Research Article | Open Access

Na Zhang, Fengde Chen, Qianqian Su, Ting Wu, "Dynamic Behaviors of a Harvesting Leslie-Gower Predator-Prey Model", *Discrete Dynamics in Nature and Society*, vol. 2011, Article ID 473949, 14 pages, 2011. https://doi.org/10.1155/2011/473949

# Dynamic Behaviors of a Harvesting Leslie-Gower Predator-Prey Model

**Academic Editor:**Prasanta K. Panigrahi

#### Abstract

A Leslie-Gower predator-prey model incorporating harvesting is studied.
By constructing a suitable Lyapunov function, we show that the unique positive
equilibrium of the system is globally stable, which means that suitable harvesting
has no influence on the persistent property of the harvesting system. After that,
detailed analysis about the influence of harvesting is carried out, and an interesting finding
is that under some suitable restriction, harvesting has no influence on the final
density of the prey species, while the density of predator species is strictly decreasing
function of the harvesting efforts. For the practical significance, the economic profit
is considered, sufficient conditions for the presence of bionomic equilibrium are given,
and the optimal harvesting policy is obtained by using the *Pontryagin's* maximal
principle. At last, an example is given to show that the optimal harvesting policy
is realizable.

#### 1. Introduction

Leslie [1, 2] introduced the following predator-prey model, where the “carrying capacity” of the predator's environment is proportional to the number of prey: where and are the density of prey species and the predator species at time , respectively. The above system admits an unique coexisting fixed point Recently, by constructing a suitable Lyapunov function, Korobeinikov [3] showed the positive equilibrium is globally stable; consequently, the system could not admits limit cycle. Such an finding is very interesting, since for predator-prey system incorporating Holling-type II or III functional response, limit cycle exists [4, 5]. After the work of Korobeinikov, many scholars have done works on Leslie-type predator prey ecosystem. Aguirre et al. [6] showed that Leslie-Gower predator-prey model with additive Allee effect is possible to admit two limit cycles; Aziz-Alaoui and Daher Okiye [7] argued that a suitable predator prey model should incorporate some kind of functional response, they proposed a predator-prey model with modified Leslie-Gower and Holling-type II schemes, they investigated the boundedness and global stability of the system; Nindjin et al. [8] further incorporated the time delay to the system considered in [7], and they showed that time delay plays important role on the dynamic behaviors of the system; Yafia et al. [9] studied the limit cycle bifurcated from time delay; Nindjin and Aziz-Alaoui [10], and Aziz-Alaoui [11] studied the dynamic behaviors of three Leslie-Gower-type species food chain system; Chen et al. [12] incorporated a prey refuge to system (1.1) and showed that the refuge has no influence on the persistent property of the system; Some scholars argued that nonautonomous case are more realistic if one consider the influence of seasonal effect of the environment. Huo and Li [13] studied the periodic solution of the nonautonomous case Leslie-Gower predator-prey system; Gakkhar and Singh [14] studied a Leslie-Gower predator-prey system with seasonally varying parameters; Song and Li [15] further considered the influence of impulsive effect. For more works on predator-prey ecosystem, one could refer to [4–27] and the references cited therein.

As was pointed out by Makinde [28]: “From the point of view of human needs, the exploitation of biological resources and harvesting of populations are commonly practiced in fishery, forestry, and wildlife management. There is a wide range of interest in the use of bioeconomic models to gain insight into the scientific management of renewable resources like fisheries and forestries.” Though there are numerous works on predator-prey system incorporating the harvesting, to this day, still no scholar studies the system (1.1) under the assumption of the harvesting on prey and predator species. In this paper, we assume that the predator and prey species in the model is both of commercial importance and they are subjected to constant effort harvesting with and , two parameters that measures the effort being spent by a harvesting agency. Thus, we formulating the system as follows: where and are the density of prey species and the predator species at time , respectively. To ensure the sustainable development, which means that we try to control the prey and predator species densities in a controllable range, but not to perish the species, it's natural to assume that , .

The rest of the paper is arranged as follows: we will study the stability property of positive equilibrium of system (1.3) in Section 2 and discuss the influence of the harvesting in Section 3. Bionomic equilibrium and optimal harvesting policy for system (1.3) are discussed in Section 4 and 5, respectively. An example of system (1.3) is given in Section 6 to show the feasibility of our results. We end this paper by a briefly discussion.

