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Screening and validating the core biomarkers in patients with pancreatic ductal adenocarcinoma

  • Pancreatic ductal adenocarcinoma (PAAD) is one of the most common malignant tumors in digestive system. To find the new therapeutic targets and explore potential mechanisms underlying PAAD, the bioinformatics has been performed in our study. The PAAD gene expression profile GSE28735 was chosen to analyze the differentially expressed genes (DEGs) between PAAD carcinoma tissues and normal adjacent tissues from 45 patients with PAAD. Gene ontology (GO) and Kyoto Encyclopedia of Genes and Genomes (KEGG) pathway analysis were performed using Database for Annotation, Visualization and Integrated Discovery (DAVID). Moreover, a protein-protein interaction (PPI) network was also constructed to help us screen the top 20 hub genes in this profile and demonstrated the underlying interactions among them. The Gene Expression Profiling Interactive Analysis (GEPIA) was further performed in order to valid the mRNA levels of top5 up-regulated and top5 down-regualted DEGs, apart from exploring their association with survival rate as well as tumor stage. Finally, Q-PCR was further employed to valid the top5 up-regulated and top5 down-regulated genes in patients with PAAD. In our study, there were a total of 444 DEGs captured (271 up-regulated genes and 173 down-regulated genes). Among these DEGs, the top5 up-regulated genes were CEACAM5, SLC6A14, LAMC2, GALNT5 and TSPAN1 while the top5 down-regulated genes were GP2, CTRC, IAPP, PNLIPRP2 and PNLIPRP1. GO analysis disclosed that the DEGs were predominantly enriched in cell adhesion, lipid metabolism, integrin binding, proteolysis and calcium ion binding. KEGG analysis disclosed that the enriched pathway included pancreatic secretion, protein digestion and absorption, fat digestion and absorption, ECM-receptor interaction, focal adhesion and PI3K-Akt signaling pathway. Survival analysis unveiled that the high expression levels of SLC6A14, GALNT5 and TSPAN1 may correlate with the poor prognosis while high expression levels of IAPP may contribute to a better prognosis in patients with PAAD. Additionally, the levels of CEACAM5, SLC6A14, LAMC2 and GALNT5 were also associated with tumor stage. Furthermore, according to the connectivity degree of these DEGs, we selected the top20 hub genes, namely ALB, FN1, EGF, MMP9, COL1A1, COL3A1, FBN1, CXCL12, POSTIN, BGN, VCAN, THBS2, KRT19, MET, MMP14, COL5A2, GCG, MUC1, MMP1 and CPB1, which were expected to be promising therapeutic targets in PAAD. Collectively, our bioinformatics analysis showed that DEGs and hub genes may be defined as new biomarkers for diagnosis and for guiding the therapeutic strategies of PAAD.

    Citation: Yan Li, Yuzhang Zhu, Guiping Dai, Dongjuan Wu, Zhenzhen Gao, Lei Zhang, Yaohua Fan. Screening and validating the core biomarkers in patients with pancreatic ductal adenocarcinoma[J]. Mathematical Biosciences and Engineering, 2020, 17(1): 910-927. doi: 10.3934/mbe.2020048

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  • Pancreatic ductal adenocarcinoma (PAAD) is one of the most common malignant tumors in digestive system. To find the new therapeutic targets and explore potential mechanisms underlying PAAD, the bioinformatics has been performed in our study. The PAAD gene expression profile GSE28735 was chosen to analyze the differentially expressed genes (DEGs) between PAAD carcinoma tissues and normal adjacent tissues from 45 patients with PAAD. Gene ontology (GO) and Kyoto Encyclopedia of Genes and Genomes (KEGG) pathway analysis were performed using Database for Annotation, Visualization and Integrated Discovery (DAVID). Moreover, a protein-protein interaction (PPI) network was also constructed to help us screen the top 20 hub genes in this profile and demonstrated the underlying interactions among them. The Gene Expression Profiling Interactive Analysis (GEPIA) was further performed in order to valid the mRNA levels of top5 up-regulated and top5 down-regualted DEGs, apart from exploring their association with survival rate as well as tumor stage. Finally, Q-PCR was further employed to valid the top5 up-regulated and top5 down-regulated genes in patients with PAAD. In our study, there were a total of 444 DEGs captured (271 up-regulated genes and 173 down-regulated genes). Among these DEGs, the top5 up-regulated genes were CEACAM5, SLC6A14, LAMC2, GALNT5 and TSPAN1 while the top5 down-regulated genes were GP2, CTRC, IAPP, PNLIPRP2 and PNLIPRP1. GO analysis disclosed that the DEGs were predominantly enriched in cell adhesion, lipid metabolism, integrin binding, proteolysis and calcium ion binding. KEGG analysis disclosed that the enriched pathway included pancreatic secretion, protein digestion and absorption, fat digestion and absorption, ECM-receptor interaction, focal adhesion and PI3K-Akt signaling pathway. Survival analysis unveiled that the high expression levels of SLC6A14, GALNT5 and TSPAN1 may correlate with the poor prognosis while high expression levels of IAPP may contribute to a better prognosis in patients with PAAD. Additionally, the levels of CEACAM5, SLC6A14, LAMC2 and GALNT5 were also associated with tumor stage. Furthermore, according to the connectivity degree of these DEGs, we selected the top20 hub genes, namely ALB, FN1, EGF, MMP9, COL1A1, COL3A1, FBN1, CXCL12, POSTIN, BGN, VCAN, THBS2, KRT19, MET, MMP14, COL5A2, GCG, MUC1, MMP1 and CPB1, which were expected to be promising therapeutic targets in PAAD. Collectively, our bioinformatics analysis showed that DEGs and hub genes may be defined as new biomarkers for diagnosis and for guiding the therapeutic strategies of PAAD.


    Dispersal of organisms is a topic of central interest in ecology and evolutionary biology. Its effects on the size, stability, and interactions of populations, as well as biological invasions and the geographical distribution of populations have attracted considerable studies. Investigation on dispersal strategies which are evolutionarily stable has been the fundamental research goal for theoretical ecologists [7,26]. The relationship between diffusion rates, spatial heterogeneity, and coupling from competition of species is the target of several recent works. To tackle these problems, continuous diffusion models expressed by reaction-diffusion systems have been considered in [1,2,3,6,9,13,14,19,20,23,25]. On the other hand, discrete diffusion models represented by systems of ODEs have been investigated in [4,5,11,12,24,31].

    Concerning the interaction between diffusion rates and the heterogeneity of the environment and mutant invasion, the following competitive Lotka-Volterra model was investigated in [18,19]:

    ut=μΔu+u[α(x)uv],vt=μΔv+v[β(x)uv], (1.1)

    under homogeneous Neumann boundary condition, where μ is the diffusion rate and functions α(x) and β(x) express the spatially dependent intrinsic growth rates or reproductive rates of u- and v-species, respectively. Therein, to study the effect of spatially heterogeneous growth rates on the competitive dynamics, the difference between intrinsic growth rates of two species was set as

    α(x)=β(x)+τg(x),

    where g(x) is a function describing resource difference between two species from the viewpoint of spatial heterogeneity, and τ>0 measures the magnitude of the difference. The case g(x)>0 on the considered domain was studied in [18], whereas the situation that g(x) changes sign was investigated in [19]. The assumption in [19],

    Ωg(x)dx>0, (1.2)

    means that the mutant u-species has better average reproductive rate than v-species, and thus the total population of u-species has higher growth rate than that of v-species when two populations are identical in the whole space Ω. However, under such a circumstance, u-species possibly fails to invade when rare for certain level of diffusion rate. Mathematically, stability of semitrivial solutions (˜u,0) and (0,˜v), which depend on the magnitudes of μ and τ, was analyzed in [19]. The stability may switch according to the varying diffusion rate μ. In particular, by measuring the level of mutation with the value of τ, theoretical analysis for the cases of tiny and large mutation was established therein. In the former case (0<τ1), multiple switches of global convergence to different equilibria was derived and the relationship between the bifurcation value of the diffusion rate and the value of τ was also established, while in the latter case (τ1), only once switch of global convergence was observed.

    The influence from magnitudes of diffusion rates on the competition outcome has been another topic of interest. It has been shown in [9] that the slower diffuser always prevails if the two species interact identically with the environment, see also [12,20,23]. To focus on the effect of diffusion rates, the birth rates for all competing species were set equal to the carrying capacity of the environment, see [2,5,6,13,15].

    Models for competitive species with dispersal expressed by discrete diffusion are also very appealing. Indeed, organisms are distributed in space, often in patches of habitat scattered over a landscape and region, and the distribution is determined by the pattern of movement between these patches. More specifically, it is interesting to see how possible interaction outcome, which can be competitive exclusion and coexistence, depends on the diffusion rates and the birth rates.

    As early as in 1934, Gause [10] formulated the competitive exclusion law which in particular states that the species with a larger birth rate will outcompete the other one, if the other properties are the same. Concerning these issues, Gourley and Kuang [11] then asked how does diffusion affect the competition outcomes of two competing species that are identical in all respects other than their strategies on how they spatially distribute their birth rates. They studied the following ODE system as a model for two neutrally competing species on two patches of habitat:

    {du1dt=u1(α1u1v1)+d(u2u1)du2dt=u2(α2u2v2)+d(u1u2)dv1dt=v1(β1u1v1)+d(v2v1)dv2dt=v2(β2u2v2)+d(v1v2) (1.3)

    where ui (resp., vi) is the population density of species-u (resp., -v) in patch i, i=1,2; the linear birth rates α1,α2,β1,β2 are positive parameters, and there is a diffusion between the two patches with same diffusivity (dispersal rate) d for both species. The two species differ only in their birth rates. Let (ˉu1,ˉu2,0,0) denote the semitrivial equilibrium with extinct v-species. The following conjectures on the global dynamics of system (1.3) were posed in [11]:

    Conjecture 1. Assume that in system (1.3), β1σ=α1<β1<β2<α2=β2+σ with 0<σ<β1, and d is sufficiently large. If u1(0)+u2(0)>0, then

    limt(u1(t),u2(t),v1(t),v2(t))=(ˉu1,ˉu2,0,0).

    Conjecture 2. Assume that in system (1.3), β1σ=α1<β1<β2<α2=β2+σ with 0<σ<β1, and d is small enough so that (1.3) has a positive steady state e. If u1(0)+u2(0)>0 and v1(0)+v2(0)>0, then

    limt(u1(t),u2(t),v1(t),v2(t))=e.

    These conjectures, if true, suggest that the species that can concentrate its birth in a single patch wins, if the diffusion rate is larger than a critical value. That is, the winning strategy is to focus as much birth in a single patch as possible. In [24], the following global dynamics and bifurcation were established, which include confirmation of Conjectures 1 and 2:

    Theorem 1.1. Suppose that the following condition holds in system (1.3),

    (C):0<α1=β1σ1<β1<β2<α2=β2+σ2with0<σ1σ2.

    Then there is a constant ˜d>0 which can be expressed or estimated by the birth rates, so that if d˜d, (ˉu1,ˉu2,0,0) is globally asymptotically stable among the initial data in R4+ satisfying u1(0)+u2(0)>0; if d<˜d, (1.3) has a unique positive steady state (u1,u2,v1,v2) which is globally asymptotically stable among the initial data in R4+ satisfying u1(0)+u2(0)>0 and v1(0)+v2(0)>0.

