Abstract
This paper investigates the problem of the global directed dynamic behaviors of a Lotka-Volterra competition-diffusion-advection system between two organisms in heterogeneous environments. The two organisms not only compete for different basic resources, but also the advection and diffusion strategies follow the dispersal towards a positive distribution. By virtue of the principal eigenvalue theory, the linear stability of the co-existing steady state is established. Furthermore, the classification of dynamical behaviors is shown by utilizing the monotone dynamical system theory. This work can be seen as a further development of a competition-diffusion system.
Keywords:
competition-diffusion-advection; principal eigenvalue; dynamic behaviors; global asymptotic stability MSC:
35B40; 35K57; 37C65; 92D25
1. Introduction
In the past few decades, the dynamic behaviors of competition-diffusion systems (see [1]) in homogeneous or heterogeneous environments have been extensively studied. Until 2017, He and Ni [2,3] studied the dynamics of two organisms by changing their random diffusion coefficients, resource carrying capacity and competitiveness, and they also described the global dynamics of two organisms. Their research has made outstanding contributions to the competition-diffusion systems. For the competition model of two organisms, either both organisms survive or win with the extinction of the other organisms (see [4,5,6]). In 2019, Du et al. [7,8] studied a Lotka-Volterra competition system with periodic habitat advection. From a biological point of view, this pulsating travel front provided a way for two competing species to interact in heterogeneous habitats. Based on the assumption that the resource function in spatial variables is decreasing, Lou et al. [9] described the competition between two aquatic organisms with different diffusion strategies for the same resource in the Lotka-Volterra reaction-diffusion-advection system in 2019. Md. Kamrujjaman [10] studied the impact of diffusion strategies on the outcome of competition between two populations while the species are distributed according to their respective carrying capacities in competition-diffusion systems. However, in the competition-diffusion-advection systems, the study of different species with different distribution functions will be more complex. Tang and Chen [11] and Xu et al. [12] studied the population dynamics of competition between two organisms from the perspective of river ecology in 2020. One interesting feature of their system was that the boundary conditions at the upstream end and downstream end can represent the net loss of individuals. In some cases, both organisms leave the site of competition, neither coexisting nor becoming extinct. Such an environment is important enough to demonstrate how organisms change their density and survival time in competition (see [13]). In 2021, Ma and Guo [14] described the feature of the coincidence of bifurcating coexistence steady-state solution branches and the effect of advection on the stability of the bifurcating solution. However, it is worthwhile to point out that all the aforementioned works focus on the global dynamic behaviors of competition-diffusion systems (see [10,15,16]) or advection systems (see [17,18]), in which the diffusion rates and spatial carrying capacity are changed, or the periodic habitat of advection systems is studied, or the upstream and downstream boundary conditions are changed.
Motivated by the effort of the aforementioned studies, we investigate the problem of the global directed dynamic behaviors of a Lotka-Volterra advection system between two organisms in heterogeneous environments, where two organisms are competing for different fundamental resources, their advection and diffusion strategies follow the dispersal towards a positive distribution, and the functions of inter-specific competition ability are variable.
Hence, we discuss the following global dynamics of the advection system:
where and are the population densities of biological organisms, location and time , which are supposed to be nonnegative; correspond to the dispersal rates of two competing organisms U and V, respectively. ∇ is the gradient operator. correspond to the advection rates of two competing organisms U and V, and are the nonconstant functions and represent the advective direction. The intrinsic growth rates of the two competing organisms are bounded functions and , respectively, two positive distributions are . show the inter-specific competition ability. The habitat is a bounded smooth domain in , ; n denotes the outward unit normal vector on the boundary . For the sake of simplicity, we can suppose the initial data and not identically zero. The system (1) satisfies no-flux boundary conditions.
When , the system (1) becomes the advection system studied by Zhou and Xiao [19]:
where and are positive constants. is the usual Laplace operator. If , readers can take a look at the relevant literature [20] and for the case , please see the references [21,22,23,24,25].
If , the system (2) becomes a diffusion model (see [2,3,5,26]):
According to the research of the above models, the purpose of our paper is to deal with a more broader model (1) in a high spatial dimensions. In this system, we consider that the diffusion and advection strategies follow the dispersal towards a positive distribution, growth rates and competitiveness of the two organisms are different. Thus, we have the following basic assumptions in this paper.
