Coexistence of a cross-diffusive West Nile virus model in a heterogenous environment

  • Received: 05 June 2018 Revised: 13 August 2018 Published: 01 December 2018
  • MSC : Primary: 35K57, 92B05; Secondary: 35J60

  • This paper is concerned with a strongly-coupled elliptic system, which describes a West Nile virus (WNv) model with cross-diffusion in a heterogeneous environment. The basic reproduction number is introduced through the next generation infection operator and some related eigenvalue problems. The existence of coexistence states is presented by using a method of upper and lower solutions. The true positive solutions are obtained by monotone iterative schemes. Our results show that a cross-diffusive WNv model possesses at least one coexistence solution if the basic reproduction number is greater than one and the cross-diffusion rates are small enough, while if the basic reproduction number is less than or equal to one, the model has no positive solution. To illustrate the impact of cross-diffusion and environmental heterogeneity on the transmission of WNv, some numerical simulations are given.

    Citation: Abdelrazig K. Tarboush, Jing Ge, Zhigui Lin. Coexistence of a cross-diffusive West Nile virus model in a heterogenous environment[J]. Mathematical Biosciences and Engineering, 2018, 15(6): 1479-1494. doi: 10.3934/mbe.2018068

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  • This paper is concerned with a strongly-coupled elliptic system, which describes a West Nile virus (WNv) model with cross-diffusion in a heterogeneous environment. The basic reproduction number is introduced through the next generation infection operator and some related eigenvalue problems. The existence of coexistence states is presented by using a method of upper and lower solutions. The true positive solutions are obtained by monotone iterative schemes. Our results show that a cross-diffusive WNv model possesses at least one coexistence solution if the basic reproduction number is greater than one and the cross-diffusion rates are small enough, while if the basic reproduction number is less than or equal to one, the model has no positive solution. To illustrate the impact of cross-diffusion and environmental heterogeneity on the transmission of WNv, some numerical simulations are given.


    1. Introduction

    Infectious diseases have been attracting considerable attention in recent years, and various epidemic models have been proposed and analyzed for prevention and control strategies, especially for vector borne diseases [15,39]. For example, a West Nile virus (WNv) is an arbovirus of the Flavivirus kind in the family Flaviviridae that causes the epidemics of febrile illness and sporadic encephalitis [7]. WNv is found in temperate and tropical regions of the world, it was first isolated and identified from the blood of a febrile Ugandan woman during research on yellow fever virus in 1937 [3].

    Although WNv is widely distributed in Africa, the Middle East, Asia and southern Europe, in North America, the first infected case was detected in 1999 during an outbreak of encephalitis in New York city [3,21,26,39]. Since 1999 this virus has spread spatially and prevail in much of North America [8,21], it is evident that the spread of WNv comes from the interplay of disease dynamics and bird and mosquito movement.

    To the best of our knowledge, currently there are no effective vaccine or medicine for WNv. To reduce the rates of WNv infection, anti-WNv efforts are primarily based on personal protective measures like insect repellent and protective clothing, and public heath measures [4].

    Many mathematical models for WNv have been proposed and analyzed, however most of the models are focused on the non-spatial transmission dynamics [4,36,39]. In fact, the spatial spreading is an important factor to affect the persistence and eradication of WNv. In 2006, Lewis et al. [21] investigated the spatial spread of WNv to describe the movement of birds and mosquitoes. The reaction-diffusion model was extended from the non-spatial model for cross infection between birds and mosquitoes that was proposed and developed by Wonham et al. in [39].

    To utilize the cooperative characteristic of cross-infection dynamics and estimate the spatial spread rate of infection, Lewis et al. in [21] proposed the following simplified WNv model

    {Ibt=D1ΔIb+αbβb(NbIb)NbImγbIb,(x,t)Ω×(0,+),Imt=D2ΔIm+αmβb(AmIm)NbIbdmIm,(x,t)Ω×(0,+), (1)

    where the positive constants Nb and Am denote the total population of birds and adult mosquitos; Ib(x,t) and Im(x,t) represent the populations of infected birds and mosquitos at the location x in the habitat ΩRN and at time t0, respectively, and Ib(x,0)+Im(x,0)>0. The parameters in the above system are defined as follows:

    αm, αb : WNv transmission probability per bite to mosquitoes and birds, respectively;

    βb : biting rate of mosquitoes on birds;

    dm : adult mosquitos death rate;

    γb : bird recovery rate from WNv.

    Here, the positive constants D1 and D2 are diffusion coefficients for birds and mosquitoes, respectively.