    Note that α1+α2 and β1+β2 measure the average birth rates of species u and v, respectively. The condition of Theorem 1.1 means α1<β1<β2<α2 and β1+β2α1+α2, and indicates that the birth rate of u-species is larger than that of v-species in the second patch, and less than that of v-species in the first patch, whereas the average birth rate of u-species is larger than or equal to that of v-species. For the situation with identical average birth rate: α1+α2=β1+β2, i.e., the case in these conjectures, Theorem 1.1 implicates that the two species coexist in a slow diffusion environment, whereas in a fast diffusion environment, the species that can concentrate its birth in a single patch drives the other species into extinction. Convincingly, the same scenario prevails when u has further competitive advantage that its average birth rate is larger than v-species: α1+α2>β1+β2. Along with such finding is that the semitrivial equilibrium (0,0,ˉv1,ˉv2) is always unstable for any diffusion rate d, as was stated in Proposition 3.11 of [24]. It becomes very interesting to see what happens when α1+α2<β1+β2, i.e., v-species has larger average birth rate.

    In this paper we will examine such interesting situation, i.e., system (1.3) under condition

    (C):0<α1=β1σ1<β1<β2<α2=β2+σ2with0<σ2<σ1.

    This condition means α1<β1<β2<α2, and α1+α2<β1+β2, due to (β1+β2)(α1+α2)=σ1σ2>0. That is, the birth rate α2 of u-species in the second patch is the biggest among all species and patches, but the average birth rate of v-species is larger than that of u-species; one may also regard this as that v-species has more total resources than u-species. Then we ask how the magnitude of the dispersal rate d is related to the species persistence or extinction. With the framework of monotone dynamics, we shall target the global dynamics of system (1.3) and the bifurcation with respect to d, under condition (C).

    It turns out that the dynamical scenarios are richer than the case under condition (C). In particular, equilibrium (0,0,ˉv1,ˉv2) switches from being unstable to stable, as d increases. On the other hand, there are up to two stability changes for equilibrium (ˉu1,ˉu2,0,0), as d increases. That is, the property described as monotone relation between the stability of (ˉu1,ˉu2,0,0) and the diffusion rate d no longer holds, cf. [19]. The main results will be summarized in Theorem 4.2. There are two dynamical scenarios (see Figure 1): (ⅰ) Under σ2β2<σ1β1, there exists an d3>0, so that the positive steady state (u1,u2,v1,v2) is globally attractive for d<d3 and the semitrivial equilibrium (0,0,ˉv1,ˉv2) becomes globally attractive for dd3. (ⅱ) Under σ2β2>σ1β1, there exist d1,d2,d3 with 0<d1<d2<d3, so that (u1,u2,v1,v2) is globally attractive for d<d1 or d2<d<d3, (ˉu1,ˉu2,0,0) is globally attractive for d1dd2, and (0,0,ˉv1,ˉv2) becomes globally attractive for dd3. In addition, d1,d2,d3 can be estimated in terms of the system parameters. Our analytical work on the model strongly suggests that, in a fast diffusion (large dispersal) environment, a species will prevail if its average birth rate is larger than the other competing species; in a slow diffusion (small dispersal) environment, the two species can coexist or one species that has the greatest birth rate among both species and patches, even with smaller average birth rate, will be able to persist and drive the other species to extinction.

    Figure 1.  Two dynamical scenarios for system (1.3): the main results, stated in Theorems 2.1 and 4.2.

    We note that α1+α2>β1+β2 in system (1.3) is analogous to condition (1.2) in PDE system (1.1). The present study, with α1+α2<β1+β2, can be compared to the results in [19] with u and v reversed. Systems with two competing species over two patches with different dispersal rates and more general competition coupling have been considered in [21,29,30]. While the effect of competition was studied in [29], herein we aim at investigating the influence of both dispersal rate and birth rates on the population dynamics and assume the same ability of competition for two species in (1.3). Predator-prey dynamics on two-patch environments were investigated in [8,16,22].

    This presentation is organized as follows. In Section 2, we characterize the existence of positive equilibrium for system (1.3). In Section 3, we analyze the stability of the semitrivial equilibria. In Section 4, we discuss the existence of positive steady state representing coexistence of two species and extinction of one species, depending on the magnitude of dispersal rate. Four numerical examples illustrating the present theory are given in Section 5. We summarize our results with some discussions in Sections 6. For reader's convenience, we review in Appendix Ⅰ the monotone dynamics theory which is to be applied to obtain our results. Some qualitative properties of the semitrivial equilibria for system (1.3) reported in [24] are recalled in Appendix Ⅱ.

    In this section, we characterize the conditions under which the positive equilibrium (u1,u2,v1,v2) of system (1.3) exists. There are five parameters α1,α2,β1,β2,d in system (1.3), which generate a complication of analysis for such existence. We first derive the following magnitude relationships which are required in the main result, Theorem 2.1, of this section.

    Lemma 2.1. The following parameter relationships hold under condition (C).

    (ⅰ) 1σ1σ2<β1β2σ2β22σ1β21 if and only if (σ2β22σ1β21)(σ1β1σ2β2)>0.

    (ⅱ) 0<α1α2σ2α22σ1α21<β1β2σ2β22σ1β21, provided σ2β22σ1β21>0.

    (ⅲ) σ1σ1σ2<β2σ1+σ2, provided σ1β1>σ2β2 and β2β1σ1+σ2.

    Proof. Recall that σ1>σ2 in condition (C).

    (ⅰ) We compute

    β1β2σ2β22σ1β211σ1σ2=β1β2(σ1σ2)(σ2β22σ1β21)(σ2β22σ1β21)(σ1σ2)=(β1+β2)(σ1β1σ2β2)(σ2β22σ1β21)(σ1σ2).

    Thus, β1β2σ2β22σ1β211σ1σ2>0 if and only if (σ2β22σ1β21)(σ1β1σ2β2)>0.

    (ⅱ) Suppose σ2β22σ1β21>0. Then

    σ2α22σ1α21>σ2β22σ1α21>σ2β22σ1β21>0.

    The assertion follows from

    α1α2σ2α22σ1α21β1β2σ2β22σ1β21=(σ2α2β2+σ1α1β1)(α1β2α2β1)(σ2α22σ1α21)(σ2β22σ1β21)<0,

    due to α1β2α2β1=α1β2(β2+σ2)(α1+σ1)<0.

    (ⅲ) If σ1β1>σ2β2, then

    β2σ1+σ2σ1σ1σ2=σ1β2σ2β2σ1(σ1+σ2)σ21σ22>σ1β2σ1β1σ1(σ1+σ2)σ21σ22=σ1[(β2β1)(σ1+σ2)]σ21σ220,

    provided β2β1σ1+σ2. The assertion thus follows.

    The following parameter condition is to be used throughout the discussions:

    Condition(P):1σ1+σ2<σ1β1σ2β22σ1β21.

    Certainly condition (P) holds only if σ2β22σ1β21>0. And a direct computation shows that condition (P) is equivalent to σ2β22σ1β21>0 with

    σ2β22<σ1β1(β1+σ1+σ2). (2.1)

    Accordingly, if condition (P) holds, Lemma 2.1(ⅰ) can be recast as

    1σ1σ2<β1β2σ2β22σ1β21σ2β2<σ1β1;

    for convenience of later use, we put this relationship as

    σ1σ2σ1σ2<σ1σ2β1β2σ2β22σ1β21σ2β2<σ1β1. (2.2)

    The condition of Lemma 2.1(ⅲ): σ1β1>σ2β2 and β2β1σ1+σ2 implies

    σ1σ1σ2<β2σ1+σ2.

    Then, by combining condition (P), we obtain

    σ1σ2σ1σ2<σ2β2σ1+σ2<σ1σ2β1β2σ2β22σ1β21. (2.3)

    On the other hand, combining σ1σ2σ1σ2>σ1σ2β1β2σ2β22σ1β21, i.e. σ2β2>σ1β1 by (2.2), with condition (P) yields

    σ2β2σ1+σ2<σ1σ2β1β2σ2β22σ1β21<σ1σ2σ1σ2. (2.4)

    Therefore, by imposing condition (P) additionally, the following relationships can be concluded.

    Lemma 2.2. Assume that conditions (C) and (P) hold.

    (ⅰ) If σ2β2<σ1β1 and β2β1σ1+σ2, then

    σ1σ2σ1σ2<σ2β2σ1+σ2<σ1σ2β1β2σ2β22σ1β21.

    (ⅱ) If σ2β2>σ1β1, then

    σ2β2σ1+σ2<σ1σ2β1β2σ2β22σ1β21<σ1σ2σ1σ2.

    It is obvious that the terms in the inequalities in Lemma 2.2 can be simplified. But it is convenient to keep these forms.

    Remark 1. In Lemma 2.2, with (2.1), the condition in (ⅱ) : σ2β2>σ1β1 leads to σ1β1β2<σ2β22<σ1β1(β1+σ1+σ2), and thus β2β1<σ1+σ2, which is contrary to the condition β2β1σ1+σ2 in (ⅱ). That is, the condition in (ⅰ) and the condition in (ⅱ) are opposite cases under assumption (P). In addition, the condition in Lemma 2.2(ⅰ): σ2β2<σ1β1 and β2β1σ1+σ2 further indicates

    σ2β22<σ1β1(β1+σ1+σ2)σ1β1β2, (2.5)

    via (2.1).

    We characterize the existence of positive equilibrium for system (1.3) in the following theorem.

    Theorem 2.1. Consider system (1.3) under conditions (C) and (P).

    (ⅰ) Under σ2β2<σ1β1 and β2β1σ1+σ2, there exists an d3>0 so that the system has a unique positive equilibrium (u1,u2,v1,v2) if and only if 0<d<d3.

    (ⅱ) Under σ2β2>σ1β1, there exist d1,d2,d3>0, with d1<d2<d3, so that the system has a unique positive equilibrium (u1,u2,v1,v2) if and only if 0<d<d1 or d2<d<d3.

    In addition,

    σ1σ2α1α2σ2α22σ1α21<d1<σ1σ2β1β2σ2β22σ1β21σ1σ2σ1σ2<d2<σ1σ1σ2+σ1σ2σ21σ22(α2α1)σ1σ2σ1σ2<d3<σ1σ1σ2(α2α1).

    Proof. System (1.3) has a positive equilibrium (u1,u2,v1,v2) if and only if

    (α1u1v1)+d(u2u11)=0,(α2u2v2)+d(u1u21)=0,(β1v1u1)+d(v2v11)=0,(β2v2u2)+d(v1v21)=0, (2.6)

    are satisfied for u1,u2,v1,v2>0. Let (u1,u2,v1,v2) be a solution of (2.6) and denote

    a:=u2u1,b:=v2v1. (2.7)

    Combining each pair of equations in (2.6), we obtain

    σ1+d(ab)=0,σ2+d(1a1b)=0. (2.8)

    This yields

    ab=σ1σ2=:k, (2.9)

    and k>1, as 0<σ2<σ1. Substituting b=k/a and a=k/b into (2.8) respectively leads to

    a=σ1+σ21+4kd22d,b=σ1+σ21+4kd22d. (2.10)

    We thus express a,b in terms of system parameters, and it can be computed that b2<k<a2 and a>1. We substitute (2.7) into (2.6) and obtain

    (α1u1v1)+d(a1)=0,a(α2au1bv1)+d(1a)=0, (2.11)
    (β1v1u1)+d(b1)=0,b(β2bv1au1)+d(1b)=0. (2.12)

    Solving the two equations in (2.11), we have

    {u1=1a2k[(aα2ad+d)k(α1+add)]v1=(α1+add)u1. (2.13)

    On the other hand, solving the two equations in (2.12), we obtain

    {u1=1kb2[(bβ2bd+d)b2(β1+bdd)]v1=(β1+bdd)u1. (2.14)

    In fact, (2.13) and (2.14) are equivalent, as it can be seen by (2.8) that α1+add=β1+bdd and

    1a2k[(aα2ad+d)k(α1+add)]=1kb2[(bβ2bd+d)b2(β1+bdd)].

    Herein, α1+add>0 since a>1. From (2.13) and (2.14), we obtain

    v1=1a2k[a2(α1+add)(aα2ad+d)]=1kb2[k(β1+bdd)(bβ2bd+d)].