#### 2. Stability Property of Positive Equilibrium

By simple computation, under the assumption , , system (1.3) admits an unique positive equilibrium Obviously, satisfies the equalities

Our result about the local stability property of this equilibrium is stated as follows.

Theorem 2.1. *The positive equilibrium of system (1.3) is locally asymptotically stable.*

*Proof. *The variational matrix of the system (1.3) is given by
So, the characteristic equation for is given by , where
It is clear that the roots of the characteristic equation are negative or have negative real parts. Hence, the unique positive equilibrium of system is stable. This completes the proof of the Theorem 2.1.

Concerned with the global stability property of the positive equilibrium, we have the following.

Theorem 2.2. *The positive equilibrium of system (1.3) is globally stable.*

*Definition 2.3. *System (1.3) is called permanent if for any positive solution of system (1.3) there exist positive constants , , , which are independent of the solution of the system, such that

Theorem 2.2 shows that
Noticing that and are only dependent on the coefficients of the system (1.3) and independent of the solution of system (1.3). Thus, (2.6) clearly shows that suitable harvesting (more precisely, with restriction , ) has no influence on the persistent property of the system.

*Proof of Theorem 2.2. *We will adapt the idea of Korobeinikov [3] to prove Theorem 2.2. More precisely, we construct the following Lyapunov function:
Obviously, is well defined and continuous for all . By simple computation, we have
Equation (2.8) shows that the positive equilibrium is the only extremum of the function in the positive quadrant. Noting that
Therefore,
Above analysis shows that is the only minimum extremum of the function in the positive quadrant. One could easily verify that
From (2.8) and (2.11), we can see that the positive equilibrium is the global minimum, that is,
for all .

Calculating the derivative of along the solution of the system (1.3), by using equalities (2.2), we have
Obviously, strictly for all except the positive equilibrium , where . Thus, satisfies Lyapunov's asymptotic stability theorem, and the positive equilibrium of system (1.3) is globally stable. This ends the proof of Theorem 2.2.

*Remark 2.4. *With the restriction , , system (1.3) always admits an unique positive equilibrium and from Theorems 2.1 and 2.2 we can see that this equilibrium is globally attractive, since it's stability property is not changed with the variation of parameter , the system could not undergoes Hopf's bifurcation and there is no limit cycle of system (1.3) in . In fact, we can also prove this declare by using *Bendixson-Dulac* theorem.

Let Obviously, , , and .

Calculating from above equations we get

According to *Bendixson-Dulac* theorem, we know that there is no limit cycle of system (1.3) in .

#### 3. The Influence of Harvesting

We will discuss this topic on three aspects.

*(1) The case of only harvesting prey species*

In this case,
Obviously, , are all continuous differentiable function of parameter and
The above inequalities show that and are both the strictly decreasing function of , that is, increasing the capture rate of prey species leads to the decreasing of the density of both prey and predator species.

*(2) The case of only harvesting predator species*

In this case,
that is, , are all continuous differentiable function of parameter . Noticing that
It is easy to see that is the strictly increasing function of parameter , while is the strictly decreasing function of , that is, increasing the capture rate of predator species leads to the increasing the density of prey species and the decreasing of predator species.

*(3) The case of harvesting predator and prey species together*

In this case, it follows from (2.1) that and are all continuous differential functions of parameters , . Though we had made the assumption , it still not an easy thing to give an detailed analysis of all of the cases. Here, we only investigate the following problem, which seems very interesting.

*Problem 3.1. *Is it possible to choose some suitable parameters such that after the harvesting of predator and prey, the densities of prey species as still has no change? That is, . If this is possible, what about the dynamic behaviors of predator species in this case?

The first part of the question is equivalent to say that in what case , that is, Solving the above equality, we obtain It means that with the suitable capture efforts ( which satisfy the equality (3.6)), prey species will converge to as .

Now substituting (3.6) into the second equality of (2.1), we have Obviously, is the continuous differential function of parameter and that is, if the capture rate of predator and prey species satisfies (3.6), then increasing the harvesting of prey (and of predator) will lead to the finally decreasing of predator densities ().

#### 4. Bionomic Equilibrium

This section is devoted to study the bionomic equilibrium of system (1.3) since it has the practical significance.

The term bionomic equilibrium is an amalgamation concepts of biological equilibrium and economic equilibrium. As we know, a biological equilibrium is given by . And the economic equilibrium is said to be achieved when the TR (total revenue obtained by selling the harvested predators and ) equals TC (the total cost for the effort devoted to harvesting).