    Considering the spatially-independent model

    {dIb(t)dt=γbIb(t)+αbβb(NbIb(t))NbIm(t),t>0,dIm(t)dt=dmIm(t)+αmβb(AmIm(t))NbIb(t),t>0, (2)

    one can see that if R0(:=αmαbβ2bAmdmγbNb )<1, the virus will vanish eventually, while for R0>1, a nontrivial epidemic level appears and is globally asymptotically stable in the positive quadrant [21].

    For the diffusive model (1), Lewis et al. proved the existence of traveling wave and calculated the spatial spread rate of infection [21]. The corresponding free boundary problem describing the expanding process has been discussed in [34]. It is worth mentioning that WNv usually spreads from one area to another because of the diffusions of birds and mosquitoes, so that its transmission is affected not only by the characteristics of pathogens, but also by the spatial difference of environment in which birds or mosquitoes reside. Considering the complexity of diffusions and the heterogeneity of environment, the system (1) can be extended to the following strongly-coupled parabolic system

    {IbtΔ[(d1+α1Ib+β1Ibγ1+Im)Ib]=αb(x)βb(x)(NbIb)NbImγb(x)Ib,(x,t)Ω×(0,+),ImtΔ[(d2+β2Imγ2+Ib+α2Im)Im]=αm(x)βb(x)(AmIm)NbIbdm(x)Im,(x,t)Ω×(0,+),Ib(x,t)=Im(x,t)=0,(x,t)Ω×(0,+),0Ib(x,0)Nb,0Im(x,0)Am,x¯Ω, (3)

    and the corresponding elliptic problem with Dirichlet boundary conditions becomes

    {Δ[(d1+α1Ib+β1Ibγ1+Im)Ib]=f1(x,Ib,Im),xΩ,Δ[(d2+β2Imγ2+Ib+α2Im)Im]=f2(x,Ib,Im),xΩ,Ib(x)=Im(x)=0,xΩ, (4)

    where

    f1(x,Ib,Im)=αb(x)βb(x)(NbIb)NbImγb(x)Ib,f2(x,Ib,Im)=αm(x)βb(x)(AmIm)NbIbdm(x)Im,

    and the parameters αb(x), βb(x), γb(x), αm(x) and dm(x) are all sufficiently smooth and strictly positive functions defined on ¯Ω. di (i=1,2) is positive constants represent the free-diffusion coefficients of population Ib(x) and Im(x), respectively. αi, βi and γi (i=1,2) are nonnegative constants; αi, βi and γi are the self-diffusion coefficients, the cross-diffusion rates and cross-diffusion pressures, respectively. The homogeneous Dirichlet boundary condition in (4) means that there is no infection on the boundary and outside of the domain Ω. More specifically, the diffusion terms can be written as

    div{(d1+2α1Ib+2β1Ibγ1+Im)Ib+β1I2b(γ1+Im)2Im},
    div{β2I2m(γ2+Ib)2Ib+(d2+2β2Imγ2+Ib+2α2Im)Im}.

    The terms

    d1+2α1Ib+2β1Ibγ1+Im,d2+2β2Imγ2+Ib+2α2Im

    represent the self-diffusions and the terms

    β1I2b(γ1+Im)2,β2I2m(γ2+Ib)2

    represent the cross-diffusions. Here the term β1I2b(γ1+Im)2Im is due to the moving of the population of birds Ib toward the location of increasing population of mosquitoes Im with rate (β1I2b(γ1+Im)2<0). Similarly, the term β2I2m(γ2+Ib)2Ib is due to the moving of the population of mosquitoes Im toward the location of increasing population of birds Ib with rate (β2I2m(γ2+Ib)2<0). The solution (Ib(x),Im(x)) to problem (4) is called a coexistence if Ib(x)>0 and Im(x)>0 for every xΩ.

    Problem (1) is weakly-coupled parabolic system which only consider the random diffusion in a homogeneous environment. However, problem (4) implies that, in addition to the dispersive force, the diffusion also depends on population pressure from other population. This means that the population in (4) are not homogeneously distributed due to the consideration of self and cross diffusion terms. Moreover, the diffusive behavior in different populations also affect the distribution of resources. Thus, the consideration of diffusion and cross-diffusion effect is very reasonable and more close to reality, see for example, [29] for mixed-culture biofilm model, [17] for the tumor-growth model and [14,16,31,40] for the competition model. There are some valuable results about the roles of diffusion and cross-diffusion in the modeling of the dynamics of strongly coupled reaction-diffusion systems [5,11,13,16,19,22,25,31,30,38,40]. For instance, Shigesada et al. [31] proposed the strongly coupled elliptic system describing two species Lotka-Volterra competition model. Ko and Ryu studied a predator-prey system with cross-diffusion, representing the tendency of prey to keep away from its predators, under the homogeneous Dirichlet boundary conditions in [19]. Fu et al. investigated the global behavior of solutions for a Lotka-Volterra predator-prey system with prey-stage structure, under the homogeneous Neumann boundary conditions [11]. In 2014, Jia et al. [16] discussed a Lotka-Volterra competition reaction-diffusion system with nonlinear diffusion effects. In 2016, Braverman and Kamrujjaman [5] introduced a competitive-cooperative models with various diffusion strategies. More recently, Li et al. studied an effect of cross-diffusion on the stationary problem of a Leslie prey-predator model with a protection zone [22].

