanent provided that
Proof. Since (65) holds, we can choose sufficiently small positive number ϵ, h4 (h4 < min (d3/β, d4/(δa − bd4)) such that
Thus we only need to find h3 > 0 such that S(t) > h3 for t large enough. We claim that for any t0 > 0, it is impossible that S(t) < h4 for all t ≥ t0. Suppose that the claim is not true. Then there is a t0 > 0 such that S(t) < h4 for all t ≥ t0. Then, from system (6) for all t ≥ t0 we have
We will show that S(t) ≥ S0* for all t > T1. Otherwise there exists a nonnegative constant T2 such that S(t) ≥ S0* for t ∈ [T1, T1 + τ + T2], S(T1 + τ + T2) = S0* and . Thus from the first equation of system (6) and (66) we easily see that
By the claim, we need to consider two cases.
Case 1. S(t) > h4 for all large t, then our aim is obtained.
Case 2. S(t) oscillates about h4 for all large t. Denote
If θ < T3, then our aim is obtained.
If T3 < θ < τ, from the first equation of system (6) we have
If θ > τ, then we have that S(t) ≥ h3 for . Then, proceeding exactly as the proof for above claim, we see that S(t) ≥ h3 for . Since this kind of interval is arbitrarily chosen (we only need to be large), we can get that S(t) ≥ h3 for all t large enough. In view of our arguments above, the choice of h3 is independent of the positive solution of system (6) which satisfies that S(t) ≥ h3 for sufficiently large t. This completes the proof of the theorem.
Corollary 14. (1) If q1 = 0, then condition (65) of Theorem 13 becomes
(2) If q2 = 0, then the condition (65) of Theorem 13 becomes
6. Discussion
In this paper, a stage-structured predator-prey model with disease in the prey is considered. The prey stands for the pest population, and the predator stands for the natural enemy population. For the purpose of integrated pest management, two types of impulsive control strategies are used. In Case 1, infective prey applications are more frequent than releases of natural enemies. In Case 2, natural enemies releases are more frequent than releases of infected prey. In both cases, we analyzed the global attractivity of the susceptible pest-eradication periodic solution of the systems, and we also obtained the condition for the permanence of the systems.
In Section 4, the first case is analyzed. By Theorem 7, the sufficient condition for the global attraction of the susceptible pest-eradication periodic solution is obtained, which means that if the release amount of infective pest and natural enemy satisfy certain conditions, then the susceptible pest will be doomed. By the result of Theorem 9, the sufficient condition for the permanence of system (5) is also obtained, which means that the pest and the natural enemy will coexist for all time, if the release amounts of infective pest and natural enemy meet some critical values. Corollary 8 and 10 show that if only one control measure is taken, that is, either only infective pests are released or only natural enemies are released, then the release amount must satisfy certain conditions.
In Section 5, we use similar method to analyze the second case. In Theorem 11, the sufficient condition for the global attraction of the susceptible pest-eradication periodic solution is obtained, which means that if the release amounts of infective pest and natural enemy meet certain conditions, then the susceptible pest will be doomed. By the result of Theorem 13, the sufficient condition for the permanence of system (6) is also obtained, which means that the pest and the natural enemy will coexist for all time, if the release amounts of infective pest and natural enemy meet some critical values. Corollary 12 and 14 show that if only one measure is taken, that is, either only infective pests are released or only natural enemies are released, then the release amount must satisfy certain conditions.
From the above theoretical results obtained in Sections 4 and 5, we can choose different control strategies to control pest. We can do it by releasing infected pest more frequently than releasing natural enemy, or vice versa. However, in practice, every control measure needs certain cost, and we must think about the cost before adopting a measure. For example, the infective prey (pest) being cultivated in the laboratory and the natural enemy (predator) being migrated from other regions all need some cost. We can choose different pest management methods with different costs. In both models, different parameters mean different cost for control measures. Therefore, we have an interesting problem. In the view point of economy, under the premise of controlling the pest, which kind of control method is more suitable? This will be an optimal control problem, and we leave it as our future work.
Acknowledgments
The authors would like to thank the referees for their helpful suggestions, which improved the quality of this paper greatly. The first author is supported by Postdoctoral Science Foundation of China (no. 2011M501428). The second author is supported by NNSF of China (nos. 11171199, 81161120403).