    Observe that u1,v1>0 imply u2,v2>0, due to (2.7) and a,b>0. Hence, system (1.3) has a unique positive equilibrium if and only if

    u1=1a2k[(aα2ad+d)k(α1+add)]=1kb2[(bβ2bd+d)b2(β1+bdd)]>0, (2.15)

    and

    v1=1a2k[a2(α1+add)(aα2ad+d)]=1kb2[k(β1+bdd)(bβ2bd+d)]>0. (2.16)

    To explore the range of d where u1,v1>0, we denote u1(d),v1(d) to express the dependence of u1,v1 on d. Let us discuss the positivity of v1 first. The terms in the brackets of (2.16) can be recast as

    {a2(α1+add)(aα2ad+d)=a2α1aα2+d(a2+1)(a1)k(β1+bdd)(bβ2bd+d)=kβ1bβ2+d(k+1)(b1). (2.17)

    We define two functions to discuss the positivity of v1:

    Fv(d):=a2α1aα2+d(a2+1)(a1),Gv(d):=kβ1bβ2+d(k+1)(b1).

    It follows from (2.16) and (2.17) that

    Fv(d)=a2kkb2Gv(d).

    In addition, v1>0 if and only if Fv(d)>0 if and only if Gv(d)>0, due to b2<k<a2. In the following discussions (a)-(e), we analyze the ranges of d within which Fv(d) and Gv(d) take positive or negative values. For some situations, analyzing Fv is more convenient than Gv, whereas the convenience is reverse in other cases.

    (a) Fv(d)>0 if d<σ1σ2α1α2σ2α22σ1α21: It can be seen that aα1>α2 implies Fv(d)>0, due to a>1. On the other hand, aα1>α2 is actually

    a=σ1+σ21+4kd22d>α2α1,

    which is equivalent to d<σ1σ2α1α2σ2α22σ1α21.

    (b) Fv(d)>0 if d>σ1σ1σ2+σ1σ2σ21σ22(α2α1): If b>1, i.e. d>σ1σ2σ1σ2 by (2.10), then a<k since ab=k. From b2<k<a2, we have 1<b<k<a<k. Then

    Fv(d)=a2α1aα2+d(a2+1)(a1)>kα1kα2+d(k+1)(k1)>0,

    if d>k(k+1)(k1)(α2α1)=σ1σ1σ2+σ1σ2σ21σ22(α2α1). It is clear that

    σ1σ1σ2+σ1σ2σ21σ22(α2α1)=σ1σ1σ2+σ1σ2σ21σ22(β2β1)+σ1σ1σ2+σ1σ2σ1σ2>σ1σ2σ1σ2.

    (c) Gv(d)>0 if σ1σ2σ1σ2<d<σ1σ2β1β2σ2β22σ1β21: Obviously, Gv(d)>0 if kβ1bβ2>0 and b>1, which is

    kβ1β2>b=σ1+σ21+4kd22d>1.

    This is equivalent to

    σ1σ2σ1σ2<d<σ1σ2β1β2σ2β22σ1β21,

    by (2.10). Such value of d exists provided σ1σ2σ1σ2<σ1σ2β1β2σ2β22σ1β21.

    (d) Gv(d)<0 if σ1σ2β1β2σ2β22σ1β21<d<σ1σ2σ1σ2: Gv(d)<0 provided kβ1bβ2<0 and b<1, which is

    kβ1β2<b=σ1+σ21+4kd22d<1,

    and equivalent to σ1σ2β1β2σ2β22σ1β21<d<σ1σ2σ1σ2, provided σ1σ2β1β2σ2β22σ1β21<σ1σ2σ1σ2.

    (e) Gv(d)<0 if d<min{σ1σ2σ1σ2,σ2β2σ1+σ2} and Gv(d)>0 if d>max{σ1σ2σ1σ2,σ2β2σ1+σ2}: From (2.10), we compute

    b=b(d)=σ1bdσ21+4kd2>0, (2.18)

    and Gv(d)=(k+1)(b1)+b(kdβ2+d). It can be seen that Gv(d)<0, provided b<1 and kdβ2+d<0, which are equivalent to d<σ1σ2σ1σ2 and d<σ2β2σ1+σ2. On the contrary, Gv(d)>0, if b>1 and kdβ2+d>0, which are equivalent to d>σ1σ2σ1σ2 and d>σ2β2σ1+σ2.

    For case (ⅰ), we will show that v1(d)>0 for all d>0 if σ2β2<σ1β1 and β2β1σ1+σ2. Recall Lemma 2.2(ⅰ): σ1σ2σ1σ2<σ2β2σ1+σ2<σ1σ2β1β2σ2β22σ1β21. From the above (c), (e), we summarize

    {Gv(d)>0ifσ1σ2σ1σ2<d<σ1σ2β1β2σ2β22σ1β21Gv(d)<0ifd<σ1σ2σ1σ2Gv(d)>0ifd>σ2β2σ1+σ2. (2.19)

    In addition, at d=σ1σ2σ1σ2, i.e., b=1, we have Gv(σ1σ2σ1σ2)=kβ1β2>0, thanks to σ2β2<σ1β1. Therefore, from (2.19), we see that Gv(d)>0 for all d>0, namely, v1(d)>0 for all d>0, under σ2β2<σ1β1, β2β1σ1+σ2, and condition (P).

    For case (ⅱ), if σ2β2>σ1β1, we will show that Gv(d)>0, and hence v1(d)>0, for d in certain range. Recall Lemma 2.2(ⅱ): σ2β2σ1+σ2<σ1σ2β1β2σ2β22σ1β21<σ1σ2σ1σ2. With the above (a), (b), (d), (e), we obtain

    {Fv(d)>0ifd<σ1σ2α1α2σ2α22σ1α21Gv(d)<0ifσ1σ2β1β2σ2β22σ1β21<d<σ1σ2σ1σ2Fv(d)>0ifd>σ1σ1σ2+σ1σ2σ21σ22(α2α1) (2.20)

    and

    {Gv(d)<0ifd<σ2β2σ1+σ2Gv(d)>0ifd>σ1σ2σ1σ2. (2.21)

    Furthermore, if dσ2β2σ1+σ2, i.e. d(k+1)β2, and dσ1σ2σ1σ2, i.e. b1, we have

    Gv(d)=kβ1bβ2d(k+1)(1b)kβ1bβ2β2(1b)=kβ1β2<0,

    by σ2β2>σ1β1. To summarize, Gv(d)<0 if σ2β2σ1+σ2dσ1σ2σ1σ2. Therefore, from (2.20) and (2.21), there exists a unique d1>0 so that Gv(d)>0 if d<d1 and Gv(d1)=0, where

    σ1σ2α1α2σ2α22σ1α21<d1<σ1σ2β1β2σ2β22σ1β21,

    and there exists a unique d2>0 so that Gv(d)>0 if d>d2, where

    σ1σ2σ1σ2<d2<σ1σ1σ2+σ1σ2σ21σ22(α2α1).

    The two cases for the assertion of v1(d)>0 are thus concluded. Now let us discuss the positivity of u1(d). From (2.15), we have

    {(aα2ad+d)k(α1+add)=aα2kα1d(k+1)(a1),(bβ2bd+d)b2(β1+bdd)=b(β2bβ1)+d(b2+1)(1b). (2.22)

    Let

    Fu(d):=aα2kα1d(k+1)(a1),Gu(d):=b(β2bβ1)+d(b2+1)(1b).

    Then

    Fu(d)=a2kkb2Gu(d).

    In addition, u1>0 if and only if Fu(d)>0 if and only if Gu(d)>0, due to b2<k<a2. Let us discuss the signs of Fu(d) and Gu(d) in the following (a')-(d').

    (a') Gu(d)>0 if dσ1σ2σ1σ2: It is clear that b1 implies Gu(d)>0, and b1 is equivalent to dσ1σ2σ1σ2. Thus, Gu(d)>0 if dσ1σ2σ1σ2.

    (b') Fu(d)<0 if d>σ1σ1σ2(α2α1): If b>1, then from 1<b<k<a<k, we have

    Fu(d)=aα2kα1d(k+1)(a1)<aα2aα1d(a+1)(a1)=a(α2α1)d(a21)<k(α2α1)d(k1)<0,

    if d>kk1(α2α1)=σ1σ1σ2(α2α1).

    (c') Case (ⅰ): σ2β2<σ1β1, i.e., σ1σ2σ1σ2<σ1σ2β1β2σ2β22σ1β21 by (2.2). We claim that Fu(d)<0 for dσ2β2σ1+σ2 and Gu(d)<0 for d>σ2β2σ1+σ2. Notably,

    Fu(d)=[β2+σ2d(k+1)]a(k+1)(a1) (2.23)

    and

    Gu(d)=(b2+1)(b1)2bbd(b1)b[d(b2+1)+2bβ1β2], (2.24)

    by direct computations, where a=a(d),b=b(d). For the first term of (2.23), we see that

    β2+σ2d(k+1)σ2>0ifdσ2β2σ1+σ2.

    In addition, from (2.10), we compute

    a=σ1adσ21+4kd2<0. (2.25)

    Hence, in (2.23), we confirm Fu(d)<0 for dσ2β2σ1+σ2, due to a>1 and a<0. Next, we discuss the third term of Gu(d) in (2.24), and claim that

    d(b2+1)+2bβ1β2>0ifd>σ2β2σ1+σ2.

    From (2.10) and b>0 shown in (2.18), a direct computation shows

    b>σ1(σ1+σ2)+σ21(σ1+σ2)2+4σ1σ2β222σ2β2ifd>σ2β2σ1+σ2.

    For d>σ2β2σ1+σ2, we compute directly

    d(b2+1)+2bβ1β2>12σ2β2(σ1+σ2)[σ21(σ1+σ2)2+2σ1σ2β22+2σ22β22]σ1β1(σ1+σ2)σ2β2β2+12σ2β2(σ1+σ2)[(2β1σ1)(σ1+σ2)σ21(σ1+σ2)2+4σ1σ2β22]=2β1σ12σ2β2[σ21(σ1+σ2)2+4σ1σ2β22σ1(σ1+σ2)]>0. (2.26)

    Note that σ1σ2σ1σ2<σ2β2σ1+σ2, according to Lemma 2.2(ⅰ). As seen in (b) above, b>1 is equivalent to d>σ1σ2σ1σ2. Thus, we see from (2.24) that Gu(d)<0 for d>σ2β2σ1+σ2, due to (2.26), b>1, and b>0.

    (d') Case (ⅱ): σ2β2>σ1β1, i.e. σ1σ2σ1σ2>σ1σ2β1β2σ2β22σ1β21 by (2.2). We claim that the third term of (2.24): d(b2+1)+2bβ1β2>0 for dσ1σ2σ1σ2. If so, then it can be seen from (2.24) and b>0, that Gu(d)<0 for dσ1σ2σ1σ2 which is equivalent to b1. For dσ1σ2σ1σ2, we obtain

    d(b2+1)+2bβ1β22σ1σ2σ1σ2+2β1β2=1σ1σ2[2σ1σ2+2σ1β12σ2β1σ1β2+σ2β2]>1σ1σ2[2σ1σ2+3σ1β12σ2β1σ1β2]>1σ1σ2[2σ1σ2+3σ1β12σ2β1σ1(β1+σ1+σ2)]=2β1σ1>0,

    due to β1>σ1>0 and β2β1<σ1+σ2, mentioned in Remark 1.

    For case (ⅰ), we summarize properties (a')-(c'):

    {Gu(d)>0ifdσ1σ2σ1σ2Fu(d)<0ifd>σ1σ1σ2(α2α1)Fu(d)<0fordσ2β2σ1+σ2Gu(d)<0ford>σ2β2σ1+σ2.