    In recent years, researches on the existence and non-existence of the positive solutions for the dynamics of strongly-coupled elliptic systems have received comprehensive attention [16,19,20,28]. There are many standard approaches to derive the coexistence for the standard semi-linear parabolic system in mathematical models, such as construction of upper and lower solutions [12,16,18,28], bifurcation theory [6], fixed point theorem [19,42], ect. The upper and lower solutions method developed by Pao [27] is concise and effective to derive the coexistence. Based on the method, the coexistence for a general strongly-coupled system has been given in [28]. In [18], Kim and Lin studied the coexistence of three species in strongly coupled elliptic system. Gan and Lin in [12] considered the competitor-competitor-mutualist three species Lotka-Volterra model. Recently, Jia et al. [16] investigated the existence of the positive steady state solution of a Lotka-Volterra competition model with cross-diffusion.

    Motivated by above problems, in this paper we are more interested in the nonnegative steady state solutions, that is, the coexistence of problem (4) describing a cross-diffusive WNv model in a heterogenous environment.

    The plan of this paper is as follows: Section 2 is devoted to the basic reproduction number of problem (4) and its properties. The existence and non-existence of coexistence to (4) are discussed in Section 3. Finally, some numerical simulations and a brief discussion are given in Section 4.


    2. Basic reproduction numbers

    In this section, we first present the basic reproduction number for problem (4) and its properties for the corresponding system in Ω. According to [10], the basic reproduction number is an expected number of secondary cases produced by a typical infected individual during its entire period of infectiousness in a completely susceptible population, and mathematically was defined as the dominant eigenvalue of a positive linear operator. Usually the basic reproduction numbers for the spatially homogenous models were calculated by the next generation matrix method [35], while for the spatially-dependent models, the numbers could be presented in the term of the principal eigenvalue of related eigenvalue problem [1] or the spectral radius of next infection operator [37,41].

    Considering the linearized problem of (3), we have

    {Ibtd1ΔIb=αb(x)βb(x)Imγb(x)Ib,xΩ,t>0,Imtd2ΔIm=Amαm(x)βb(x)NbIbdm(x)Im,xΩ,t>0,Ib(x)=Im(x)=0,xΩ. (5)

    We now consider the following linear reaction-diffusion system

    {utDΔu=F(x)uV(x)u,xΩ,t>0,u(x)=0,xΩ, (6)

    where

    u=(IbIm),D=(d100d2),
    F(x)=(0αb(x)βb(x)Amαm(x)βb(x)Nb0),V(x)=(γb(x)00dm(x)).

    In addition, the interval evolution of individuals in the infectious compartments is governed by the following linear system

    {utDΔu=V(x)u,xΩ,t>0,u(x)=0,xΩ. (7)

    Let X1:=C(¯Ω,R2) and X+1:=C(¯Ω,R2+). Set T(t) be the solution semigroup on X1 associated with system (7). We let Ψ=(ϕ,ψ) is the density distribution of u at the spatial location xΩ, we then see that T(t)Ψ:=(T(t)ϕ,T(t)ψ) represents the remaining distribution of infective birds and mosquitoes at time t. Therefore, the distribution of total new infective members is

    0F(x)[T(t)Ψ](x)dt.

    Following the idea of [37,41], we define the linear operator

    L(Ψ)(x):=0F(x)[T(t)Ψ](x)dt.

    It follows from the definition, we know that L is a continuous and positive operator which maps the initial infection distribution Ψ to the distribution of the total members produced during the infection period. Consequently, we define the spectral radius of L as the basic reproduction number of system (5), that is,

    RD0=ρ(L).

    As in [23], to ensure the existence of the basic reproduction numbers we consider the following linear eigenvalue problem:

    {d1Δϕ=αb(x)βb(x)Rψγb(x)ϕ+μϕ,xΩ,d2Δψ=Amαm(x)βb(x)NbRϕdm(x)ψ+μψ,xΩ,ϕ(x)=ψ(x)=0,xΩ. (8)

    For any R>0, the system is strongly cooperative, that is, αb(x)βb(x)>0 and αm(x)βb(x)AmNb>0 for all x¯Ω. According to [2,9,33], for any R>0, there exists a unique value μ:=μ1(R), and called the principal eigenvalue, such that problem (8) admits a unique solution pair (ϕR,ψR) (subject to constant multiples) with ϕR>0 and ψR>0 in Ω. Moreover, μ1(R) is algebraically simple and dominant, and the following properties hold.

