Abstract
We study the permanence and extinction of a generalized Gause-type predator-prey system with periodic coefficients. We provide a sufficient and necessary condition to guarantee the predator and prey species to be permanent and a sufficient condition for the existence of a periodic solution. In addition we prove that when the predator population tends to extinction, the prey population keeps oscillating above a positive population level.
1. Introduction
Permanence of a dynamical system has always been a hot issue in the past few decades. The concept of permanence has been introduced and investigated by several authors, each using his own terminology: “cooperativity” in the earlier papers of Schuster et al. [1], and Hofbauer [2], “permanent coexistence” by Hutson and Vickers [3], “uniform persistence” in Butler et al. [4], and “ecological stability” by Svirezhev and Logofet [5–7] (for more detailed statements of the concept see [8]).
Many important results have been found in recent years [1–37]. Some authors (see [21, 28, 31, 33]) have considered the following two species periodic Lotka-Volterra predator-prey system where and are periodic functions on with common period and for all . They have established sufficient and necessary conditions for the existence of positive -periodic solutions of the system by using different methods, respectively. Teng [31] has given sufficient and necessary conditions for the uniform persistence of the system.
Cui [16] has considered the permanence of the following Lotka-Volterra predator-prey model with periodic coefficients: He provided a sufficient and necessary condition to guarantee the predator and prey species to be permanent. In Theorems 2.2 and 2.3 he set a precondition that . This restricts the application of the theorems more or less, since many researchers often neglect the logistic term in the predator equation when the population level of the predator is relatively low and the competition between predators can be ignored, and it proved to be an unnecessary precondition in our paper. However, the research methods in his work inspired me, and many proofs, especially in the first half of this paper, are analogous to [16].
In this paper we consider the permanence of the following generalized Gause-type predator-prey system, where , for all and , , are all periodic continuous functions with common period ; , are positive, and is nonnegative. We emphasize that our model includes the case when .
In the absence of predators, system (1.3) becomes where is a real-valued function defined on
Vance and Coddington [35] have studied system (1.4) and proved the existence of a unique periodic solution under some assumptions. Apart from the assumption we mentioned above that is -periodic, the other assumptions with a mild modification are as follows. (A1)Function is continuous and differentiable with respect to on , and is continuous on . (A2)There are continuous functions and with for and for , such that (A3)There exist constants and , such that (i) for ,(ii) for . (A4)There exist constants and, such that(i)for ,(ii) for .
In addition, we assume that the -periodic function satisfies the following. (A5)Function is continuous with respect to , and is strictly monotonely increasing with respect to , (A6) is nonnegative and is positive when , (A7) is bounded on , that is there is a constant such that for .
Traditionally represents a grazing rate. Usually when the amount of prey increases, the grazing rate increases and eventually tends to a maximal value as the prey population tends to infinity. But here we do not emphasize that is bounded because its unnecessary theoretically. Assumption (A5) means that there is a higher capture rate when there is a larger amount of prey. is directly proportional to the mean possibility density for each individual prey being captured, and (A6), (A7) suggest that there is a possibility, but its not definitely for each individual prey being captured. In ecology is called the functional response or grazing function, for example, the Holling-type grazing function: where or the generalized form where ; the Ivlev grazing function where . Obviously all of these functions satisfy (A5)–(A7).
In this paper we will establish sufficient and necessary conditions for the permanence of system (1.3). In the next section we state our main results. These results are proved in Section 3. Two applications are given in Section 4.
2. Main Results
Throughout this paper, we will assume that all the functions , , , , and are continuous and periodic with common period . For any continuous -periodic function defined on , we denote In order to describe our main results, we first introduce a lemma.
Lemma 2.1 (see [35]). Suppose that satisfies (A1)–(A4). Then system (1.4) possesses a unique -periodic positive solution which is globally asymptotically stable with respect to the positive -axis.
Theorem 2.2. Suppose that satisfies (A1)–(A4), satisfies (A5)–(A7). Then system (1.3) is permanent provided that where is the unique periodic solution of (1.4) given by Lemma 2.1.
Theorem 2.3. Suppose that satisfies (A1)–(A4), satisfies (A5)–(A7), and then(i),(ii), for any solution of system (1.3) with positive initial conditions, where is the unique periodic solution of (1.4) given by Lemma 2.1.
By Theorems 2.2 and 2.3, we have the following corollary.
Corollary 2.4. Suppose that satisfies (A1)–(A4), satisfies (A5)–(A7). Then system (1.3) is permanent if and only if (2.2) holds.
Lemma 2.5 (see Theorem 15.5 in [30, 37]). Consider a periodic system where satisfies a local Lipschitz condition. If all solutions of the above system exist in the future and one of them is bounded, then there exists a periodic solution of period .
By Theorem 2.2 and Lemma 2.5, we have the following corollary (see proof in Section 3).