    Recall Lemma 2.2(ⅰ): σ1σ2σ1σ2<σ2β2σ1+σ2<σ1σ2β1β2σ2β22σ1β21, and that Gu(d) and Fu(d) have identical sign. There are two possibilities:

    (Ⅰ) Fu(σ2β2σ1+σ2)0, i.e., Gu(σ2β2σ1+σ2)0: As Gu(d)<0 for d>σ2β2σ1+σ2 and Fu(d)<0 if d>σ1σ1σ2(α2α1), there exists a unique d3>0 such that Gu(d3)=0, where

    σ2β2σ1+σ2d3<σ1σ1σ2(α2α1).

    (Ⅱ) Fu(σ2β2σ1+σ2)<0, i.e., Gu(σ2β2σ1+σ2)<0: As Fu(d)>0 for dσ1σ2σ1σ2, Fu(d)<0 for dσ2β2σ1+σ2, and Gu(d)<0 for d>σ2β2σ1+σ2, we confirm that there exists a unique d3>0 such that Gu(d3)=0, where

    σ1σ2σ1σ2<d3<min{σ2β2σ1+σ2,σ1σ1σ2(α2α1)}.

    Both (Ⅰ) and (Ⅱ) indicate that there exists a unique d3>0 such that Gu(d3)=0, Gu(d)>0 if d<d3, and Gu(d)<0 if d>d3, where

    σ1σ2σ1σ2<d3<σ1σ1σ2(α2α1).

    For case (ⅱ), from the above (a'), (b'), and (d'), we summarize

    {Gu(d)>0ifdσ1σ2σ1σ2Fu(d)<0ifd>σ1σ1σ2(α2α1)Gu(d)<0ifdσ1σ2σ1σ2.

    We thus conclude that there exists a unique d3>0 such that Gu(d3)=0, Gu(d)>0 if d<d3, and Gu(d)<0 if d>d3, where

    σ1σ2σ1σ2<d3<σ1σ1σ2(α2α1).

    From (2.13), we see that v1α1+add>0 as u10+, i.e., u1 and v1 can not be zero simultaneously. From the above discussions, we confirm that d2<d3.

    Combining the above discussions of two scenarios for v1(d)>0, and one single scenario for u1(d)>0, the assertions are thus justified, see Figure 2.

    Figure 2.  The existence of u1(d),u2(d),v1(d),v2(d) with respect to d, in cases (ⅰ) and (ⅱ) of Theorem 2.1 respectively.

    Remark 2. (Ⅰ) Under conditions (C) and (P), the proof of Theorem 2.1 actually indicate:

    (ⅰ) If σ2β2<σ1β1 and β2β1σ1+σ2, then (u1,u2,v1,v2)(0,0,ˉv1,ˉv2), as d(d3), i.e., the positive equilibrium (u1,u2,v1,v2) degenerates and merges into the semitrivial equilibrium (0,0,ˉv1,ˉv2) at d=d3.

    (ⅱ) If σ2β2>σ1β1, then (u1,u2,v1,v2)(ˉu1,ˉu2,0,0), as d(d1), i.e., the positive equilibrium (u1,u2,v1,v2) degenerates and merges into the semitrivial equilibrium (ˉu1,ˉu2,0,0) at d=d1; (ˉu1,ˉu2,0,0)(u1,u2,v1,v2), as d(d2)+, i.e., the semitrivial equilibrium (ˉu1,ˉu2,0,0) becomes the positive equilibrium (u1,u2,v1,v2) as the value of d increases through d2; (u1,u2,v1,v2)(0,0,ˉv1,ˉv2), as d(d3), i.e., the positive equilibrium (u1,u2,v1,v2) again degenerates and merges into the semitrivial equilibrium (0,0,ˉv1,ˉv2) at d=d3.

    (Ⅱ) In Theorem 2.1(ⅱ), combining σ2β2>σ1β1 and condition (P) yields β2β1<σ1+σ2, which is contrary to condition β2β1σ1+σ2 in case (ⅰ), as mentioned in Remark 1.

    (Ⅲ) Although the same symbol d3 is used in Theorem 2.1 (ⅰ) and (ⅱ), they represent different values under assumptions in (ⅰ) and (ⅱ), respectively.

    (Ⅳ) With the setting a:=u2u1,b:=v2v1, and subsequently ab=σ1σ2=:k, we always have b2<k<a2. Notably, in [24], 0<k1 under assumption σ1σ2, and hence b<1. This is disparate from the situation in Theorem 2.1 that k>1, and hence a>1, due to σ1>σ2.

    In this section, we analyze the stability of the semitrivial equilibria for system (1.3). We denote by (ˉu1,ˉu2,0,0) and (0,0,ˉv1,ˉv2) the semitrivial (boundary) equilibria for system (1.3), and by ˉui(d) and ˉvi(d),i=1,2, to express the dependence of ˉui and ˉvi on d. In Appendix Ⅱ, we recall some properties of semitrivial equilibria of system (1.3) in Propositions 3.7-3.10 of [24], which are independent of the order between σ1 and σ2. Herein, we add the following additional properties for the semitrivial equilibria, which shall be employed to discuss the stability of semitrivial equilibria.

    Proposition 3.1. (ⅰ) If α1<α2, then ˉu1(d)>0,ˉu2(d)<0, ˉu1(d)<0, and ˉu2(d)>0, for all d>0.

    (ⅱ) If β1<β2, then ˉv1(d)>0,ˉv2(d)<0, ˉv1(d)<0, and ˉv2(d)>0, for all d>0.

    Proof. (ⅰ) (ˉu1,ˉu2,0,0) is an equilibrium of (1.3) if and only if ˉu1 and ˉu2 satisfy

    ˉu1(α1ˉu1)+d(ˉu2ˉu1)=0ˉu2(α2ˉu2)+d(ˉu1ˉu2)=0. (3.1)

    Differentiating (3.1) with respect to d, we obtain

    (α12ˉu1d)ˉu1+dˉu2+ˉu2ˉu1=0(α22ˉu2d)ˉu2+dˉu1+ˉu1ˉu2=0, (3.2)

    where ˉui,i=1,2, represent the derivatives of ˉui with respect to d. Thus,

    ˉu1=(α22ˉu2)(ˉu1ˉu2)(α12ˉu1d)(α22ˉu2d)d2, (3.3)
    ˉu2=(α12ˉu1)(ˉu2ˉu1)(α12ˉu1d)(α22ˉu2d)d2. (3.4)

    Note that

    (α12ˉu1d)(α22ˉu2d)d2=(α12ˉu1)(α22ˉu2)d(α12ˉu1)d(α22ˉu2)>0,

    by Proposition A.3 (in Appendix Ⅱ). Thus ˉu1>0 and ˉu2<0. More detailed descriptions for ˉu1 and ˉu2 can be found in Proposition A.5. We further differentiate (3.3) with respect to d, and obtain

    (α12ˉu1d)ˉu1+dˉu2=2ˉu12ˉu2+2(ˉu1)2(α22ˉu2d)ˉu2+dˉu1=2ˉu22ˉu1+2(ˉu2)2.

    Thus,

    ˉu1=2(α22ˉu2d)[ˉu1ˉu2+(ˉu1)2]d[ˉu2ˉu1+(ˉu2)2](α12ˉu1d)(α22ˉu2d)d2ˉu2=2(α12ˉu1d)[ˉu2ˉu1+(ˉu2)2]d[ˉu1ˉu2+(ˉu1)2](α12ˉu1d)(α22ˉu2d)d2.

    Let us focus on the numerators. For ˉu1, we have

    (α22ˉu2d)[ˉu1ˉu2+(ˉu1)2]d[ˉu2ˉu1+(ˉu2)2]=(α22ˉu2d)(ˉu1)2+(α22ˉu2)(ˉu1ˉu2)d(ˉu2)2<0,

    due to ˉu1>0, ˉu2<0 for all d>0, and Proposition A.3. Thus, ˉu1<0. For ˉu2, with (3.5) and (3.6), we have

    (α12ˉu1d)[ˉu2ˉu1+(ˉu2)2]d[ˉu1ˉu2+(ˉu1)2]=(α12ˉu1d)(ˉu2)2+(α12ˉu1)(ˉu2ˉu1)d(ˉu1)2=(α12ˉu1d)[(α12ˉu1)(ˉu2ˉu1)(α12ˉu1d)(α22ˉu2d)d2]2+(α12ˉu1)[(α12ˉu1)(ˉu2ˉu1)(α12ˉu1d)(α22ˉu2d)d2(α22ˉu2)(ˉu1ˉu2)(α12ˉu1d)(α22ˉu2d)d2]d[(α22ˉu2)(ˉu1ˉu2)(α12ˉu1d)(α22ˉu2d)d2]2=(ˉu2ˉu1)[(α12ˉu1d)(α22ˉu2d)d2]2{(α12ˉu1d)(α2ˉu1ˉu2)(α12ˉu1)2d(α12ˉu1)3+(α22ˉu2)2[(α12ˉu1d)(α12ˉu1)d(ˉu2ˉu1)]}.

    For the first two terms in the bracket,

    (α12ˉu1d)(α2ˉu1ˉu2)(α12ˉu1)2d(α12ˉu1)3>0,

    by Proposition A.3. For the third term, using (3.1), we have

    (α22ˉu2)2[(α12ˉu1d)(α12ˉu1)d(ˉu2ˉu1)]=(α22ˉu2)2[(α12ˉu1)2d(α12ˉu1)d(ˉu2ˉu1)]=(α22ˉu2)2{[d(1ˉu2ˉu1)ˉu1]2d[d(1ˉu2ˉu1)ˉu1]d(ˉu2ˉu1)}=(α22ˉu2)2{d2(1ˉu2ˉu1)2d2(1ˉu2ˉu1)+dˉu2+ˉu21}>0,

    since ˉu1<ˉu2. Thus, ˉu2>0.

    Part (ⅱ) can be obtained by arguments similar to those for (ⅰ), using

    ˉv1(β1ˉv1)+d(ˉv2ˉv1)=0ˉv2(β2ˉv2)+d(ˉv1ˉv2)=0. (3.5)

    This completes the proof.

    Propositions A.3-A.6, in Appendix Ⅱ, and Proposition 3.1 are independent of the order between σ1 and σ2. Some of the following properties for the semitrivial equilibria hold under σ2<σ1. The following notations will be helpful to recognize various related quantities:

    d1:=σ1σ2(σ21+σ22+σ2β2σ1β1)(σ1σ2)(σ21+σ22),d2:=σ1σ2σ1σ2,d3:=β1β2(σ2β1+σ1β2)(β2β1)(β21+β22),d4:=σ1σ2β1β2σ2β22σ1β21,d5:=σ1σ2(σ2α2σ1α1)(σ1+σ2)(σ1σ2).

    Proposition 3.2. Under conditions (C) and (P), the following relationships among parameters hold:

    (Ⅰ) ˉu2ˉu1(d) is strictly decreasing with respect to d.

    (Ⅱ) If ˉu2ˉu1=σ1σ2, then d=d1.

    (Ⅲ) If ˉu2ˉu1=β2β1, then d=d3.

    (Ⅳ) If ˉu2ˉu1=σ1σ2, then d=d5.

    (Ⅴ) (ⅰ) If σ2β2<σ1β1, then σ1σ2>β2β1 and d1<d2<d3<d4.

    (ⅱ) If σ2β2>σ1β1, then σ1σ2<β2β1 and d1>d2>d3>d4.

    (ⅲ) If σ2β2=σ1β1, then σ1σ2=β2β1 and d1=d2=d3=d4.

    Proof. (Ⅰ) The assertion follows from ˉu1(d)>0,ˉu2(d)<0, as in the proof of Proposition 3.1.