    Lemma 2.1. μ1(R) is continuous and strictly increasing.

    With the above definition, we have the following relation between the two principal eigenvalues.

    Theorem 2.2. sign(1RD0) = sign(λ0), where RD0=RD0(Ω,γb(x),dm(x)) is the principal eigenvalue of the eigenvalue problem

    {d1Δϕ=αb(x)βb(x)RD0ψγb(x)ϕ,xΩ,d2Δψ=Amαm(x)βb(x)NbRD0ϕdm(x)ψ,xΩ,ϕ(x)=ψ(x)=0,xΩ (9)

    and λ0 is the principal eigenvalue of the eigenvalue problem

    {d1Δϕ=αb(x)βb(x)ψγb(x)ϕ+λ0ϕ,xΩ,d2Δψ=Amαm(x)βb(x)Nbϕdm(x)ψ+λ0ψ,xΩ,ϕ(x)=ψ(x)=0,xΩ. (10)

    Proof. In fact, λ0=μ1(1). On the other hand, one can easily deduce from the monotonicity with respect to the coefficients in (8) that limR0+μ1(R)<0 and limR+μ1(R)>0, therefore RD0 is the unique positive root of the equation μ1(R)=0. The result follows from the monotonicity of μ1(R) with respect to R.

    Remark 1. Recalling that μ1 is monotonically increasing with respect to βb(x), in the sense that μ1(βb,1(x))<μ1(βb,2(x)) if βb,1(x)βb,2(x) and βb,1(x))βb,2(x) in Ω, we deduce from Lemma 2.1 that RD0 is monotonically increasing with respect to βb(x), and RD0>1 if βb(x) is sufficiently large.

    If all coefficients are constant, we can provide an explicit formula for RD0, which is known as the basic reproduction number for the corresponding diffusive WNv model.

    Theorem 2.3. If αb(x)=αb, αm(x)=αm,βb(x)=βb, γb(x)=γb and dm(x)=dm, then the principal eigenvalue RD0 for (9), or the basic reproduction number for model (4), is expressed by

    RD0(Ω)=Amαb(βb)2αmNb[d1λ+γm][d2λ+dm] , (11)

    where λ is the principal eigenvalue of Δ in Ω with null Dirichlet boundary condition.

    Proof. Let ψ be the eigenfunction corresponding to the principal eigenvalue (λ) of Δ in Ω with null Dirichlet boundary condition and

    P=Amαb(βb)2αmNb[d1λ+γb][d2λ+dm],
    ϕ=αbβbR[d1λ+γb]ψ.

    Then we know that (ϕ,ψ) is a positive solution of problem (9) with RD0=P, and (11) follows directly from the uniqueness of the principal eigenvalue of (9).


    3. Coexistence

    In this section, inspired by [16,18,20,28], we first study the existence of a coexistence solution to problem (4) by constructing upper and lower solutions and then we establish the non-existence of the coexistence solution to problem (4). For the convenience, we let

    S={(Ib,Im)C(¯Ω)×C(¯Ω);(^Ib,^Im)(Ib,Im)(~Ib,~Im),x¯Ω)},

    where (^Ib,^Im) and (~Ib,~Im) are given in the following definition.

    Next we are going to give a sufficient condition for problem (4) to possess a positive solution by constructing upper and lower solutions as in [28]. To achieve this, we first give an equivalent form of problem (4):

    {Δ[H1(Ib,Im)]=f1(x,Ib,Im),xΩ,Δ[H2(Ib,Im)]=f2(x,Ib,Im),xΩ,Ib(x)=Im(x)=0,xΩ. (12)

    where

    H1(Ib,Im)=(d1+α1Ib+β1Ibγ1+Im)Ib,
    H2(Ib,Im)=(d2+β2Imγ2+Ib+α2Im)Im.

    Taking

    u=H1(Ib,Im),v=H2(Ib,Im),

    then the Jacobian J of the transformation (Ib,Im)(u,v) is given by

    J=(u,v)(Ib,Im)=|d1+2α1Ib+2β1Ibγ1+Imβ1I2b(γ1+Im)2β2I2m(γ2+Ib)2d2+2β2Imγ2+Ib+2α2Im|=(d1+2α1Ib+2β1Ibγ1+Im)(d2+2β2Imγ2+Ib+2α2Im)β1β2I2bI2m(γ1+Im)2(γ2+Ib)2d1d2+4β1β2IbIm(γ1+Im)(γ2+Ib)β1β2I2bI2m(γ1+Im)2(γ2+Ib)2d1d2>0for(Ib,Im)(0,0).

