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Partial permanence and extinction on stochastic Lotka-Volterra competitive systems
Advances in Difference Equations volume 2015, Article number: 266 (2015)
Abstract
This paper discusses an autonomous competitive Lotka-Volterra model in random environments. The contributions of this paper are as follows. (a) Some sufficient conditions for partial permanence and extinction on this system are established; (b) By using some novel techniques, the conditions imposed on permanence and extinction of one-species are weakened. Finally, a numerical experiment is conducted to validate the theoretical findings.
1 Introduction
It is a usual phenomenon for two or more species who live in proximity share the same basic requirements and compete for resources, habitat, food, or territory. It is therefore very important to study the competitive models for multi-species. As we know, the well-known Lotka-Volterra model concerning ecological population modeling has received great attention and has been studied extensively owing to its theoretical and practical significance. A deterministic, competitive Lotka-Volterra system with n interacting species is described by the n-dimensional differential equation
where \(x_{i}(t)\) represents the population size of species i at time t, \(b_{i}\) is the growth rate of species i, and \(a_{ij}\) represents the effect of interspecific (if \(i\ne j\)) or intraspecific (if \(i=j\)) interaction.
On the other hand, from the biological point of view, population systems in the real world are inevitably affected by environmental noise. In practice, the growth rates are often subject to environmental noise. To obtain a more accurate description of such systems, we usually consider the stochastic perturbation of the growth rate \(b_{i}\) by an average value plus an error term. Then the intrinsic growth rate depending on time becomes
where \(\dot{B}_{i}(t)\) is a white noise. As a result, system (1.1) becomes a stochastic Lotka-Volterra competition system with n interacting components as follows:
where \(b_{i}\), \(a_{ij}\), \(\sigma_{i}\) are non-negative for \(i,j=1,2,\ldots,n\), and \(\sigma_{i}^{2}\) will be called the noise intensity matrix. Throughout this paper we always assume that the following hypothesis holds:
It is therefore necessary to reveal how the noise affects the population systems. As a matter of fact, stochastic Lotka-Volterra competitive systems have recently been studied by many authors, for example, [1–6].
In the study of population systems, permanence and extinction are two important and interesting topics, respectively meaning that the population system will survive or die out in the future, which have received much attention (see [7–18]). Luo and Mao [15] revealed that a large white noise will force the stochastic Lotka-Volterra systems to become extinct while the population may be persistent under a relatively small white noise. Li and Mao [9] investigated a non-autonomous stochastic Lotka-Volterra competitive system, and some sufficient conditions on stochastic permanence and extinction were obtained. Li et al. [11] showed that both stochastic permanence and extinction have close relationships with the stationary probability distribution of the Markov chain. Tran and Yin [16] investigated stochastic permanence and extinction for stochastic competitive Lotka-Volterra systems using feedback controls.
However, most of the existing criteria are established for stochastic general Lotka-Volterra system. Hence, one natural question arises: How to derive some criteria with less conservatism for stochastic Lotka-Volterra competitive systems? This issue constitutes the first motivation of this paper.
Moreover, most of the existing criteria are established for total permanence and total extinction. To the best of our knowledge, partial permanence and partial extinction have scarcely been investigated, which are very important properties. Is it feasible to obtain some partial permanence and extinction conditions for stochastic Lotka-Volterra competitive systems? Thus, the second purpose of this paper is to solve this interesting problem.
The rest of the paper is arranged as follows. The main results of this paper are stated in Sections 3 and 4. In Section 2, some preliminaries, definitions and lemmas are given. Sufficient conditions on persistence in mean and extinction for one-species are obtained in Section 3. Based on these sufficient conditions on one-species, sufficient criteria on partial permanence and extinction on system (1.2) are established in Section 4. Section 5 provides some numerical examples to check the effectiveness of the derived results.
2 Notation
Throughout this paper, unless otherwise specified, let \((\Omega,\mathscr{F},\{\mathscr{F}_{t}\}_{t\geq0},\mathbb{P})\) be a complete probability space with a filtration \(\{\mathscr{F}_{t}\}_{t\geq0}\) satisfying the usual conditions (i.e., it is increasing and right continuous, while \(\mathscr{F}_{0}\) contains all \(\mathbb{P}\)-null sets). Let \({B}(t)=(B_{t}^{1},\ldots,B_{t}^{m})\) be an m-dimensional Brownian motion defined on the probability space. Let \(R_{+}^{n}=\{ x\in R^{n}: x_{i}>0\mbox{ for all }1\leqslant i\leqslant n\}\).