    (Ⅱ) If ˉu2ˉu1=σ1σ2, with ˉu1 and ˉu2 satisfying (3.1), we have

    α1ˉu1+d(σ1σ21)=0α2σ1σ2ˉu1+d(σ2σ11)=0.

    By eliminating ˉu1, we have

    d(σ31σ21σ2σ32+σ1σ22σ21σ2)=σ2α2σ1α1σ1.

    Then

    d=σ1σ2(σ21+σ22+σ2β2σ1β1)(σ1σ2)(σ21+σ22)=d1,

    due to α2=β2+σ2 and α1=β1σ1.

    Cases (Ⅲ) and (Ⅳ) can be obtained by arguments similar to those for (Ⅱ). Now let us prove (Ⅴ), and the assertions will be justified by the following (a)-(c):

    (a) It is clear that

    d2d1{>0ifσ2β2<σ1β1=0ifσ2β2=σ1β1<0ifσ2β2>σ1β1.

    (b) We see that

    d3d2{>0ifσ2β2<σ1β1=0ifσ2β2=σ1β1<0ifσ2β2>σ1β1.

    as, by a direct calculation,

    d3d2=(σ1β22+σ2β21)(σ1β1σ2β2)(σ1σ2)(β2β1)(β21+β22).

    (c) It holds that

    d4d3{>0ifσ2β2<σ1β1=0ifσ2β2=σ1β1<0ifσ2β2>σ1β1

    due to

    d4d3=(σ1+σ2)β21β22(σ1β1σ2β2)(σ2β22σ1β21)(β2β1)(β21+β22).

    This completes the proof.

    In Appendix Ⅰ, we compute the Jacobian matrix for system (1.3). At (ˉu1,ˉu2,0,0), the Jacobian matrix is

    [α12ˉu1ddˉu10dα22ˉu2d0ˉu200β1ˉu1dd00dβ2ˉu2d], (3.6)

    and at (0,0,ˉv1,ˉv2), the Jacobian matrix is

    [α1ˉv1dd00dα2ˉv2d00ˉv10β12ˉv1dd0ˉv2dβ22ˉv2d]. (3.7)

    First, let us focus on the stability of semitrivial equilibrium (ˉu1,ˉu2,0,0), by calculating the eigenvalues of the following submatrices in (3.6):

    [α12ˉu1dddα22ˉu2d]and[β1ˉu1dddβ2ˉu2d]. (3.8)

    Theorem 3.3. Assume that conditions (C) and (P) hold for system (1.3).

    (ⅰ) If σ2β2<σ1β1 and β2β1σ1+σ2, the semitrivial equilibrium (ˉu1,ˉu2,0,0) is unstable for all d>0.

    (ⅱ) If σ2β2>σ1β1, there exist ˉd1,ˉd2>0, with ˉd1<ˉd2, so that the semitrivial equilibrium (ˉu1,ˉu2,0,0) is unstable when d<ˉd1 or d>ˉd2 and is asymptotically stable when ˉd1<d<ˉd2.

    In addition,

    σ1σ2α1α2σ2α22σ1α21<ˉd1<σ1σ2β1β2σ2β22σ1β21,σ1σ2σ1σ2<ˉd2<σ1σ1σ2+σ1σ2σ21σ22(α2α1).

    Proof. Under condition (C), the two eigenvalues of the first matrix in (3.11) are negative by Gerschgorin's Theorem and Proposition A.3. Thus, the stability of (ˉu1,ˉu2,0,0) is determined by the two eigenvalues, denoted by λ, of the second matrix in (3.11). By a direct calculation, the two eigenvalues are

    λ:=12[(β1ˉu1+β2ˉu22d)(β1ˉu1β2+ˉu2)2+4d2].

    First, we consider λ=λ(d) and claim λ(d)<0 for all d>0. From condition (C) and Proposition A.3, we have

    β1ˉu1+β2ˉu2=(σ1σ2)+d[2(ˉu2ˉu1+ˉu1ˉu2)],

    and

    β1ˉu1β2+ˉu2=(σ1+σ2)+d(ˉu1ˉu2ˉu2ˉu1).

    Then

    λ=12[(β1ˉu1+β2ˉu22d)(β1ˉu1β2+ˉu2)2+4d2]=12[(σ1σ2)d(ˉu2ˉu1+ˉu1ˉu2)[(σ1+σ2)+d(ˉu1ˉu2ˉu2ˉu1)]2+4d2]<12[(σ1σ2)d(ˉu2ˉu1+ˉu1ˉu2)|(σ1+σ2)+d(ˉu1ˉu2ˉu2ˉu1)|].

    We obtain

    λ<σ2dˉu1ˉu2<0,

    if (σ1+σ2)+d(ˉu1ˉu2ˉu2ˉu1)0, and

    λ<σ1dˉu2ˉu1<0,

    if (σ1+σ2)+d(ˉu1ˉu2ˉu2ˉu1)<0. Consequently, λ(d)<0 for all d>0.

    Next, we identify the sign of λ+=λ+(d). Note that λ+(d)0 if and only if

    |β1ˉu1+β2ˉu22d|(β1ˉu1β2+ˉu2)2+4d2,

    equivalently,

    (β1ˉu1)(β2ˉu2)d(β1ˉu1+β2ˉu2)0. (3.9)

    As β1=α1+σ1 and β2=α2σ2, (3.12) can be expressed by

    (α1ˉu1+σ1)(α2ˉu2σ2)d[α1ˉu1+α2ˉu2+(σ1σ2)]0,

    i.e.,

    [d(1ˉu2ˉu1)+σ1][d(1ˉu1ˉu2)σ2]d2[2(ˉu2ˉu1+ˉu1ˉu2)]d(σ1σ2)0,

    using (3.1). This inequality can be simplified to

    d(σ2ˉu2ˉu1σ1ˉu1ˉu2)σ1σ20. (3.10)

    From (3.10), we define

    g(d):=d(σ2ˉu2(d)ˉu1(d)σ1ˉu1(d)ˉu2(d))σ1σ2. (3.11)

    Then λ+(d)0 if and only if g(d)0. According to Propositions A.3 and A.5, we have that 1<ˉu2(d)ˉu1(d)<α2α1 and ˉu2(d)ˉu1(d) decreases from α2α1 to 1 as d increases from 0 to . Thus, g(0)=σ1σ2 and g(d) as d, because of σ1>σ2. More precisely,

    g(d)=d(σ2ˉu2ˉu1σ1ˉu1ˉu2)σ1σ2<d(σ2α2α1σ1α1α2)σ1σ2=d(σ2α22σ1α21α1α2)σ1σ20,ifdσ1σ2α1α2σ2α22σ1α21. (3.12)

    Note that ˉu2(d)ˉu1(d)σ1σ2 is equivalent to dd5, by Proposition 3.2. Hence,

    g(d)=d(σ2ˉu2ˉu1σ1ˉu1ˉu2)σ1σ2d(σ2σ1σ2σ1σ2σ1)σ1σ2=σ1σ2<0,ifˉu2(d)ˉu1(d)σ1σ2.

    Thus,

    g(d)<0ifdd5. (3.13)

    A direct calculation yields

    g(d)=(σ2ˉu2ˉu1σ1ˉu1ˉu2)+d[σ2(ˉu2ˉu1)σ1(ˉu1ˉu2)]. (3.14)

    We know σ2(ˉu2ˉu1)σ1(ˉu1ˉu2)<0 for all d>0, due to ˉu1>0 and ˉu2<0, as in the proof of Proposition 3.1 or by Proposition A.5; σ2ˉu2ˉu1σ1ˉu1ˉu2=σ2α22σ1α21α1α2>0 when d=0, by Lemma 2.1(ⅱ); σ2ˉu2ˉu1σ1ˉu1ˉu2σ2σ1<0 as d+, due to Propositions A.3 and A.5. That is, σ2ˉu2ˉu1σ1ˉu1ˉu2 decreases from σ2α22σ1α21α1α2 to (σ1σ2) as d increases from 0 to +. On the other hand, by (3.1), we have

    {d(ˉu2ˉu1)=1ˉu2ˉu1+ˉu1d(ˉu1ˉu2)=1ˉu1ˉu2+ˉu2. (3.15)

    With (3.15), we reexpress (3.14) as

    g(d)=σ2σ1+σ2ˉu1σ1ˉu2. (3.16)

    It follows that

    g(d)=σ2ˉu1σ1ˉu2<0,

    by Proposition 3.1. Thus, the graph of g(d) is concave downward. Therefore, there are two possible situations based on the above analysis: (ⅰ) g(d)<0 for all d>0, (ⅱ) there exist ˉd1,ˉd2>0 such that g(ˉd1)=g(ˉd2)=0, and

    {g(d)<0ifd<ˉd1ord>ˉd2g(d)>0ifˉd1<d<ˉd2.

    The graphs of g(d) are illustrated in Figure 3. Accordingly, there are two possibilities for λ+: (ⅰ) λ+>0 for all d>0, (ⅱ) there exist ˉd1,ˉd2>0 such that λ+=0 for d=ˉd1,ˉd2 and

    {λ+>0ifd<ˉd1ord>ˉd2λ+<0ifˉd1<d<ˉd2.
    Figure 3.  Two situations for g(d), regarding the sign of λ+ in Theorem 3.3.

    Now we investigate the two situations by analyzing g(d) and the stationary equation (3.1). To determine the behavior of g, we seek for its equivalent expression. Let w:=ˉu2ˉu1. As ˉu2ˉu1(d) is strictly decreasing with respect to d, by Proposition 3.2(Ⅰ), the one-to-one correspondence between d and w can be derived from the stationary equation for ˉu1 and ˉu2 in (3.1):

    d=α2wα1(w1)(w+1w), (3.17)

    where 1<w<α2α1, by Proposition A.5. Then

    g(d)=d(σ2ˉu2ˉu1σ1ˉu1ˉu2)σ1σ2=d(σ2w2σ1w)σ1σ2=(α2wα1(w1)(w+1w))(σ2w2σ1w)σ1σ2=(α2wα1)(σ2w2σ1)σ1σ2(w1)(w2+1)(w1)(w2+1)=:f(w).

    Let us define q(w):=(α2wα1)(σ2w2σ1)σ1σ2(w1)(w2+1), which is the numerator of f(w), and thus f(w)=q(w)(w1)(w2+1). Note that

    q(w)=(α2wα1)(σ2w2σ1)σ1σ2(w1)(w2+1)=(β2wβ1)(σ2w2σ1)+w(σ1+σ2)(σ2wσ1), (3.18)

    by β1=α1+σ1 and β2=α2σ2. Thus, we have

    g(d)<0f(w)<0q(w)<0, (3.19)

    due to (w1)(w2+1)>0. In addition,

    f(w)=q(w)(w1)(w2+1)q(w)(3w22w+1)(w1)2(w2+1)2, (3.20)

    where

    q(w)=2σ2w(β2wβ1)+(σ1+σ2)(σ2wσ1)σ2β1w2+σ2(σ1+σ2)w+σ1β1. (3.21)

    Notice that

    f(w)<0g(d)>0,

    according to Proposition 3.2(Ⅰ). By a direct computation in (3.18), we obtain

    q(σ1σ2)=σ1σ22(σ1σ2)(σ2β2σ1β1){<0ifσ2β2<σ1β1=0ifσ2β2=σ1β1>0ifσ2β2>σ1β1, (3.22)

    and

    q(β2β1)=β2β21(σ1+σ2)(σ2β2σ1β1){<0ifσ2β2<σ1β1=0ifσ2β2=σ1β1>0ifσ2β2>σ1β1. (3.23)

    Case (ⅰ) σ2β2<σ1β1, i.e., β2β1<σ1σ2: We first claim that q(w)<0 for all β2β1<w<σ1σ2. Consider w=β2β1+δ, with δ>0 satisfying β2β1<w=β2+δβ1β1<σ1σ2. Hence, σ2(β2+δβ1)<σ1β1. From (3.18), we obtain

    q(w=β2+δβ1β1)=δβ1[σ2(β2+δβ1)2σ1β21]1β21{(σ1+σ2)(β2+δβ1)[σ1β1σ2(β2+δβ1)]}<0,

    by σ2β22σ1β21>0 from condition (P), and σ1β1>σ2(β2+δβ1).