    Therefore, the inverse Ib=g1(u,v), Im=g2(u,v) exist whenever (Ib,Im)(0,0). Hence, problem (4) reduces to the following equivalent form

    {Δu+k1u=F1(x,Ib,Im),xΩ,Δv+k2v=F2(x,Ib,Im),xΩ,Ib=g1(u,v),Im=g2(u,v),xΩ,u(x)=v(x)=0,xΩ, (13)

    where Fi(x,Ib,Im)=kiHi(Ib,Im)+fi(x,Ib,Im) (i=1,2) with ki>0 (i=1,2) chosen later.

    In addition, from an elementary computations one can check that

    Ibu=d2+2α2Im+2β2Imγ2+IbJ,Ibv=β1I2b(γ1+Im)2J,
    Imu=β2I2m(γ2+Ib)2J,Imv=d1+2α1Ib+2β1Ibγ1+ImJ,

    which shows that Ib=g1(u,v) is nondecreasing in both u and v, while Im=g2(u,v) is also nondecreasing in both u and v for all (Ib,Im)(0,0).

    For the later analysis, we present the definition of upper and lower solutions to problem (13) as follows.

    Definition 3.1. Assume that F1 and F2 are nondecreasing with respect to Ib and Im. A pair of 4- nonnegative functions (~Ib,~Im,˜u,˜v), (^Ib,^Im,ˆu,ˆv) in C2(Ω)C(¯Ω) are called ordered upper and lower solutions of (13), if

    (0,0)(ˆIb,ˆIm)(˜Ib,˜Im)(Nb,Am),(ˆu,ˆv)(˜u,˜v)

    and

    {Δ˜u+k1˜uF1(x,~Ib,~Im),xΩ,Δ˜v+k2˜vF2(x,~Ib,~Im),xΩ,Δˆu+k1ˆuF1(x,^Ib,^Im),xΩ,Δˆv+k2ˆvF2(x,^Ib,^Im),xΩ,˜Ibg1(˜u,˜v),ˆIbg1(ˆu,ˆv),xΩ,˜Img2(˜u,˜v),ˆImg2(ˆu,ˆv),xΩ,˜u(x)0ˆu(x),˜v(x)0ˆv(x),xΩ. (14)

    For definiteness, we select

    ˜Ib=g1(˜u,˜v),  ˜Im=g2(˜u,˜v);
    ˆIb=g1(ˆu,ˆv),  ˆIm=g2(ˆu,ˆv),

    which is equivalent to

    ˜u=H1(˜Ib,˜Im),  ˜v=H2(˜Ib,˜Im);
    ˆu=H1(ˆIb,ˆIm),  ˆv=H2(ˆIb,ˆIm).

    Then the requirements of (˜Ib,˜Im) and (ˆIb,ˆIm) in (14) are satisfied and those of (˜u,˜v),(ˆu,ˆv) are reduced to

    {Δ[H1(˜Ib,˜Im)]+k1H1(˜Ib,˜Im)F1(x,~Ib,~Im),xΩ,Δ[H2(˜Ib,˜Im)]+k2H2(˜Ib,˜Im)F2(x,~Ib,~Im),xΩ,Δ[H1(ˆIb,ˆIm)]+k1H1(ˆIb,ˆIm)F1(x,^Ib,^Im),xΩ,Δ[H2(ˆIb,ˆIm)]+k2H2(ˆIb,ˆIm)F2(x,^Ib,^Im),xΩ,˜Ib(x)0ˆIb(x),˜Im(x)0ˆIm(x),xΩ. (15)

    Now we consider the monotonicity of Fi (i=1,2). From direct computations it is easy to see that

    F1Ib=k1(d1+2α1Ib+2β1Ibγ1+Im)αb(x)βb(x)ImNbγb(x),
    F2Im=k2(d2+2β2Imγ2+Ib+2α2Im)αm(x)βb(x)IbNbdm(x).

    If we choose

    k1=maxx¯Ω{αbβbAm+NbγbNbd1}(x),k2=maxx¯Ω{αmβb+dmd2}(x),

    then F1 and F2 are increasing with respect to Ib and Im, respectively, as long as (0.0)(Ib,Im)(Nb,Am). On the other hand, the direct calculations show that

    F1Im=k1β1(γ1+Im)2I2b+αb(x)βb(x)NbIbNb,
    F2Ib=k2β2(γ2+Ib)2I2mαm(x)βb(x)AmImNb.