Corollary 2.6. Suppose satisfies (A1)–(A4), satisfies (A5)–(A7), and condition (2.2) holds. If in addition satisfies a local Lipschitz condition with respect to , then system (1.3) has a positive -periodic solution.
3. Proof of Our Main Results
Lemma 3.1 (see [31]). If , are -periodic functions, , for all and , then has a unique nonnegative -periodic solution which is globally asymptotically stable with respect to the positive -axis. Moreover, if , then , for all and if , then .
Lemma 3.2. Suppose that satisfies (A1)–(A4), then there exists an , such that for any , system possesses an -periodic positive solution which is globally asymptotically stable with respect to the positive -axis.
Proof. It suffices to show that if satisfies (A1)–(A4), then there exists an such that for any , satisfies (A1)–(A4). Obviously satisfies (A1)–(A3). Taking , its easy to see satisfies (A4).
From a comparison theorem (see Theorem 1.1 [37]) and Lemma 2.1, we have the following lemma.
Lemma 3.3. Suppose that satisfies (A1)–(A4), is a solution of system (1.4), and is the periodic solution of system (1.4) given by Lemma 2.1. Let be a right maximal (right minimal) solution of a scalar differential equation on , where is continuous and satisfies for all . Then the following conclusions hold.(i)If , then for .(ii)For all there exist a such that for .
Proposition 3.4. Under assumptions (A1)–(A7), there exist and , such that for all solutions of system (1.3) with positive initial values.
Proof. Obviously, is a positively invariant set of system (1.3). Given any solution of (1.3) with positive initial values, from system (1.3) we have The following equation has a globally asymptotically stable positive -periodic solution by Lemma 2.1. By Lemma 3.3, there exists , such that Let . We have By (1.3), Denote . Then we have Since , for, there is a constant , such that by the continuity of . It follows that Notice that . It's easy to show that there is an , such that By the notation , taking and , Proposition 3.4 is proved.
Proposition 3.5. Under assumption (A1)–(A7), there exists a positive constant , such that for all solutions of (1.3) with positive initial values.
Proof. Suppose that (3.13) is not true. Then there is a sequence , such that
where is the solution of (1.3) with . By assumption (A5), we can choose sufficiently small positive numbers and , such that
where
By (3.14), for the given , there exists a positive integer , such that
For the rest of this proof, we assume that . Equation (3.18) implies there exists , such that
and further
By (3.15), and Lemma 3.1, any solution of the following equation,
with positive initial conditions satisfies
Hence,
by Lemma 3.3. So there is a , such that
It follows that
By (3.16), the equation
has an -periodic positive solution which is globally asymptotically stable. Hence,
for sufficiently large and , which is a contradiction with (3.14). This completes the proof of Proposition 3.5.
Proposition 3.6. Under assumption (A1)–(A7), there exists a positive constant , such that for all solutions of system (1.3) with positive initial values.
Proof. Suppose that (3.28) is not true, then there exists a sequence such that
On the other hand, by Proposition 3.5, we have
Hence, there are time sequences and satisfying
By Proposition 3.4, for a given positive integer , there is a , such that
Because of as , there is a positive integer such that since , and hence
for . Integrating (3.33) from to yields
or
If , it leads to a contradiction. Otherwise, , we have
since is bounded. By (3.15) and (3.16), there are constants and , such that
for , , and . Equation (3.37) implies
for , . For the positive satisfying (3.16) and (3.38), we have the following two cases:(i) for all ;(ii)there exists , such that .If (i) holds, by (3.38) and (3.39) we have
This is a contradiction.
If (ii) holds, we now claim that
Otherwise, there exists such that
By the continuity of , there must exist such that
Denote as the nonnegative integer, such that . By (3.15), we have
This contradiction establishes that (3.41) is true, particularly (3.41) holds for .
By (3.31) and (3.38), we have
which is also a contradiction. This completes the proof of Proposition 3.6.
Proposition 3.7. Suppose satisfies (A1)–(A4), satisfies (A5)–(A7), and (2.2) holds. Then there exists a positive constant such that for all solutions of (1.3) with positive initial values.
Proof. By assumption (A5) and (2.2) we can choose a constant such that
where
Consider the following equation with positive parameter α:
By Lemma 3.2, (3.49) has a unique positive -periodic solution which is globally asymptotically stable since . Let be the solution of (3.49) with initial condition in which is the unique periodic solution of (1.4) given by Lemma 2.1. Hence, for the above , there exists sufficiently large , such that
By the continuity of the solution in the parameter, we have uniformly in as . Hence, for there exists , such that
So, we have
Notice that and are all -periodic, hence
Choosing a constant , we have
Suppose that (3.46) is not true. Then there exists , such that
where is the solution of (1.3) with . So, there exist , such that
and hence
Let be the solution of (3.49) with and , then
By the global asymptotic stability of , for the given , there exists , such that
Hence
and hence
from (3.54). This implies
Integrating the above inequality from to yields
as from (3.47) which is a contradiction. This completes the proof of Proposition 3.7.