- , , and are positive constants;
- ;
- , in ;
- is nonconstant, and , are also nonconstant.
Conditions ensure that the distribution of resources is heterogeneous for two species and the positivity is imposed here to guarantee the existence of two semi-trivial steady states for later discussion convenience. Under the conditions of , we show a complete classification of the global dynamics of the system (1). The rest of this paper is arranged as follows. In Section 2, we mainly do some preparatory work. Some related properties of the system (1) are deduced from the properties of a single organisms model (4). Besides, some lemmas are proved. In Section 3, we investigate our main results. By using principal eigenvalue theory, we obtain the linear stability of coexisting steady states (see Theorem 2). Then, the most important thing is that in virtue of the monotone dynamical system theory (see [4]), we show the classification of global dynamic behaviors (see Theorem 3). A discussion on the main results and problems that deserve future investigation is presented in Section 4.
2. Preliminaries
In order to describe our main results, we show a competition-diffusion-advection system for a single organisms as follows:
where and , is bounded. According to the relevant description in [27] and the case that , there is a unique positive steady state in the system (4). If we apply this result to the system (1) and the conditions , there are two semi-trivial steady states and respectively.
Lemma 1.
Assume that and , is bounded. The elliptic boundary value Problem:
has a unique positive solution denoted by θ.
Proof.
It is known in [27] that the problem (5) admits a solution and the solution is positive, denoted by , owning to the positivity of . Next, assume that , are any two positive solutions of (5) and . It is not difficult to see that
Then
We deduce
Therefore, . □
To give a complete classification of the global dynamic system (1), we define
Based on the approach in [2], we define
We first recall the well-known Krein-Rutman Theorem:
Theorem 1
(Krein-Rutman Theorem [28]). Let X be a Banach space, a total cone and a compact linear operator that is positive (i.e., ) with positive spectral radius . Then is an eigenvalue with an eigenvector . Moreover, is an eigenvalue of with an eigenvector .
In order to better describe the linear stability of semi-trivial steady states, we give the definition of elliptic eigenvalue problem:
where and Let
Since A is uniformly strongly elliptic operator, we declare that the operator A satisfies the conditions in Theorem 1. An eigenvalue of the problem (8) is called a principal eigenvalue if and for any eigenvalue with , we have . Hence, the problem (8) has a principal eigenvalue, denoted by , and its corresponding eigenfuntion in . The principal eigenvalue is expressed as
Next, we give a useful lemma related to eigenvalue comparison results, which is used for Lemma 3 and Theorem 3.
Lemma 2
([5]). If within Ω, then and the equality holds if and only if in Ω.
According to the description of theory of monotone semi-flow in the literature [6], let X denote the standard Banach space consisting of all continuous functions from to , i.e., , and be the set of all non-negative continuous functions from to . Define as the usual cone for the study of competitive systems with nonempty interior. Then we define the notion of linear stability of a given steady state . Linearizing the steady state problem of (1) at , we obtain
Similar to the scalar problem (8), we can define the principal eigenvalue for the system (10), that is, an eigenvalue of the problem (10) is called a principal eigenvalue if and for any eigenvalue with , we have . Based on the approach in [6], by using Theorem 1, the problem (10) has a principal eigenvalue . In fact, we can select the corresponding eigenfunction , which satisfies in . Here, for the convenience of readers to better understand the problem (10), we provide a simple illustration. Let us do this simple transformation
then the problem (10) can be changed to
which is a linear cooperative elliptic system. Suppose now L is the elliptic operator, let
According to [28,29], the problem (11) has coefficients and is strictly uniformly elliptic in the bounded domain which has boundary. Let K be the positive cone in consisting of nonnegative functions. For any , then we can deduce that defined by is a positive compact linear operator. By applying Theorem 1 for positive compact linear operators and the Neumann type boundary condition, the problem (11) admits a principal eigenvalue , and the corresponding eigenfunction can be chosen to satisfy and in . Notice that is the solution of the problem (11). Moreover, since the off-diagonal elements and are strictly positive in , it can be further concluded that is simple and it is the unique eigenvalue corresponding to a pair of strictly positive eigenfunctions, i.e., and in . In fact, we have and in due to Hopf boundary lemma, which in turn allows us to choose in . See [30] using semi-group theory and [31] using maximum principle, [1,6] for detailed explanation. For the principal eigenvalue theory of general linear cooperative elliptic systems, we refer the interested readers to [29]. If is an eigenvalue of (10) and the boundary condition is Neumann type, then in the coexistence case.