    Note F1Im(Nb,Am)0 and F2Ib(Nb,Am)0 for any β1γ1 and β2γ2, respectively. To ensure that F1Im0, F2Ib0, we have to modify the upper solution and we seek ((1δ0)Nb,(1δ0)Am) as a new upper solution, where

    δ0minx¯Ω{12,γbαbβbAm,dmαbβb}

    from the first and second inequalities of (15). Let

    β1=minx¯Ωδ0αbβbγ21N2bk1(x),β2=minx¯Ωδ0αmβbγ22NbAmk2(x),

    then for β1β1 and β2β2, Fi (i=1,2) is monotone nondecreasing with respect to Ib and Im. Consequently, (~Ib,~Im,˜u,˜v), (^Ib,^Im,ˆu,ˆv) are a pair of ordered upper and lower solutions of problem (13).

    To present the existence of a positive solution to (4), it suffices to find a pair of upper and lower solutions of (4). We seek such as in the form (˜Ib,˜Im)=(M1,M2), (ˆIb,ˆIm)=(g1(δd1ϕ,δd2ψ),g2(δd1ϕ,δd2ψ)) where Mi (i=1,2) and δ are some positive constants with δ small enough, (ϕ,ψ)(ϕ(x),ψ(x)) is (normalized) positive eigenfunction corresponding to λ0, and λ0 is the principal eigenvalue of eigenvalue problem (10).

    Indeed, (M1,M2) and (g1(δd1ϕ,δd2ψ),g2(δd1ϕ,δd2ψ)) satisfy the inequalities in (15) if

    {Δ[(d1+α1M1+β1M1γ1+M2)M1]αb(x)βb(x)(NbM1)NbM2γb(x)M1,Δ[(d2+β2M2γ2+M1+α2M2)M2]αm(x)βb(x)(AmM2)NbM1dm(x)M2,Δ[d1ϕ]αb(x)βb(x)(NbˆIb)Nbd2ψ/(d2+α2ˆIm+β2ˆImγ2+ˆIb)γb(x)d1ϕ/(d1+α1ˆIb+β1ˆIbγ1+ˆIm),Δ[d2ψ]αm(x)βb(x)(AmˆIm)Nbd1ϕ/(d1+α1ˆIb+β1ˆIbγ1+ˆIm)dm(x)d2ψ/(d2+α2ˆIm+β2ˆImγ2+ˆIb). (16)

    The first two inequalities in (16) hold if we set

    (M1,M2)=((1δ0)Nb,(1δ0)Am). (17)

    Next, we notice that the relations

    δd1ϕ=(d1+α1ˆIb+β1ˆIbγ1+ˆIm)ˆIb, δd2ψ=(d2+α2ˆIm+β2ˆImγ2+ˆIb)ˆIm

    imply that 0<ˆIbδϕ and 0<ˆImδψ.

    If RD0>1, the principal eigenvalue of problem (10) is λ0<0, therefore we can choose δ sufficiently small such that the last two inequalities in (16) hold. Consequently, the pair (˜Ib,˜Im)=(M1,M2), (ˆIb,ˆIm)=(g1(δd1ϕ,δd2ψ),g2(δd1ϕ,δd2ψ)) are ordered upper and lower solutions of problem (4), respectively.

    Using Theorem 2.1 of [28] leads to the following existence result :

    Theorem 3.2. If RD0>1, problem (4) admits at least one coexistence solution (Ib(x),Im(x)) provided that β1 and β2 are sufficiently small.

    To establish the non-existence of the coexistence solution to problem (4), we have the following result.

    Theorem 3.3. If RD0(Ω,γb(x)11+(α1d1+β1γ1d1)Nb,dm(x)11+(α2d2+β2γ2d2)Am)1, problem (4) has no positive solution.

    Proof. Suppose (Ib(x),Im(x)) is a coexistence solution of problem (4), that is, (Ib(x),Im(x))>(0,0) in Ω and satisfies

    {Δ[(d1+α1Ib+β1Ibγ1+Im)Ib]=αb(x)βb(x)(NbIb)NbImγb(x)Ib,xΩ,Δ[(d2+β2Imγ2+Ib+α2Im)Im]=αm(x)βb(x)(AmIb)NbIbdm(x)Im,xΩ,Ib(x)=Im(x)=0,xΩ. (18)

    First by the upper and lower solution method we know that (Ib,Im)(Nb,Am).