    Next, we will show that f(σ1σ2)<0 and f(β2β1)>0. From (3.20) and (3.21), at w=σ1σ2, a direct calculation yields

    q(w)(w1)(w2+1)q(w)(3w22w+1)=σ1(σ1σ2)σ22{σ21σ22[σ2β2+σ2β1+σ2(σ1+σ2)]+[σ1β1+σ2β1+σ2(σ1+σ2)]}+σ1(σ1σ2)σ22(2σ1σ2+1)(σ2β2σ1β1)<σ1(σ1σ2)σ22[σ2(σ21σ22+1)(β2β1σ1σ2)(2σ1σ2+1)(σ1β1σ2β2)]<0,

    because of β2β1σ1+σ2. Thus, f(σ1σ2)<0 by (3.20) and (w1)2(w2+1)2>0. Similarly, from (3.20) and (3.21), at w=β2β1, we have

    q(w)(w1)(w2+1)q(w)(3w22w+1)=1β41[(σ1+σ2)(σ1β1σ2β2)(2β32+β31β1β22)]+1β41{(β2β1)(β21+β22)[σ2β22+σ1β21+σ2β2(σ1+σ2)]}>1β41[(σ1+σ2)(σ1β1σ2β2)(β32+2β31β21β2)]>0,

    due to condition (P), i.e., σ2β22<σ1β1(β1+σ1+σ2) in (2.1). Thus, f(β2β1)>0 by (3.20) and (w1)2(w2+1)2>0.

    Consequently, by (3.19) and concavity of g(d), we conclude f(w)<0 for all 1<w=ˉu2ˉu1<α2α2, i.e., g(d)<0 for all d>0, i.e., λ+>0 for all d>0.

    Case (ⅱ) σ2β2>σ1β1, i.e., β2β1>σ1σ2: We claim that q(w)>0 for all β2β1>w>σ1σ2. Consider w=σ1σ2+δ, with δ>0 satisfying β2β1>w=σ1+δσ2σ2>σ1σ2. Hence, σ2β2>(σ1+δσ2)β1. From (3.18), we compute

    q(w=σ1+δσ2σ2)=δ(σ1+σ2)(σ1+δσ2)+1σ22[(σ1+δσ2)2σ1σ2][σ2β2(σ1+δσ2)β1]>0,

    owing to σ1>σ2 and σ2β2>(σ1+δσ2)β1. Combining (3.22) with (3.23), we have q(w)>0 for all β2β1wσ1σ2. That is, g(d)>0 if d3dd1, by Proposition 3.2. Recall the relationship between d and w(d)=ˉu2(d)ˉu1(d) in Proposition 3.2. We will use the relationship to estimate ˉd1 and ˉd2. For ˉu2ˉu1>β2β1, we have d<d3 by Proposition 3.2, and then

    g(d)>d(σ2β22σ1β21β1β2)σ1σ20ifdd4.

    Thus, g(d)>0 if d4d<d3. According to (3.12), (3.13), and Proposition 3.2, we obtain

    {g(d)<0ifdσ1σ2α1α2σ2α22σ1α21g(d)>0ifd4=σ1σ2β1β2σ2β22σ1β21dσ1σ2(σ21+σ22+σ2β2σ1β1)(σ1σ2)(σ21+σ22)=d1g(d)<0ifdσ1σ2(σ2α2σ1α1)(σ1+σ2)(σ1σ2)=d5.

    In addition, we recall Lemma 2.1(ⅱ): 0<σ1σ2α1α2σ2α22σ1α21<σ1σ2β1β2σ2β22σ1β21=d4. Accordingly, there exist ˉd1,ˉd2>0 such that λ+=0 for d=ˉd1,ˉd2 and

    {λ+>0ifd<ˉd1ord>ˉd2λ+<0ifˉd1<d<ˉd2

    where

    σ1σ2α1α2σ2α22σ1α21<ˉd1<σ1σ2β1β2σ2β22σ1β21=d4d1=σ1σ2(σ21+σ22+σ2β2σ1β1)(σ1σ2)(σ21+σ22)<ˉd2<σ1σ2(σ2α2σ1α1)(σ1+σ2)(σ1σ2)=d5.

    By a direct computation, it can be seen that

    σ1σ2(σ2α2σ1α1)(σ1+σ2)(σ1σ2)<σ1σ1σ2+σ1σ2σ21σ22(α2α1).

    Thus, for consistency with the ranges in the assertion of Theorem 2.1, we also write

    σ1σ2α1α2σ2α22σ1α21<ˉd1<σ1σ2β1β2σ2β22σ1β21σ1σ2σ1σ2<ˉd2<σ1σ1σ2+σ1σ2σ21σ22(α2α1).

    This completes the proof.

    Remark 3. Assume that conditions (C) and (P) hold. Under σ2β2=σ1β1 and β2β1=σ1+σ1, we do have g(d1=d2=d3=d4)=0 and g(d1=d2=d3=d4)=0, by Proposition 3.2(Ⅴ)(ⅲ) and the proof of Theorem 3.3, and hence ˉd1=ˉd2.

    Next, let us focus on the boundary equilibrium (0,0,ˉv1,ˉv2). We shall discuss the stability of (0,0,ˉv1,ˉv2) through analyzing the eigenvalues of the following submatrices in (3.7):

    [α1ˉv1dddα2ˉv2d]and[β12ˉv1dddβ22ˉv2d]. (3.24)

    Theorem 3.4. Consider system (1.3) under condition (C). There exists a ˉd3>0 so that the boundary equilibrium (0,0,ˉv1,ˉv2) is unstable if d<ˉd3 and asymptotically stable if d>ˉd3. In addition,

    σ1σ2σ1σ2<ˉd3<σ1σ1σ2(α2α1).

    Proof. Under condition (C), it can be shown that the two eigenvalues of the second matrix in (3.24) are negative, by Gerschgorin's Theorem and Proposition A.4 in Appendix Ⅱ. The stability of (0,0,ˉv1,ˉv2) is thus determined by the two eigenvalues, denoted by λ, of the first matrix in (3.24). By a direct calculation, these two eigenvalues are

    λ:=12[(α1ˉv1+α2ˉv22d)(α1ˉv1α2+ˉv2)2+4d2].

    From Proposition A.4, we have

    α1ˉv1+α2ˉv22d=(β1ˉv1+β2ˉv2)(σ1σ2)2d<0,

    and thus λ<0 for all d>0. Next, let us identify the sign of λ+=λ+(d). Note that λ+(d)0 if and only if

    α1ˉv1+α2ˉv22d∣≤(α1ˉv1α2+ˉv2)2+4d2(α1ˉv1)(α2ˉv2)d(α1ˉv1+α2ˉv2)0. (3.25)

    By α1=β1σ1, α2=β2+σ2, (3.5), and Proposition A.4, (3.25) is equivalent to

    [d(1ˉv2ˉv1)σ1][d(1ˉv1ˉv2)+σ2]d2[2(ˉv2ˉv1+ˉv1ˉv2)]+d(σ1σ2)0.

    This inequality can be simplified to

    d(σ1ˉv1ˉv2σ2ˉv2ˉv1)σ1σ20. (3.26)

    Now, we define

    h(d):=d(σ1ˉv1ˉv2(d)σ2ˉv2ˉv1(d))σ1σ2. (3.27)

    Then λ+(d)0 if and only if h(d)0. By Propositions A.4 and A.6, we have 1<ˉv2ˉv1(d)<β2β1 and ˉv2ˉv1(d) decreases from β2β1 to 1, as d increases from 0 to . Hence, h(0)=σ1σ2<0 and h(d) as d due to σ1>σ2.

    A direct calculation yields

    h(d)=(σ1ˉv1ˉv2(d)σ2ˉv2ˉv1(d))+d[σ1(ˉv1ˉv2)(d)σ2(ˉv2ˉv1)(d)]. (3.28)

    Notably, σ1(ˉv1ˉv2)σ2(ˉv2ˉv1)>0 for all d>0, σ1ˉv1ˉv2σ2ˉv2ˉv1=σ2β22σ1β21β1β2 if d=0, and σ1ˉv1ˉv2σ2ˉv2ˉv1σ1σ2>0 as d; namely, σ1ˉv1ˉv2σ2ˉv2ˉv1 increases from σ2β22σ1β21β1β2 to σ1σ2 as d increases from 0 to . Moreover, from (3.5), the equations for ˉvi, we have

    {d(ˉv2ˉv1)=1ˉv2ˉv1+ˉv1d(ˉv1ˉv2)=1ˉv1ˉv2+ˉv2. (3.29)

    With (3.28) and (3.29), we obtain

    h(d)=(σ1σ2)+σ1ˉv2σ2ˉv1, (3.30)

    and then

    h(d)=σ1ˉv2σ2ˉv1>0,

    by Proposition 3.1. Thus, the graph of h(d) is concave upward. Therefore, from the above discussions, there is a unique ˉd3>0 such that

    {h(d)<0ifd<ˉd3h(d)=0ifd=ˉd3h(d)>0ifd>ˉd3

    and accordingly,

    {λ+(d)>0ifd<ˉd3λ+(d)=0ifd=ˉd3λ+(d)<0ifd>ˉd3. (3.31)

    The graph of h(d) is illustrated in Figure 4.

    Figure 4.  The graph of h(d), regarding the sign of λ+ in Theorem 3.4.

    Now, we estimate the range for the values of ˉd3. Function h in (3.30) can be expressed by

    h(d)=σ2(β1ˉv1)σ1(β2ˉv2)+d(σ1σ2)σ1σ2, (3.32)

    via (3.5). Thus, inequality (3.26) is equivalent to

    σ2(β1ˉv1)σ1(β2ˉv2)+d(σ1σ2)σ1σ20. (3.33)

    According to Proposition A.4(ⅰ), we have

    σ1(β2β1)<σ2(β1ˉv1)σ1(β2ˉv2)<0.

    Then,

    h(σ1σ2σ1σ2)<σ1σ2σ1σ2=0,

    and

    h(σ1σ1σ2(α2α1))>σ1(β2β1)+σ1(α2α1)σ1σ2=σ1(β2β1)+σ1(β2β1+σ1+σ2)σ1σ2=σ21>0.

    Consequently, (3.31) holds with

    σ1σ2σ1σ2<ˉd3<σ1σ1σ2(α2α1).

    This completes the proof.

    Let us summarize the main results in Sections 2 and 3:

    (ⅰ) Under σ2β2<σ1β1 and β2β1σ1+σ2, the positive equilibrium (u1,u2,v1,v2) exists if d<d3 (Theorem 2.1); the semitrivial equilibrium (ˉu1,ˉu2,0,0) is unstable for all d>0 (Theorem 3.3) and the semitrivial equilibrium (0,0,ˉv1,ˉv2) is unstable if d<ˉd3 and asymptotically stable if d>ˉd3 (Theorem 3.4). Besides, the estimated range of d3 in Theorem 2.1 coincides with the one of ˉd3 in Theorem 3.4.

    (ⅱ) Under σ2β2>σ1β1, the positive equilibrium (u1,u2,v1,v2) exists if d<d1 or d2<d<d3 (Theorem 2.1); the semitrivial equilibrium (ˉu1,ˉu2,0,0) is unstable if d<ˉd1 or d>ˉd2, and asymptotically stable if ˉd1<d<ˉd2 (Theorem 3.3); the semitrivial equilibrium (0,0,ˉv1,ˉv2) is unstable if d<ˉd3 and asymptotically stable if d>ˉd3 (Theorem 3.2). In addition, the estimated ranges of d1 and d2 in Theorem 2.1 respectively coincide with those of ˉd1 and ˉd2 in Theorem 3.3.