Based on [26], (Corollary 2.10), the following lemma is about the linear stability of and .
Lemma 3.
The linear stability of , and in the system (1) are determined by the sign of , and
Proof.
For the linear stability of , when in (10), we have
Let be an principal eigenvalue of (12) with the eigenfunction . We get
If , then belonging to an eigenvalue of the second equation in (12), is real and the inequality holds. Perhaps, if , then and is an eigenvalue of the first equation, we get -4.6cm0cm
Due to the fact that is real and satisfies . It follows
If now , letting be the first eigenfunction corresponding to , then is an eigenvalue of (12) with the eigenfunction , which deduces .
Suppose that . Let be the first eigenfunction corresponding to , then is an eigenvalue of (12) with the eigenfunction , that means . Here satisfies
The existence of is inferred from
So our claim is right. Owing to (6) and (9), it is inferred that . Hence, according to Lemma 2, we gain
then has the same sign as the first eigenvalue . Applying the definition of and linear stability, we deduce that the linear stability of is determined by the sign of .
Through completely similar arguments, we demonstrate that the stability of and , is determined by respectively. □
Remark 1.
From the variational characteristics of the first eigenvalue, we can see that is linearly unstable for any .
Therefore, we give equivalent descriptions of (7) below:
The neutrally stable case is defined as follows
By the definition, it is easy to see =.
In the following, “” is used to mean that the steady state is globally asymptotically stable among all non-negative and not identically zero initial conditions.
Lemma 4
([5]). For any , assume that hold and every coexistence steady state of the system (1), if it exists, is asymptotically stable. Then one of the following alternatives holds:
- (i)
- There exists a unique coexistence steady state of (1) that is .
- (ii)
- The system (1) has no coexistence steady state and either one of or is , while the other is unstable.
3. Main Results
In this section, we present the results which are related to the co-existence steady state and the classification of global dynamic behaviors of the system (1).
Theorem 2.
Suppose that hold. For any , then every co-existence steady state of the system if exists, is linearly stable, i.e., .
Theorem 3.
Suppose that hold. Then we have the mutually disjoint decomposition of Γ:
Moreover, the following statements hold for the system :
- For all , is ;
- For all , is ;
- For all , the system has a unique coexistence steady state that is ;
- For all , in and the system has a compact global attractor consisting of a continuum of steady statesconnecting with two semi-trivial steady states.
3.1. Co-Existence Steady State
In order to prove Theorem 2, we assume that is the co-existence steady state of the following system (1):
Similar to the problem (10), then we get the linear eigenvalue model by linearize system (1) at ,
According to the problem (8) and using Theorem 1, we can deduce that the problem (18) has a principal eigenvalue . Moreover, we can choose the corresponding eigenfunction , it satisfies in .
Now, we are ready to discuss Theorem 2.
Proof of Theorem 2.
Obviously, as long as we can obtain when . Multiplying the first equation in and by and , respectively, and subtracting the obtained equations, we obtain
In the similar way, it can be derived from the second equation in and that
Furthermore, multiplying by , then we integrate over and deduce (for simplicity, we replace with ∫)
By using the similarly method for the Equation (20), we get
We now simplify the formulas and Then we find
and
Next, we have
By the similar method, we deduce that
and
Thus
By a similar method, one obtains
Replace (21) and (22) with (26) and (27), respectively. Multiplying (22) by and subtracting it from (21), we can obtain
where we use the facts and in . So, .
Next, we will prove that can not happen. According to (28), we infer that if and only if
which means that
i.e.,
Then, one obtains
In addition, by applying (30) to (17), and the uniqueness of the positive steady state of the system (4), it can be concluded that
Noting that , we deduce
Based on (31), one can easily check
and
According to the assumption , we get . Therefore, . □
3.2. Classification of the Global Dynamics
In this subsection, we are ready to discuss the complete global dynamic behaviors of the system (1).