    Second, letting w=(d1+α1Ib+β1Ibγ1+Im)Ib/d1 and z=(d2+β2Imγ2+Ib+α2Im)Im/d2, we have

    {d1Δw=αb(x)βb(x)(NbIb)Nbd2zd2+β2Imγ2+Ib+α2Imγb(x)d1wd1+α1Ib+β1Ibγ1+Im,xΩ,d2Δz=αm(x)βb(x)(AmIb)Nbd1wd1+α1Ib+β1Ibγ1+Imdm(x)d2zd2+β2Imγ2+Ib+α2Im,xΩ,w=Ib(x)=0,xΩ,z=Im(x)=0,xΩ, (19)

    which means that

    {d1Δw<αb(x)βb(x)zγb(x)1+(α1d1+β1γ1d1)Nbw,xΩ,d2Δz<αm(x)βb(x)AmNbwdm(x)1+(α2d2+β2γ2d2)Amz,xΩ,w=z=0,xΩ. (20)

    On the other hand, the principal eigenvalue λ0 in problem (10) meets

    {d1Δϕ=αb(x)βb(x)ψγb(x)1+(α1d1+β1γ1d1)Nbϕ+λ0ϕ,xΩ,d2Δψ=αm(x)βb(x)AmNbϕdm(x)1+(α2d2+β2γ2d2)Amψ+λ0ψ,xΩ,ϕ(x)=ψ(x)=0,xΩ. (21)

    Comparing (20) with (21), we can easily deduce from the monotonicity with respect to the coefficients in (21) that λ0 is monotone decreasing with respect to βb(x), which implies that λ0<0. Recalling Theorem 2.2 we can get that RD0>1, which is contrary to RD01.

    Remark 2. Assume that all coefficients of (4) are spatially-independent. RD0 is represented by (11). If αb,αm or βb is big, then RD0>1 and problem (4) admits at least one coexistence solution provided that β1 and β2 are sufficiently small. On the other hand, if αb,αm or βb is small enough, then RD01 and problem (4) has no positive solution.

    Next we apply the monotone iterative schemes to construct the true solutions of (4). It follows from RD0>1, we know that (M1,M2) and

    (g1(δd1ϕ,δd2ψ),g2(δd1ϕ,δd2ψ))

    are ordered upper and lower solution of problem (4), respectively. Using (ˉI(0)b,ˉI(0)m)=((1δ0)Nb,(1δ0)Am) and (I_(0)b,I_(0)m)=(g1(δd1ϕ,δd2ψ),g2(δd1ϕ,δd2ψ)) as two initial iterations, we can construct two sequences {(ˉu(n),ˉv(n))} and {(u_(n),v_(n))} from the iteration process

    {Δˉu(n)+k1ˉu(n)=F1(x,ˉI(n1)b,ˉI(n1)m),xΩ,Δˉv(n)+k2ˉv(n)=F2(x,ˉI(n1)b,ˉI(n1)m),xΩ,Δu_(n)+k1u_(n)=F1(x,I_(n1)b,I_(n1)m),xΩ,Δv_(n)+k2v_(n)=F2(x,I_(n1)b,I_(n1)m),xΩ,ˉI(n)b=g1(ˉu(n),ˉv(n)),I_(n)b=g1(u_(n),v_(n)),xΩ,ˉI(n)m=g2(ˉu(n),ˉv(n)),I_(n)m=g2(u_(n),v_(n)),xΩ,ˉu(n)(x)=u_(n)(x)=0,ˉv(n)(x)=v_(n)(x)=0,xΩ, (22)

    where n=1,2,.

    As in Lemma 3.1 of [28], the sequences {(ˉu(n),ˉv(n))} and {(u_(n),v_(n))} governed by (22) are well-defined and possess the monotone property

    (ˆu,ˆv)(u_(n1),v_(n1))(u_(n),v_(n))(ˉu(n),ˉv(n))
    (ˉu(n1),ˉv(n1))(˜u,˜v) for  n=1,2,.

    Hence, the pointwise limits

    limn(ˉu(n),ˉv(n)))=(ˉu,ˉv),limn(u_(n),u_(n))=(u_,v_)

    exist and their limits possess the relation

    (ˆu,ˆv)(u_(n),v_(n))(u_,v_)(ˉu,ˉv)(ˉu(n),ˉv(n))(˜u,˜v) (23)

    for every n=1,2,.