    In fact, the following theorem reveals that these critical values of d are consistent in determining the existence of the positive equilibrium and the stability of semitrivial equilibria, namely, d1=ˉd1, d2=ˉd2 and d3=ˉd3. Such interesting consistency makes precise the global dynamics of this competitive species model (1.3), under the framework of monotone dynamics. Let us elaborate.

    Theorem 4.1. d1=ˉd1, d2=ˉd2 and d3=ˉd3.

    Proof. From (3.26), we see that h(d)=0 if and only if

    d(σ1ˉv1ˉv2(d)σ2ˉv2ˉv1(d))=σ1σ2.

    That is, ˉd3 satisfies

    ˉd3(σ1ˉv1ˉv2(ˉd3)σ2ˉv2ˉv1(ˉd3))=σ1σ2.

    Let ˉb:=ˉv2ˉv1(ˉd3), then

    ˉd3(σ11ˉbσ2ˉb)=σ1σ2.

    Thus,

    ˉb=σ1+σ21+4kˉd232ˉd3, (4.1)

    where k=σ1σ2. Recall the definition of b in (2.7). From Remark 2(Ⅰ), we see that (u1,u2,v1,v2)(0,0,ˉv1,ˉv2), as d(d3). Therefore, recalling (2.10), we obtain

    σ1+σ21+4k(d3)22d3=limd(d3)b(d)=limd(d3)v2v1(d)=ˉv2ˉv1(d3).

    Noting that b=b(d) in (2.10) is monotone in d (shown in (2.18)), with (4.1), we thus conclude that d3=ˉd3.

    If σ2β2>σ1β1, from (3.11), we have g(ˉd1)=0 and g(ˉd2)=0. Let ˉa1:=ˉu2ˉu1(ˉd1), ˉa2:=ˉu2ˉu1(ˉd2). Then

    ˉa1=σ1+σ21+4kˉd212ˉd1,ˉa2=σ1+σ21+4kˉd222ˉd2. (4.2)

    It follows from Remark 2(Ⅰ) that (u1,u2,v1,v2)(ˉu1,ˉu2,0,0), as d(d1). Notice that a=a(d) in (2.10) is monotone in d, shown in (2.25). Therefore, recalling (2.10), we have

    σ1+σ21+4k(d1)22d1=limd(d1)a(d)=limd(d1)u2u1(d)=ˉu2ˉu1(d1).

    With (4.2), we thus conclude that d1=ˉd1. In addition, by Remark 2(Ⅰ), we see that (ˉu1,ˉu2,0,0)(u1,u2,v1,v2) as d(d2)+. Consequently,

    limd(d2)+ˉa2(d)=limd(d2)+ˉu2ˉu1(d)=u2u1(d2)=σ1+σ21+4k(d2)22d2.

    With (4.2), we conclude that d2=ˉd2. This completes the proof.

    Combining the discussions in Sections 2 and 3 with the assertion in Theorem 4.1, we conclude that for system (1.3), either there exists a positive equilibrium representing the coexistence of two species or one species drives the other to extinction, depending on the magnitude of the dispersal rate d.

    Theorem 4.2. Consider system (1.3) under conditions (C), and (P).

    (Ⅰ) Assume that σ2β2<σ1β1 and β2β1σ1+σ2 hold.

    (ⅰ) If d<d3, then the positive equilibrium (u1,u2,v1,v2) is stable, and

    limt(u1(t),u2(t), v1(t),v2(t))=(u1,u2,v1,v2), for all (u1(0),u2(0),v1(0),v2(0))R4+ with u1(0)+u2(0)>0 and v1(0)+v2(0)>0.

    (ⅱ) If dd3, then limt(u1(t),u2(t),v1(t),v2(t))=(0,0,ˉv1,ˉv2), for all (u1(0),u2(0), v1(0),v2(0))R4+ with v1(0)+v2(0)>0.

    (Ⅱ) Assume that σ2β2>σ1β1 holds.

    (ⅰ) If d<d1 or d2<d<d3, then the positive equilibrium (u1,u2,v1,v2) is stable, and limt(u1(t),u2(t), v1(t),v2(t))=(u1,u2,v1,v2), for all (u1(0),u2(0),v1(0),v2(0))R4+ with u1(0)+u2(0)>0 and v1(0)+v2(0)>0.

    (ⅱ) = If d1dd2, then limt(u1(t),u2(t),v1(t),v2(t))=(ˉu1,ˉu2,0,0), for all (u1(0),u2(0), v1(0),v2(0))R4+ with u1(0)+u2(0)>0.

    (ⅲ) If dd3, then limt(u1(t),u2(t),v1(t),v2(t))=(0,0,ˉv1,ˉv2), for all (u1(0),u2(0), v1(0),v2(0))R4+ with v1(0)+v2(0)>0.

    Proof. The assertions are based on the monotone dynamics theory which is reviewed in Appendix I. (I) Assume that σ2β2<σ1β1 and β2β1σ1+σ2 hold. (ⅰ) If d<d3, then the positive equilibrium (u1,u2,v1,v2) is unique, by Theorem 2.1, the semitrivial equilibrium (ˉu1,ˉu2,0,0) is unstable, by Theorem 3.3, and the semitrivial equilibrium (0,0,ˉv1,ˉv2) is unstable, by Theorem 3.4. Therefore, the assertion follows from Theorem A.1. (ⅱ) If d>d3, then case (a) of the trichotomy in Theorem A.2 does not hold, since the positive steady state does not exist, by Theorem 2.1; case (b) does not hold since (ˉu1,ˉu2,0,0) is unstable, by Theorem 3.3, and (0,0,ˉv1,ˉv2) is asymptotically stable, by Theorem 3.4. Therefore, the assertion follows from case (c) of Theorem A.2.

    (Ⅱ) Assume that σ2β2>σ1β1. (ⅰ) If d<d1 or d2<d<d3, the argument is similar to the one in (Ⅰ)(ⅰ). (ⅱ) If d1<d<d2, then case (a) of the trichotomy in Theorem A.2 does not hold, since the positive equilibrium does not exist, by Theorem 2.1; case (c) does not hold since (0,0,ˉv1,ˉv2) is unstable, by Theorem 3.2, and (ˉu1,ˉu2,0,0) is asymptotically stable, by Theorem 3.1. Therefore, the assertion follows from case (b) of Theorem A.2. (ⅲ) If d>d3, the argument is similar to the one in (Ⅰ)(ⅱ). This completes the proof.

    Remark 4 That the equilibrium (0,0,ˉv1,ˉv2) is globally asymptotically stable for d>d3 now follows from Theorem 4.2. In fact it also holds true for d=d3. In this case, the stability for (0,0,ˉv1,ˉv2) can be concluded by some comparison argument. In addition, there is no positive equilibrium and the equilibrium (ˉu1,ˉu2,0,0) is unstable in both cases (Ⅰ) and (Ⅱ), by Theorem 2.1 and Theorem 3.3. Hence the trichotomy in Theorem A.2 implies the global convergence to (0,0,ˉv1,ˉv2). Similarly, we see that the equilibrium (ˉu1,ˉu2,0,0) is globally asymptotically stable for d=d1 or d2.

    We arrange two examples to illustrate the global dynamics of system (1.3), and the bifurcation with respect to the dispersal rate d, which are concluded in Theorem 4.2. We also present two more examples to demonstrate that the established scenarios still hold without satisfying condition (P).

    Example 1. Consider system (1.3) with α1=1, α2=3, β1=1.5 and β2=2.8, i.e., σ1=0.5 and σ2=0.2. Let us examine the conditions in Theorem 4.2(Ⅰ): condition (C): α1=1<β1=1.5<β2=2.8<α2=3 with σ2=0.2<σ1=0.5; condition (P): σ2β2σ1+σ2=0.8<σ1σ2β1β2σ2β22σ1β21=0.948; σ2β2=0.56<σ1β1=0.75, and β2β1=1.3σ1+σ2=0.7. We depict in Figure 5 the bifurcation diagram with respect to the dispersal rate d. It appears that d31.22, which is consistent with Theorems 2.1(ⅰ) and 4.2(Ⅰ): σ1σ2σ1σ2=0.333<d3<σ1σ1σ2(α2α1)=3.33. The globally stable positive equilibrium (u1,u2,v1,v2) exists for d<d3 and collides with the semitrivial equilibrium (0,0,ˉv1,ˉv2) at d=d3. For dd3, the semitrivial equilibrium (0,0,ˉv1,ˉv2) becomes globally attractive.

    Figure 5.  Bifurcation diagram, with respect to d, for the equilibria of system (1.3) with α1=1,α2=3,β1=1.5,β2=2.8,σ1=0.5 and σ2=0.2, where σ2β2<σ1β1.

    Example 2. Consider system (1.3) with α1=1, α2=3, β1=1.7 and β2=2.5, i.e., σ1=0.7 and σ2=0.5. Let us examine the conditions in Theorem 4.2(Ⅱ): condition (C): α1=1<β1=1.7<β2=2.5<α2=3 with σ2=0.5<σ1=0.7; condition (P): σ2β2σ1+σ2=1.042<σ1σ2β1β2σ2β22σ1β21=1.35; σ2β2=1.25>σ1β1=1.19. The bifurcation diagram with respect to the dispersal rate d is depicted in Figure 6. It appears that d10.91, d21.92, d34.15, which are consistent with Theorem 2.1(ⅱ) and 4.2(Ⅱ): σ1σ2α1α2σ2α22σ1α21=0.276<d1<σ1σ2β1β2σ2β22σ1β21=1.35, σ1σ2σ1σ2=1.75<d2<σ1σ1σ2+σ1σ2σ21σ22(α2α1)=6.36, and σ1σ2σ1σ2=1.75<d3<σ1σ1σ2(α2α1)=7. The globally stable positive equilibrium (u1,u2,v1,v2) exists for d<d1 and collides with the semitrivial equilibrium (ˉu1,ˉu2,0,0) at d=d1. For d1dd2, the equilibrium (ˉu1,ˉu2,0,0) becomes globally attractive and for d2<d<d3, the globally stable positive equilibrium (u1,u2,v1,v2) exists. For dd3, the semitrivial equilibrium (0,0,ˉv1,ˉv2) becomes globally attractive.

    Figure 6.  Bifurcation diagram, with respect to d, for the equilibria of system (1.3) with α1=1,α2=3,β1=1.7,β2=2.5,σ1=0.7 and σ2=0.5, where σ2β2>σ1β1.

    Example 3. Consider system (1.3) with α1=1, α2=3, β1=1.4 and β2=2.85, i.e., σ1=0.4 and σ2=0.15. For such parameter values, condition (C) holds: α1=1<β1=1.4<β2=2.85<α2=3 with σ2=0.15<σ1=0.4. In addition, σ2β2=0.4275<σ1β1=0.56. Such parameter values violate condition (P), as σ2β2σ1+σ2=0.777>σ1σ2β1β2σ2β22σ1β21=0.551. Nevertheless, the same dynamical scenario as Example 1 takes place, as seen in Figure 7.

    Figure 7.  Bifurcation diagram, with respect to d, for the equilibria of system (1.3) with α1=1,α2=3,β1=1.4,β2=2.85,σ1=0.4 and σ2=0.15, where σ2β2<σ1β1.

    Example 4. Consider system (1.3) with α1=1, α2=3, β1=1.35 and β2=2.8, i.e., σ1=0.35 and σ2=0.2. With such parameter values, condition (C) holds: α1=1<β1=1.35<β2=2.8<α2=3 with σ2=0.2<σ1=0.35; σ2β2=0.56>σ1β1=0.4725. These parameter values violate condition (P), as σ2β2σ1+σ2=1.0182>σ1σ2β1β2σ2β22σ1β21=0.2845. Nevertheless, the dynamical scenario shown in Figure 8 remains identical to Example 2.