Some symbols should be given at first.

Let is the price per unit biomass of the prey , is the price per unit biomass of the predator , is the fishing cost per unit effort of the prey , is the fishing cost per unit effort of the predator .

Then, the economic rent (revenue at any time) is given by where , , that is, and represent the net revenues for the population and , respectively.

For convenience, we take the price per unit biomass of the predators and the fishing cost per unit effort of the predators to be constant. So, the bionomic equilibrium is given by the following simultaneous equations

Since the price and the cost of the predators are not sure, we will consider the following cases in order to determine the bionomic equilibrium.

*Case 4.1. *If
that is,
holds, that is to say the total cost exceed the total revenue for the harvesting of prey, obviously, the prey harvesting will be stopped (i.e., ) and the predator harvesting remains operational if .

Then, from (4.4), we have
Substituting it into (4.3), it follows that
Again, substituting (4.7) and (4.8) into (4.2) leads to

So, if and hold together, we have the bionomic equilibrium .

*Case 4.2. *If
that is,
holds, that is to say the total cost exceeds the total revenue for the harvesting of predator obviously, the prey harvesting will be stopped (i.e., ) and the predator harvesting remains operational if .

Then, it follows from (4.4) that
Substituting (4.12) into (4.3), we obtain
Again, substituting (4.12) and (4.13) into (4.2) leads to
So, if hold, we have the bionomic equilibrium .

*Case 4.3. *If
that is,
hold, then it is equivalent to say that the total cost exceeds the total revenue for two populations.

Obviously, the harvesting will be stopped, that is, , . In this case, there is no bionomic equilibrium.

*Case 4.4. *If
that is,
hold. In this case, the total revenue exceeds the total cost for two populations and the harvesting is in operational because it can bring profit for fishery.

From (4.4), we have

Substituting above equalities into (4.2) and (4.3), it is easy to obtain
So, if
hold together, we have the bionomic equilibrium .

It is obviously that the bionomic equilibrium may exist if the intrinsic growth rates of two species exceed some value.

#### 5. Optimal Harvesting Policy

In order to determine the optimal harvesting policy, we consider the present value of a continuous time-stream of revenue where denotes the instantaneous annual rate of discount and are the control variables, which are subject to the assumption , .

Now, our objective is to maximize subject to the state equations (1.3) by invoking *Pontryagin's* maximal principle.

The Hamiltonian for the problem is given at first where and are the adjoint variables.

Obviously, the control variables and appear linearly in the Hamiltonian function . So, the conditions are necessary for the singular control to be optimal.

Then, we have that is Therefore, the shadow prices do not vary with time in the optimal equilibrium. Hence they remain bounded as .

By the maximal principle, the adjoint variables satisfy and , for all , that is. Substituting (5.4) and (5.5) into (5.6), we get From (5.7), we may find the positive values of .

Substituting into (1.3), we get the equations as follows: Then, we may have the optimal equilibrium effort levels and , if

It means that, under the harvesting for both prey and predator species, the optimal harvesting policy can be obtained only under the assumption that the intrinsic growth rates of two species exceed some values.

The optimal harvesting policy means the economic rent for harvesting will be the maximal in the future time, with the populations of the system are permanent.

#### 6. Numerical Example

Let

From Section 6, we know that the optimal harvesting policy can be obtained by substituting the result of (5.7) into (1.3). So, the corresponding equations of (5.7) are

Solving (6.2) by using Maple, we get the results as follow:

From the above results, one could easily see that there is only one result (, ) meeting the condition In this case, the corresponding system of (1.3) is as follows:

Substituting (, ) into (6.5), we obtain the optimal harvesting efforts

#### 7. Conclusion

A Leslie-Gower predator-prey model incorporating harvesting is studied in this paper. We first show that suitable harvesting has no influence on the persistent property of the harvesting system. After that, we try to give the detail analysis of harvesting on the dynamic behaviors of the system. Our study shows that for the system having both harvesting on predator and prey species, it admits some interesting phenomenon; maybe such a finding could be applied to help human improving the scientific management of renewable resources such as fisheries and forest trees. Then, for the practical significance, we consider the economic profit of the harvesting. The bionomic equilibrium and optimal harvesting policy are studied. The results show that the optimal harvesting policy may exist. Finally, an example is given to show that the optimal harvesting policy of system (1.3) is realizable.