Lemma 2.1
([19])
Assume that condition (1.3) holds. Then, for any given initial value \(x(0)\in R_{+}^{n}\), there is a unique solution \(x(t)\) to system (1.2) and the solution will remain in \(R_{+}^{n}\) with probability 1, namely
Lemma 2.2
([19])
Assume that condition (1.3) holds. Then, for any given initial value \(x(0)\in R_{+}^{n}\), the solution \(x_{i}(t)\) to system (1.2) obeys
Definition 2.1
System (1.2) is said to be persistent in mean if there exist positive constants \(\alpha_{i}\), \(\beta_{i}\) such that the solution to system (1.2) has the following property:
To proceed with our study, we consider two auxiliary stochastic differential equations
where
Lemma 2.3
Assume that condition (1.3) holds. Let \(x(t)\) be a solution to system (1.2) with \(x(0)\in R_{+}^{n}\), then we have
i.e.,
Proof
By the Itô formula, we derive that
which means
Applying the Itô formula to equation (2.2) yields
From the representations of \(y_{i}(t)\) and \(z_{i}(t)\), and by (2.6) we have
□
3 Persistence in mean and extinction of one-species
3.1 Persistence of one-species
In this section, we investigate persistence in mean and extinction of one-species for system (1.2). Now, let us present some lemmas which are essential to the proof of Theorem 3.1.
Lemma 3.1
([20])
Let condition (1.3) hold. The solution \(y_{i}(t)\) to equation (2.1) has the following property:
With the help of Lemma 3.1, we slightly improve Lemma 3.1 of [8] by weakening hypotheses posed on the coefficients of equation (2.2) as follows.
Lemma 3.2
Let condition (1.3) hold, and assume that \(b_{i}-\frac{\sigma_{i}^{2}}{2}>0\), \(b_{j}-\frac{\sigma _{j}^{2}}{2}\geq0\) (\(i\neq j\)) and \(b_{i}-\frac{\sigma_{i}^{2}}{2}-\sum_{j\ne i}\frac{a_{ij}}{a_{jj}}((b_{j}-\frac{\sigma_{j}^{2}}{2})\wedge 0)>0\). Then the solution to equation (2.2) has the property
Theorem 3.1
Let condition (1.3) and assumptions in Lemma 3.2 hold. Then the solution to system (1.2) has the following property:
which means the species i of system (1.2) is persistent in mean.
Proof
Applying the Itô formula to equation (2.1) yields
Then we have
Dividing both sides of (3.5) by t yields
Note that
This implies
By Lemma 3.1, letting \(t\to\infty\) on both sides of (3.6) yields
Combining Lemma 3.2 and (3.7), we can claim that
Now we process to show assertion (3.3). Applying the Itô formula to \(\log z_{i}(t)\) yields
Dividing both sides of (3.9) by t yields
Using Lemma 3.2 and the law of strong large number for martingale, we have
Combining Lemma 3.2 and (3.11), letting \(t\to+\infty\) on both sides of (3.9) yields
Since \(x_{i}(t)\geqslant z_{i}(t)\), assertion (3.3) is true. Therefore this theorem is proved. □
Remark 3.1
Compared with the existing literature [8], the conditions imposed on the permanence of one-species are weaker.
Applying Lemma 3.1 to system (1.2), we have the following corollary, which coincides with Theorem 3.1 in [8].
Corollary 3.1
Let condition (1.3) hold and assume that \(b_{i}-\frac{\sigma_{i}^{2}}{2}>0\), \(b_{i}-\frac{\sigma_{i}^{2}}{2}-\sum_{j\ne i}\frac{a_{ij}}{a_{jj}}(b_{j}-\frac{\sigma_{j}^{2}}{2})>0\) for all \(i=1,\ldots,n\). System (1.2) is persistent in mean.
3.2 Extinction of one-species
Theorem 3.2
Let condition (1.3) hold and \(x_{i}(t)\) be the solution to system (1.2) with positive initial value \(x_{i}(0)\). Then we have the following assertions:
-
(i)
If \(\sigma_{i}^{2}>2b_{i}\), the solution \(x_{i}(t)\) to system (1.2) has the property that
$$ \limsup_{t\to\infty}\frac{\log x_{i}(t)}{t}\leqslant b_{i}-\frac{\sigma_{i}^{2}}{2} \quad\textit{a.s.} $$(3.13)That is, the species i of system (1.2) will become extinct.
-
(ii)
If \(\sigma_{i}^{2}=2b_{i}\), the solution \(x_{i}(t)\) to system (1.2) has the property that
$$ \lim_{t\to\infty}x_{i}(t)=0 \quad \textit{a.s.} $$(3.14)That is, the species i of system (1.2) still become extinct with probability one.