Proof of Theorem 3.
According to the proof procedure (see [2,5]), this proof can be divided into two steps.
Step 1. On the proof of the disjoint decomposition in (16).
Obviously, we can get the decomposition in (16). According to the relevant conclusion and definitions, we only need to prove
By Lemma 3, the linear stability of , can be determined by the sign of , respectively. For the sake of convenience of in writing, let
According to the properties of the variational characterization and (6), we obtain
and
Since , combining with (33) and (34) together, we have
where all the inequalities become equalities if and only if
It follows from (35) that the conclusion (32) holds.
Step 2. On the proof of the statements .
Firstly, we will prove the statements hold. In consideration of (16) in Theorem 3 and (35) in step 1, we see that for any there are five possibilities as follows:
- , i.e., is linearly stable, is linearly unstable;
- , i.e., is linearly unstable, is linearly stable;
- , i.e., both and are linearly unstable;
- , i.e., is neutrally stable, is linearly unstable;
- , i.e., is neutrally stable, is linearly unstable.
By Lemma 4, we immediately deduce the following conclusion:
and are based on the assumptions and , respectively, and there is a unique co-existence steady state under the condition .
We now claim that there is no coexistence steady state under the condition or the condition . Then we can infer that and are also based on the assumptions and , respectively, from Lemma 4.
We only need to verify the above statement for the case (). Indeed, if the system (1) has a co-existence steady state for some and satisfies
We have
Define the operator
with
From (36), and Theorem 2, it yields that
Applying implicit function theorem, one gets is closed to . We have a positive solution to the equation . Let us choose , which implies (the solution corresponding to is denoted by )
Let us see the following auxiliary problem
which has the same semi-trivial steady states and . From (37), (38) and Lemma 2, it then follows that
According to the case , is also in the system (39) which contradicts with the existence of . Therefore, there is no coexistence steady state under the condition . Similarly, we can get the conclusion that there is also no coexistence steady state under the condition . The above descriptions of the cases represent the expected results described in the statements .
Secondly, we prove the statement . We will show
It makes the same description of , which means the expected result in the statement .
Let , then
Based on the proof of Theorem 2, we get
which implies . When (42) holds, the last three inequalities in (35) become equalities, we have
which shows . Hence, the equality (41) is confirmed.
Let and be the corresponding coexistence steady state of (1). We claim that constant. Let be a principal eigenvalue for . Moreover, we choose the corresponding eigenfunction , which satisfies in and . In order to prove it, it is enough to show that (29) holds. Suppose that (29) is not true. Then (28) means . Similar to the proof of the case , we get (29) holds, i.e., constant. This yields that
Therefore, we conclude that for any , the set of equilibria of (1) is
where is a repeller by Remark 1. Then each solution of (1) converges to a single equilibrium . □
4. Discussion
In this paper, by using principal eigenvalue theory and monotone dynamical system theory, we mainly analyzed the global directed dynamic behaviors of a Lotka-Volterra competition-diffusion-advection system between two organisms in heterogeneous environments. The two organisms compete for different fundamental resources, their advection and diffusion strategies follow a positive diffusion distribution, the functions of inter-specific competition ability are variable. Our work can be seen as a further development of Wang [5] for the competition-diffusion system, where we bring new ingredients in the arguments to overcome the difficulty caused by the involvement of advection.
In the future, exploring the global directed dynamic behaviors under the condition of cross-diffusion may be an interesting research point. We leave this challenge to future investigations.
Author Contributions
All authors contributed equally and significantly in writing this article. Conceptualization, L.C.; Formal analysis, Y.Z.; Funding acquisition, L.C.; Writing—original draft, S.L. All authors have read and agreed to the published version of the manuscript.
Funding
This research was funded by Shandong Provincial Natural Science Foundation under grant ZR2020MA006 and the Introduction and Cultivation Project of Young and Innovative Talents in Universities of Shandong Province.
Institutional Review Board Statement
Not applicable.
Informed Consent Statement
Not applicable.
Data Availability Statement
Not applicable.
Acknowledgments
We would like to express our thanks to the anonymous referees and the editor for their constructive comments and suggestions, which greatly improved this article
Conflicts of Interest
The authors declare no conflict of interest.
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