#### Acknowledgments

The author is grateful to the anonymous referees for their excellent suggestions, which greatly improved the presentation of the paper. Also, this work was supported by the Technology Innovation Platform Project of Fujian Province (2009J1007) and the Start-up Foundation of Science and Technology of Mingjiang University (YKQ08001).

#### References

- P. H. Leslie, “Some further notes on the use of matrices in population mathematics,”
*Biometrika*, vol. 35, pp. 213–245, 1948. View at: Google Scholar | Zentralblatt MATH - P. H. Leslie, “A stochastic model for studying the properties of certain biological systems by numerical methods,”
*Biometrika*, vol. 45, no. 1-2, pp. 16–31, 1958. View at: Google Scholar | Zentralblatt MATH - A. Korobeinikov, “A Lyapunov function for Leslie-Gower predator-prey models,”
*Applied Mathematics Letters*, vol. 14, no. 6, pp. 697–699, 2001. View at: Publisher Site | Google Scholar | Zentralblatt MATH - T. Kumar Kar, “Stability analysis of a prey-predator model incorporating a prey refuge,”
*Communications in Nonlinear Science and Numerical Simulation*, vol. 10, no. 6, pp. 681–691, 2005. View at: Publisher Site | Google Scholar | Zentralblatt MATH - Y. Huang, F. Chen, and L. Zhong, “Stability analysis of a prey-predator model with Holling type III response function incorporating a prey refuge,”
*Applied Mathematics and Computation*, vol. 182, no. 1, pp. 672–683, 2006. View at: Publisher Site | Google Scholar | Zentralblatt MATH - P. Aguirre, E. González-Olivares, and E. Sáez, “Two limit cycles in a Leslie-Gower predator-prey model with additive Allee effect,”
*Nonlinear Analysis: Real World Applications*, vol. 10, no. 3, pp. 1401–1416, 2009. View at: Publisher Site | Google Scholar | Zentralblatt MATH - M. A. Aziz-Alaoui and M. Daher Okiye, “Boundedness and global stability for a predator-prey model with modified Leslie-Gower and Holling-type II schemes,”
*Applied Mathematics Letters*, vol. 16, no. 7, pp. 1069–1075, 2003. View at: Publisher Site | Google Scholar | Zentralblatt MATH - A. F. Nindjin, M. A. Aziz-Alaoui, and M. Cadivel, “Analysis of a predator-prey model with modified Leslie-Gower and Holling-type II schemes with time delay,”
*Nonlinear Analysis: Real World Applications*, vol. 7, no. 5, pp. 1104–1118, 2006. View at: Publisher Site | Google Scholar | Zentralblatt MATH - R. Yafia, F. El Adnani, and H. T. Alaoui, “Limit cycle and numerical similations for small and large delays in a predator-prey model with modified Leslie-Gower and Holling-type II schemes,”
*Nonlinear Analysis: Real World Applications*, vol. 9, no. 5, pp. 2055–2067, 2008. View at: Publisher Site | Google Scholar | Zentralblatt MATH - A. F. Nindjin and M. A. Aziz-Alaoui, “Persistence and global stability in a delayed Leslie-Gower type three species food chain,”
*Journal of Mathematical Analysis and Applications*, vol. 340, no. 1, pp. 340–357, 2008. View at: Publisher Site | Google Scholar | Zentralblatt MATH - M. A. Aziz-Alaoui, “Study of a Leslie-Gower-type tritrophic population model,”
*Chaos, Solitons and Fractals*, vol. 14, no. 8, pp. 1275–1293, 2002. View at: Publisher Site | Google Scholar | Zentralblatt MATH - F. Chen, L. Chen, and X. Xie, “On a Leslie-Gower predator-prey model incorporating a prey refuge,”
*Nonlinear Analysis: Real World Applications*, vol. 10, no. 5, pp. 2905–2908, 2009. View at: Publisher Site | Google Scholar | Zentralblatt MATH - H.-F. Huo and W.-T. Li, “Periodic solutions of delayed Leslie-Gower predator-prey models,”
*Applied Mathematics and Computation*, vol. 155, no. 3, pp. 591–605, 2004. View at: Publisher Site | Google Scholar | Zentralblatt MATH - S. Gakkhar and B. Singh, “Dynamics of modified Leslie-Gower-type prey-predator model with seasonally varying parameters,”