    Figure 8.  Bifurcation diagram, with respect to d, for the equilibria of system (1.3) with α1=1,α2=3,β1=1.35,β2=2.8,σ1=0.35 and σ2=0.2, where σ2β2>σ1β1.

    We have exhibited the global dynamics for a model on two-species competition in a two-patch environment, under certain conditions. The main condition (C): α1<β1<β2<α2, (β1+β2)(α1+α2)=σ1σ2>0, indicates that the birth rate of u-species in the second patch is the largest among all birth rates of two species on two patches, yet the average birth rate of v-species is larger than u-species. This means that the birth rate for v-species is larger than u-species in the first patch. The present investigation exploited analytically two dynamical scenarios for such competition, as demonstrated in Examples 1 and 2, respectively. The first scenario takes place under σ2β2<σ1β1. As expressed by σ1σ2>β2β1>1, it indicates that the value of σ1 is larger than the value of σ2 in a way that its ratio exceeds the ratio of β2 over β1. This includes the situation that σ1 is much bigger than σ2, which is denoted by (β1+β2)(α1+α2). The second scenario comes about under σ2β2>σ1β1. On the contrary, as expressed by 1<σ1σ2<β2β1, it indicates that the value of σ1 may be merely a little over the value of σ2, depending on the ratio of β2 over β1. In this case, the average birth rate of v-species may be merely a little more than and close to the average birth rate of u-species; we denote this situation by (β1+β2)(α1+α2).

    In the first case, including the sense (β1+β2)(α1+α2), coexistence of two species occurs for dispersal rate d<d3, and (0,0,ˉv1,ˉv2) is globally attractive for dd3, where d3 has been estimated by system parameters. In this situation, (ˉu1,ˉu2,0,0) is unstable for any d>0 and an eigenvalue of the linearized system at (0,0,ˉv1,ˉv2) changes from positive to negative as d, being increasing from 0, exceeds d3, and (0,0,ˉv1,ˉv2) becomes stable for dd3.

    In the second case, including the sense (β1+β2)(α1+α2), the coexistence of two species takes place for d<d1 or d2<d<d3, (ˉu1,ˉu2,0,0) is globally attractive for d1dd2, and (0,0,ˉv1,ˉv2) becomes globally attractive for dd3, where d1,d2,d3 have been estimated. An eigenvalue of the linearized system at (ˉu1,ˉu2,0,0) changes from positive to negative at d1, and then back to positive at d2. In addition, an eigenvalue of the linearized system at (0,0,ˉv1,ˉv2) changes from positive to negative at d3, and d2<d3.

    Our analytical investigation on this model strongly suggests that, in high-dispersal situations, one species will prevail if its average birth rate is larger than the other competing species, whereas in low-dispersal situations, the two species can coexist or one species that has the greatest birth rate in one patch among all species and patches will be able to persist and drive the other species to extinction, even though its average birth rate is lower. Such findings may illuminate some insights into how species learn to compete and point out the evolution directions.

    Condition (C) is a basic assumption for the present results. Although there are additional conditions (P) and β2β1σ1+σ2, due to mathematical technicality, it is believed that such scenarios remain true under condition (C) only. However, it is very difficult to remove these additional conditions, as the algebraic operations involving five parameters are rather involved. In Examples 3, 4, we have demonstrated exactly the same dynamical scenarios for parameter values which do not satisfy condition (P).

    To compare our results with those in [19], we set σ1=ξσ2, ξ>1, according to condition (C). The resource difference between two species can be depicted as (σ1,σ2) among two patches, where β1α1=σ1>0 means that v-species has an advantage over v-species in competing the resource in patch-1, while β2α2=σ2<0 means that it is disadvantageous for v-species to compete with u-species for the resource in patch-2. We rewrite it as σ2(ξ,1) with fixed ξ>1, and now the value of σ2 measures the difference between two species and resembles the value of τ in [19]. We accordingly rewrite the conditions in Theorem 4.2 to explore how the magnitude of resource difference affects the invasion of mutant species:

    (P)σ2>σ2:=β22ξβ21ξ(ξ+1)β1,β2β1(<)σ1+σ2σ2(>)σ2:=β2β1ξ+1,σ2β2<(>)σ1β1ξ>(<)β2β1.

    Note that the criteria in Theorem 4.2(Ⅱ) imply β2β1<σ1+σ2. Therefore, by increasing the dispersal rate d, the global convergence of system (1.1) switches in case (Ⅰ) of Theorem 4.2 from the coexistence to extinction of u-species when σ2<σ2σ2 and ξ>β2β1; on the other hand, in case (Ⅱ), the dynamics switches three times from global convergence to the coexistence to extinction of (mutant) v-species, again to the coexistence and then the persistence of v-species, when σ2>σ2 and ξ<β2β1. This result enhances the understanding on the dynamics of competitive species from the viewpoint of patchy habitat in the following aspects: Compared to concluding global convergence under small magnitude of spatial heterogeneity (τ) in [19,Theorem 1.2], our result in Theorem 4.2 admits a large range of magnitudes (σ2) depicting spatial heterogeneity. The multiple stability switches in Theorem 4.2 are global dynamics, as compared to local dynamics in [19,Theorem 1.1].

    The authors are grateful to Chao-Nien Chen for suggesting to study the case with average birth rates of opposite order, and to Yuan Lou for very helpful discussions. The authors are supported, in part, by the Ministry of Science and Technology, Taiwan.

    All authors declare no conflicts of interest in this paper.

    For reader's convenience, we review some theory on monotone dynamical systems from [17] and [28]. Denote by Rn+={x=(x1,,xn)Rn:xi0,1in} the first octant of Rn. For x,yRn+, define the following order: xmy if yxKm, and xmy whenever yxIntKm, where

    Km={xRn:xi0,1ik,andxj0,k+1jn}=Rk+×(Rnk+).

    If xmy, we define [x,y]m={zRn+:xmzmy} and (x,y)m={zRn+:xmzmy}.

    A semiflow ϕ is said to be of type-K monotone with respect to Km, provided

    ϕt(x)mϕt(y)wheneverxmy,t0.

    A system of ODEs ˙x=f(x) is called a type-K monotone system with respect to Km if the Jacobian matrix of f takes the form

    [A1A2A3A4],

    where A1 is an k×k matrix, A4 an (nk)×(nk) matrix, A2 an k×(nk) matrix, A3 an (nk)×k matrix, every off-diagonal entry of A1 and A4 is nonnegative, and A2 and A3 are nonnegative matrices, for some k with 1kn. It was shown in [27] that the flow ϕt(x) generated by the type-K monotone system is type-K monotone with respect to the cone Km, i.e., if x,yRn+ with xiyi for 1ik and xjyj for k+1jn, then for any t>0, (ϕt(x))i(ϕt(y))i for 1ik and (ϕt(x))j(ϕt(y))j for k+1jn.

    System (1.3) is a type-K monotone system with respect to

    Km={(u1,u2,v1,v2):ui0,vi0,i=1,2},

    since its Jacobian matrix is

    [α12u1v1ddu10dα22u2v2d0u2v10β12v1u1dd0v2dβ22v2u2d].

    For system (1.3), let us denote by e0:=(0,0,0,0) the trivial equilibrium, by eˉu:=(ˉu1,ˉu2,0,0), and eˉv:=(0,0,ˉv1,ˉv2), ˉui, ˉvi>0, i=1,2, the semitrivial equilibria. If (u1,u2,v1,v2)R4+, then (0,0,v1,v2)m(u1,u2,v1,v2)m(u1,u2,0,0), and therefore,

    ϕt((0,0,v1,v2))mϕt((u1,u2,v1,v2))mϕt((u1,u2,0,0)),

    for all t0. Since ϕt((0,0,v1,v2))eˉv and ϕt((u1,u2,0,0))eˉu as t, for (u1,u2,v1,v2)R4+, and u1+u2>0, v1+v2>0, it follows that all points in R4+ are attracted to the set

    Γ:=[0,ˉu1]×[0,ˉu2]×[0,ˉv1]×[0,ˉv2]=[eˉv,eˉu]m={wR4+:eˉvmwmeˉu}.

    If w=(u1,u2,v1,v2) with u1,u2,v1,v2>0, then ϕt(w)0 for t>0. Define E and E+ the sets of all nonnegative equilibria and all positive equilibria for ϕt, respectively. Obviously, [eˉv,eˉu]m contains E and e(eˉv,eˉu)m for any eE+. The following theorem restates Corollary 4.4.3 in [28] for system (1.3), see also [27,31].

    Theorem A.1. If eˉu and eˉv are both linearly unstable, then system (1.3) is permanent. More precisely, there exist positive equilibria e and e, not necessarily distinct, satisfying

    eˉvmememeˉu.

    The order interval

    [e,e]m:={w:emwme}

    attracts all solutions evolved from w=(u1,u2,v1,v2)R4+, with u1+u2>0 and v1+v2>0. In particular, if e=e, then e attracts all such solutions.

    It was shown in [17] that, for models of two competing species, either there is a positive equilibrium representing coexistence of two species, or one species drives the other to extinction. Note that system (1.3) satisfies conditions (H1)-(H4) in [17], and thus Theorem B in [17] can be restated as follows.

    Theorem A.2. Consider system (1.3). The ω-limit set of every orbit evolved from a point in R4+ is contained in Γ and exactly one of the following holds:

    (a) There exists a positive equilibrium e of in Γ.

    (b) ϕt(w)eˉu as t, for every w=(u1,u2,v1,v2)Γ with u1+u2>0.

    (c) ϕt(w)eˉv as t, for every w=(u1,u2,v1,v2)Γ with v1+v2>0.

    In addition, if (b) or (c) holds, then either ϕt(w)eˉu or ϕt(w)eˉv, as t, for w=(u1,u2,v1,v2)R4+Γ.

    We recall some qualitative properties of the semitrivial equilibria for system (1.3) in [24]. The following results are independent of the order between σ1 and σ2.

    Proposition A.3 (Proposition 3.7 [24]). If α1<α2, the following hold for all d>0.

    (ⅰ) α1<ˉu1<ˉu2<α2.

    (ⅱ) (α1ˉu1)(α2ˉu2)=d(ˉu21ˉu22)ˉu1ˉu2<0, (α1ˉu1)+(α2ˉu2)=d[2(ˉu2ˉu1+ˉu1ˉu2)]<0.

    (ⅲ) α1<ˉu1<α1+α22<ˉu2<α2.

    Proposition A.4 (Proposition 3.8 [24]). If β1<β2, the following hold for all d>0.

    (ⅰ) β1<ˉv1<ˉv2<β2.

    (ⅱ) (β1ˉv1)(β2ˉv2)=d(ˉv21ˉv22)ˉv1ˉv2<0, (β1ˉv1)+(β2ˉv2)=d[2(ˉv2ˉv1+ˉv1ˉv2)]<0.

    (ⅲ) β1<ˉv1<β1+β22<ˉv2<β2.

    Proposition A.5 (Proposition 3.9 [24]). If α1<α2, the following hold:

    (ⅰ) ˉu1,ˉu2α1+α22 as d.

    (ⅱ) d is strictly decreasing with respect to ˉu2 on (α1+α22,α2), and d is strictly increasing with respect to ˉu1 on (α1,α1+α22).

    Proposition A.6 (Proposition 3.10 [24]). If β1<β2, the following hold:

    (ⅰ) ˉv1,ˉv2β1+β22 as d.

    (ⅱ) d is strictly decreasing with respect to ˉv2 on (β1+β22,β2), and d is strictly increasing with respect to ˉv1 on (β1,β1+β22).



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