Proof
The proof is rather technical, so we will divide it into two steps. The first step is to show the exponential extinction of species i when \(\sigma_{i}^{2}>2b_{i}\). The second step is to show the extinction in the case of \(\sigma_{i}^{2}=2b_{i}\).
Step 1. Applying the Itô formula to \(\log x_{i}(t)\) yields
Dividing both sides of (3.15) by t yields
Using the law of strong large number for martingales, we can claim that
Letting \(t\to\infty\) yields
Step 2. Now, let us finally show assertion (3.14). Decompose the sample space into three mutually exclusive events as follows:
When \(\sigma_{i}^{2}=2b_{i}\), equation (3.16) has the following form:
Furthermore, we decompose the sample space into the following two mutually exclusive events according to the convergence of \(\int_{0}^{\infty}x_{i}(s)\,ds\):
The proof of \(\lim_{t\to\infty}x_{i}(t)=0\) a.s. is equivalent to showing \(E_{i1}\subset\Omega_{i3}\), \(E_{i2}\subset\Omega_{i3}\) a.s. The strategy of the proof is as follows.
- ⋆:
-
First, by using the techniques proposed in [21], we show that \(E_{i1}\subset\Omega_{i3}\). It is sufficient to show \(P(E_{i1}\cap\Omega_{i1})=0\) and \(P(E_{i1}\cap\Omega_{i2})=0\).
- ⋆:
-
Second, using some novel techniques, we prove that \(P(E_{i2}\cap\Omega_{i1})=0\) and \(P(E_{i2}\cap\Omega_{i2})=0\), which means \(E_{i2}\subset\Omega_{i3}\) a.s.
Now we realize this strategy as follows.
Case 1. Let us now show \(E_{i1}\subset\Omega_{i3}\). Clearly, \(x_{i}(t)\in C(R_{+},R)\) a.s. It is straightforward to see from \(E_{i1}\) that \(\liminf_{t\to\infty}x_{i}(t)=0\) a.s. Therefore, we have obtained that \(P(E_{i1}\cap\Omega_{i1})=0\). Now we only need to prove that \(P(E_{i1}\cap\Omega_{i2})=0\). We prove it by contradiction.
If \(P(E_{i1}\cap\Omega_{i2})>0\), there exists a number \(\epsilon>0\) such that
where \(J_{1}=\{\limsup_{t\to\infty}x_{i}(t)>2\epsilon\}\). Let us now define a sequence of stopping times
From \(E_{i1}\), we also have \(E(I_{E_{i1}}\int_{0}^{\infty}x_{i}(s)\,ds)<\infty\), then we compute it
where \(I_{A}\) is the indicator function for all sets A. Since \(\tau_{2k}<\infty\) whenever \(\tau_{2k-1}<\infty\), by the above formula, so we have
On the other hand, integrating equation (1.2) from 0 to t yields
A simple computation shows that
and
where \(K_{2}\) and \(K_{4}\) are defined in Lemma 2.2. Using the Hölder inequality and Burkholder-Davis-Gundy inequality (see [19]), we compute
Furthermore, we choose \(T=T(\epsilon)>0\) sufficiently small for
It then follows from (3.22) that
where
Recalling the fact that \(\tau_{k}<\infty\), for \(k=1,2,\ldots\) , whenever \(\omega\in J_{1}\), we further compute
If \(\omega\in\{\tau_{2k-1}<\infty\}\cap\{ H^{c}_{k}\cap E_{i1}\}\), note that
We derive from (3.20) and (3.24) that
which is a contraction. So that \(P(E_{i1}\cap\Omega_{i2})=0\) holds. Therefore, we obtain that \(E_{i1}\subset\Omega_{i3}\).
Case 2. Now, we turn to prove that \(E_{i2}\subset\Omega_{i3}\) a.s. It is sufficient to show \(P(E_{i2}\cap\Omega_{i1})=0\) and \(P(E_{i2}\cap\Omega_{i2})=0\). We prove it by contradiction.
If \(P(E_{i2}\cap\Omega_{i1})>0\) holds, for any \(\omega\in E_{i2}\cap\Omega_{i1}\), \(\epsilon_{0}\in(0,\frac{\gamma_{i}}{2})\), there exists \(T=(\epsilon_{0},\omega)\) such that
It then follows from (3.17) that
Letting \(t\to\infty\), we obtain that
This implies
which contradicts the definition of \(E_{i2}\) and \(\Omega_{i1}\). So \(P(E_{i2}\cap\Omega_{i1})=0\) must hold.