*Chaos, Solitons and Fractals*, vol. 27, no. 5, pp. 1239–1255, 2006. View at: Publisher Site | Google Scholar - X. Song and Y. Li, “Dynamic behaviors of the periodic predator-prey model with modified Leslie-Gower Holling-type II schemes and impulsive effect,”
*Nonlinear Analysis: Real World Applications*, vol. 9, no. 1, pp. 64–79, 2008. View at: Publisher Site | Google Scholar | Zentralblatt MATH - P. D. N. Srinivasu and I. L. Gayatri, “Influence of prey reserve capacity on predator-prey dynamics,”
*Ecological Modelling*, vol. 181, no. 2-3, pp. 191–202, 2005. View at: Publisher Site | Google Scholar - W. Ko and K. Ryu, “Qualitative analysis of a predator-prey model with Holling type II functional response incorporating a prey refuge,”
*Journal of Differential Equations*, vol. 231, no. 2, pp. 534–550, 2006. View at: Publisher Site | Google Scholar - T. Kumar Kar, “Modelling and analysis of a harvested prey-predator system incorporating a prey refuge,”
*Journal of Computational and Applied Mathematics*, vol. 185, no. 1, pp. 19–33, 2006. View at: Publisher Site | Google Scholar | Zentralblatt MATH - E. González-Olivares and R. Ramos-Jiliberto, “Dynamic consequences of prey refuges in a simple model system: more prey, fewer predators and enhanced stability,”
*Ecological Modelling*, vol. 166, no. 1-2, pp. 135–146, 2003. View at: Publisher Site | Google Scholar - F. Chen and M. You, “Permanence, extinction and periodic solution of the predator-prey system with Beddington-DeAngelis functional response and stage structure for prey,”
*Nonlinear Analysis: Real World Applications*, vol. 9, no. 2, pp. 207–221, 2007. View at: Publisher Site | Google Scholar | Zentralblatt MATH - F. Chen and J. Shi, “On a delayed nonautonomous ratio-dependent predator-prey model with Holling type functional response and diffusion,”
*Applied Mathematics and Computation*, vol. 192, no. 2, pp. 358–369, 2007. View at: Publisher Site | Google Scholar | Zentralblatt MATH - F. Chen, Y. Chen, and J. Shi, “Stability of the boundary solution of a nonautonomous predator-prey system with the Beddington-DeAngelis functional response,”
*Journal of Mathematical Analysis and Applications*, vol. 344, no. 2, pp. 1057–1067, 2008. View at: Publisher Site | Google Scholar | Zentralblatt MATH - X. Li and W. Yang, “Permanence of a discrete model of mutualism with infinite deviating arguments,”
*Discrete Dynamics in Nature and Society*, vol. 2010, Article ID 931798, 7 pages, 2010. View at: Google Scholar | Zentralblatt MATH - R. Wu and L. Li, “Permanence and global attractivity of discrete predator-prey system with Hassell-Varley type functional response,”
*Discrete Dynamics in Nature and Society*, vol. 2009, Article ID 323065, 17 pages, 2009. View at: Publisher Site | Google Scholar - L. Chen and L. Chen, “Permanence of a discrete periodic Volterra model with mutual interference,”
*Discrete Dynamics in Nature and Society*, vol. 2009, Article ID 205481, 9 pages, 2009. View at: Google Scholar | Zentralblatt MATH - Q. Zhan, X. Xie, and Z. Zhang, “Stability results for a class of differential equation and application in medicine,”
*Abstract and Applied Analysis*, vol. 2009, Article ID 187021, 8 pages, 2009. View at: Google Scholar | Zentralblatt MATH - L. Chen, J. Xu, and Z. Li, “Permanence and global attractivity of a delayed discrete predator-prey system with general Holling-type functional response and feedback controls,”
*Discrete Dynamics in Nature and Society*, vol. 2008, Article ID 629620, 17 pages, 2008. View at: Google Scholar | Zentralblatt MATH - O. D. Makinde, “Solving ratio-dependent predator-prey system with constant effort harvesting using Adomian decomposition method,”
*Applied Mathematics and Computation*, vol. 186, no. 1, pp. 17–22, 2007. View at: Publisher Site | Google Scholar | Zentralblatt MATH

#### Copyright

Copyright © 2011 Na Zhang et al. This is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.