Proposition 3.8. Suppose satisfies (A1)–(A4), satisfies (A5)–(A7), and (2.2) holds. Then there exists a positive constant , such that for all solutions of (1.3) with positive initial values.
Proof. Otherwise, there exists a sequence , such that
However
from Proposition 3.7. Hence there are two time sequence and satisfying the following conditions:
By Proposition 3.4, for a given integer there is a , such that
Because of as , there is a positive integer , such that as , and hence
for and . Integrating the above inequality from to , we have
or
Because of the boundedness of the function , we know that
By (3.47), there is a constant and a positive integer such that
for , and . Further,
In addition,
Let be the solution of (3.49) with and . Then
by Lemma 3.3. Further, by Propositions 3.4 and 3.6, we can choose such that
for . For , (3.49) has a unique positive -periodic solution which is globally asymptotically stable. In addition, by the periodicity of (3.49), the periodic solution is uniformly asymptotically stable with respect to the compact set . Hence, for the given in Proposition 3.7, there exists which is independent of and , such that
Thus
from (3.54). By (3.72), there exists a positive integer such that for and . So we have
since and . Hence,
from (3.75) and (3.81). Integrationg the above inequality from to yields
and hence
from (3.74). This is a contradiction. This completes the proof of Proposition 3.8. The result of Theorem 2.2 follows from Propositions 3.4–3.8.
Proof of Theorem 2.3. Suppose is the periodic solution of (1.4). For any solution of (1.3) with a positive initial value, there are two possible cases:(i)for all , ;(ii)there is a , such that . Now, in the above two cases, we prove , respectively.
(i) Suppose is a solution of (1.4) with . Comparing (1.3) and (1.4) we have and . Hence
From system (1.3), for all , we have
So
where . By the boundedness of , , and the continuity of and , there must exist a constant , such that
For all , by (3.85), there exists a , such that , for all . Hence
for . Denote , . Then
Hence
This implies
On the other hand,
By the boundedness of , , , and the continuity of , μ, and , there exists a such that
Hence
For all , , we have
It follows that
Considering (3.92), we have
(ii) Consider the second case. We claim that once there is a such that , then
Otherwise, suppose . Then
This is a contradiction since from (1.3) and (1.4) we know at the same point . So (3.99) holds.
Now we show for all , there is a such that
Otherwise, suppose for all , . Then
Suppose is a solution of (1.4) with . Then by Lemma 2.1 we have , where denotes the periodic solution of (1.4). Since , there exists a such that for . Denote . From (3.99) we know for . We show that there is an such that . In fact,
where . By the boundedness of , , and the continuity of and , there exists an , such that . Hence
Choosing , such that for , and letting
Then whenever for , we have . By assumption (A6), , and this implies . Choosing , such that and for , we have
Hence, there exists an such that
for , by (2.3) and assumption (A5). So
This is a contradiction and it implies that (3.101) holds.
Second, we show that
where
Otherwise, there exists , such that
By the continuity of , there must exist such that and for . Let be the nonnegative integer, such that , and by (2.3), (A6) and (3.99), we have
which is a contradiction. This completes the proof of conclusion (i) of Theorem 2.3.
Now we prove the second conclusion. Since (i) holds and is bounded, there is a and an , such that for . Then we have
The following auxiliary equation,
has a globally asymptotically stable positive -periodic solution by Lemma 3.2. So by Lemma 3.3, we have
This completes the proof of conclusion (ii) of Theorem 2.3.
Proof of Corollary 2.6. We claim that once satisfies a local Lipschitz condition with respect to , then the function satisfies a local Lipschitz condition with respect to and . Considering assumptions (A1)–(A7), this is clearly the case. So the uniqueness of solutions of system (1.3) is guaranteed, and by Lemma 2.5 we know that there exsits an -periodic solution . From the proof of Theorem 15.5 in [37], the initial value of the periodic solution is the fix point of the mapping which is the limit point of a subsequence of the sequence , where is the initial value of a bounded solution of system (1.3). Taking any positive and , by Theorem 2.2 we know the solution started from this point is bounded and the limit of any subsequence of the sequence is positive. So the periodic solution is positive.
4. Examples
Example 4.1. Suppose , , , . The corresponding system is We know that the periodic solution of system (3.1) is Using this formula we can compute the periodic solution of (4.1) in absence of predators System (4.1) is permanent.
Example 4.2. Taking , and the same as in Example 4.1, and , we can compute
In this circumstance the predator population tends to extinction and the prey population keeps oscillating.
Simulation results of the two examples are shown in Figure 1.
(a)
(b)