Now we process to show \(P(E_{i2}\cap\Omega_{i2})>0\) is false. For this purpose, we need more notations as follows:
where \(m(\Pi_{t}^{\epsilon}(i))\) indicates the length of \(\Pi_{t}^{\epsilon}(i)\). It is easy to see that \(\Delta^{0}(i)=E_{i2}\cap\Omega_{i2}\). For any \(\epsilon _{1}<\epsilon_{2}\), simple computations show that
which implies
It is easy to observe from the continuity of probability that
If \(P(E_{i2}\cap\Omega_{i2})>0\), there exists \(\epsilon>0\) such that \(P(D^{\epsilon})>0\). For any \(\omega\in\Delta^{\epsilon}(i)\), simple computations show that
By letting \(t\to\infty\), we have
Substituting (3.26) into (3.17), we obtain that
This contradicts the definition of \(E_{i2}\) and \(\Omega_{i2}\). It yields the desired assertion \(P(E_{i2}\cap\Omega_{i2})=0\) immediately. Combining the fact \(E_{i1}\subset\Omega_{i3}\), \(P(E_{i2}\cap\Omega_{i1})=0\) and \(P(E_{i2}\cap\Omega_{i2})=0\), we can claim that
The proof is completed. □
Remark 3.2
In comparison with [8] and [11], we point out that species i is still extinct when \(\sigma_{i}^{2}=2b_{i}\) by using some novel stochastic analysis techniques.
Corollary 3.2
Let condition (1.3) hold and \(x(t)\) be a solution to system (1.2) with positive initial value \(x(0)\). Assume that there exists an integer m, \(1\leqslant m< n\), such that
Then we have the following assertions:
-
(i)
For all \(i=1,\ldots,n\), the solution \(x_{i}(t)\) to system (1.2) has the property that
$$ \lim_{t\to\infty}\frac{\log x_{i}(t)}{t}= b_{i}- \frac{\sigma_{i}^{2}}{2} \quad\textit{a.s. } i=1,\ldots,m. $$(3.29) -
(ii)
For all \(i=m+1,\ldots,n\), the solution \(x_{i}(t)\) to system (1.2) has the property that
$$ \lim_{t\to\infty}\frac{\log x_{i}(t)}{t}=0 \quad\textit{a.s. } i=m+1,\ldots,n. $$(3.30)
Proof
By virtue of Theorem 3.2, for all \(\sigma_{i}^{2}>2b_{i}\), \(i=1,\ldots,m\), we obtain that
This shows that for any \(\epsilon_{i}\in(0,\frac{\sigma_{i}^{2}}{2}-b_{i})\) there is a positive random variable \(T(\epsilon_{i})\) such that
which means
Then letting \(t\to\infty\) on both sides of (3.16) yields
which is the required assertion (3.29).
Now we process to show assertion (3.30). By utilizing Theorem 3.1 and conditions (3.28), we derive
This implies
By the law of strong large numbers for martingales and (3.31), letting \(t\to\infty\) on both sides of (3.17) yields
The proof is completed. □
4 Partial permanence and extinction
Now in this section we present conditions for system (1.2) to be partially permanent and extinct. To proceed with our study, we consider the following auxiliary stochastic equation:
Theorem 4.1
Let condition (1.3) hold. Assume that there exists an integer m, \(1\leqslant m< n\), such that
Then we have the following assertions:
-
(i)
For all \(i=1,\ldots,m\), the solution \(x(t)\) to system (1.2) has the property that
$$\begin{aligned}& \limsup_{t\to\infty}\frac{1}{t}\int _{0}^{t}x_{i}(s)\,ds\leqslant \frac{1}{a_{ii}}\biggl(b_{i}-\frac{\sigma_{i}^{2}}{2}\biggr) \quad\textit{a.s. } i=1,\ldots,m. \end{aligned}$$(4.4)$$\begin{aligned}& \begin{aligned}[b] &\liminf_{t\to\infty}\frac{1}{t}\int _{0}^{t}x_{i}(s)\,ds \\ &\quad\geqslant \frac{1}{a_{ii}}\Biggl[\biggl(b_{i}-\frac{\sigma_{i}^{2}}{2}\biggr) -\sum _{k\ne i}^{m}\frac{a_{ik}}{a_{kk}} \biggl(b_{k}-\frac{\sigma_{k}^{2}}{2}\biggr)\Biggr] \quad\textit{a.s. } i=1, \ldots,m. \end{aligned} \end{aligned}$$(4.5)That is, for each \(i=1,\ldots,m\), the species i of system (1.2) is persistent in mean;
-
(ii)
For all \(i=m+1,\ldots,n\), the solution \(x(t)\) to system (1.2) has the property that
$$\begin{aligned} \limsup_{t\to\infty}\frac{\log x_{i}(t)}{t} \leqslant& b_{i}-\frac{\sigma_{i}^{2}}{2} -\sum_{j=1}^{m} \frac{a_{ij}}{a_{jj}} \Biggl[\biggl(b_{j}-\frac{\sigma_{j}^{2}}{2}\biggr) \\ &{} -\sum_{k\ne j}^{m}\frac{a_{jk}}{a_{kk}} \biggl(b_{k}-\frac{\sigma_{k}^{2}}{2}\biggr)\Biggr] \quad\textit{a.s. } i=m+1, \ldots,n. \end{aligned}$$(4.6)That is, for each \(i=m+1,\ldots,n\), the species i will become extinct.
Proof
We will divide the proof into two steps. The first step is to show the permanence of the top m species of system (1.2). The second step is to show the extinction for the bottom \(n-m\) species of system (1.2).
Step 1. Applying the Itô formula to (4.1) yields
Simple computations show that
Applying the Itô formula to \(V(t)=\sum_{i=1}^{m}|\log x_{i}(t)-\log\varPhi_{i}(t)|\) yields
Hence we get
where \(\mu=\min_{1\leqslant i\leqslant m}(a_{ii}-\sum_{j\ne i}^{m}a_{ji})>0\), \(\theta_{l}=\sum_{i=1}^{m}a_{il}\geqslant0\). We therefore have
Letting \(t\to\infty\) on both sides of (4.10) yields
By Theorem 3.2 and condition (4.3), we have
Substituting (4.12) into (4.11) yields
By virtue of the similar techniques proposed in Step 2 of Theorem 3.2, we have
When condition (4.2) is satisfied, by applying Corollary 3.1 to system (4.1), we have
A simple computation shows that
Therefore, we obtain that \(x_{i}(t)\) is persistent in mean, for all \(i=1,\ldots,m\).
Step 2. For all \(i=m+1,\ldots,n\), applying Itô to \(\log x_{i}(t)\) yields
It follows from (4.15) that
By letting \(t\to\infty\) on both sides of (4.16) yields. We can conclude that
Finally, we can get \(x_{i}(t)\) will become extinct for all \(i=m+1,\ldots,n\). The proof is completed. □
5 Numerical simulations
In this paper, we have discussed the persistence in mean and extinction of system (1.2). Moreover, sufficient conditions have been established in Theorems 3.1, 3.2 and 4.1. Thus, in this section, we give out the numerical experiment for the case \(n=2\) as follows to support to our results.
The existence and uniqueness of the solution follows from Lemma 2.1. We consider the solution with initial data \(x_{1}(0)=1.4\), \(x_{2}(0)=2.1\). By Matlab software, we simulate the solution to system (5.1) with different values of \(\sigma_{1}\) and \(\sigma_{2}\).
In Figure 1, \(\sigma_{1}=0.03\), \(\sigma_{2}=\sqrt{2.2}\). By Theorems 3.1, 3.2 and 4.1, species 1 is persistent in mean and species 2 is extinct with zero exponential extinction rate.
In Figure 2, \(\sigma_{1}=0.03\), \(\sigma_{2}=0.04\). By the conditions of Corollary 3.1, all of the species are persistent in mean.
In Figure 3, \(\sigma_{1}=\sqrt{1.8}\), \(\sigma_{2}=1.7\). By Corollary 3.2, species 1 is extinct with zero exponential extinction rate and species 2 is exponentially extinct.
6 Conclusions
This paper is devoted to partial permanence and extinction on a stochastic Lotka-Volterra competitive model. Firstly, by using some novel techniques, we established some weaker sufficient conditions on the persistence in mean and extinction for one-species. Secondly, based on these sufficient conditions for one-species and some stochastic analysis techniques, sufficient criteria for ensuring the partial permanence and extinction of the populations of the n different species in the ecosystem have been obtained. Finally, numerical experiment is provided to illustrate the effectiveness of our results.
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Acknowledgements
The authors would like to thank the editor and referees for their very important and helpful comments and suggestions. We also thank the National Natural Science Foundation of China (Grant Nos. 61304070, 11271146, 61104045, 51190102), the National Key Basic Research Program of China (973 Program) (2013CB228204).
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Dong, C., Liu, L. & Sun, Y. Partial permanence and extinction on stochastic Lotka-Volterra competitive systems. Adv Differ Equ 2015, 266 (2015). https://doi.org/10.1186/s13662-015-0608-2
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DOI: https://doi.org/10.1186/s13662-015-0608-2