    The last three equations of (22) give

    ˉI(n)b=g1(ˉu(n),ˉv(n)),I_(n)b=g1(u_(n),v_(n)),
    ˉI(n)m=g2(ˉu(n),ˉv(n)),I_(n)m=g2(u_(n),v_(n)),

    which is equivalent to

    ˉu(n)=H1(ˉI(n)b,ˉI(n)m),u_(n)=H1(I_(n)b,I_(n)m),ˉv(n)=H2(ˉI(n)b,ˉI(n)m),v_(n)=H2(I_(n)b,I_(n)m). (24)

    Now, by the above relation, letting n and applying the standard regularity argument for elliptic boundary problems, we derive that (ˉIb,ˉIm) and (I_b,I_m) satisfy

    {Δ[H1(ˉIb,ˉIm)]+k1H1(ˉIb,ˉIm)=F1(x,ˉIb,ˉIm),xΩ,Δ[H2(ˉIb,ˉIm)]+k2H2(ˉIb,ˉIm)=F2(x,ˉIb,ˉIm),xΩ,Δ[H1(I_b,I_m)]+k1H1(I_b,I_m)=F1(x,I_b,I_m),xΩ,Δ[H2(I_b,I_m)]+k2H2(I_b,I_m)=F2(x,I_b,I_m),xΩ,ˉIb(x)=I_b(x)=0,ˉIm(x)=I_m(x)=0,xΩ, (25)

    which is equivalent to

    {Δ[H1(ˉIb,ˉIm)]=f1(x,ˉIb,ˉIm),xΩ,Δ[H2(ˉIb,ˉIm)]=f2(x,ˉIb,ˉIm),xΩ,Δ[H1(I_b,I_m)]=f1(x,I_b,I_m),xΩ,Δ[H2(I_b,I_m)]=f2(x,I_b,I_m),xΩ,ˉIb(x)=I_b(x)=0,ˉIm(x)=I_m(x)=0,xΩ. (26)

    Therefore (ˉIb,ˉIm) and (I_b,I_m) are true solutions of (4). Moreover, (ˉIb,ˉIm) and (I_b,I_m) are maximal and minimal solutions in the sense that (Ib,Im) is any other solution of (4) in the sector

    <(ˆIb,ˆIm),(˜Ib,˜Im)>:={(Ib,Im)C(¯Ω);(ˆIb,ˆIm)(Ib,Im)(˜Ib,˜Im)on¯Ω},

    then (I_b,I_m)(Ib,Im)(ˉIb,ˉIm) on ¯Ω. Furthermore, if ˉIb=I_b or ˉIm=I_m, then (ˉIb,ˉIm)=(I_b,I_m):=(Ib,Im) and (Ib,Im) is the unique solution of (4) in ¯Ω. To achieve this, in fact, a subtraction of the third equation from the first equation in (26) yields that

    Δ[β1(Ib)2(ˉImI_m)(γ1+ˉIm)(γ1+I_m)]=αb(x)βb(x)(NbIb)Nb(ˉImI_m)

    In light of β1>0, Ib>0, αb(x)βb(x)>0, NbIb>0 and (ˉImI_m)=0 on Ω, the above equation gives ˉImI_m in Ω. Similarly as above one can show that ˉIbI_b in Ω. Therefore, (ˉIb,ˉIm)=(I_b,I_m):=(Ib,Im) which is unique solution of (4) in S.

    The above conclusions lead to the following theorem.

    Theorem 3.4. Let (˜Ib,˜Im) and (ˆIb,ˆIm) be a pair of ordered upper and lower solutions of (4), respectively, then the sequences {(ˉI(n)b,ˉI(n)m)} and {(I_(n)b,I_(n)m)} provided from (22) converge monotonically from above to a maximal solution (ˉIb,ˉIm) and from below to a minimal solution (I_b,I_m) in S, respectively, and satisfy the relation

    (^Ib,^Im)(I_(n)b,I_(n)m)(I_(n+1)b,I_(n+1)m)(I_b,I_m)(ˉIb,ˉIm)(ˉI(n+1)b,ˉI(n+1)m)
    (ˉI(n)b,ˉI(n)m)(~Ib,~Im) for n=1,2,

    additionally, if ˉIb=I_b or ˉIm=I_m, then (ˉIb,ˉIm)=(I_b,I_m)(=(Ib,Im)) and (Ib,Im) is the unique solution of (4) in S.


    4. Numerical simulation and discussion

    In this section, in order to illustrate our theoretical results, we simulate problem (4) with the following coefficients and parameters:

    d1=0.2,d2=0.4,α1=0.03,α2=0.04,γ1=1,γ2=1,
    αb=1+0.88sin(π100x),αm=1+0.16sin(π100x),
    γb=1+0.6sin(π100x),dm=1+0.029sin(π100x),

    and βb=1+0.09sin(π100x) and we also take the ratio Am/Nb=20 as in [21].

    From Fig. 1, it is easy to see that there exist the upper solution sequence {(¯I(n)b,¯I(n)m)} which is monotone decreasing and the lower solution sequence {(I_(n)b,I_(n)m)} which is monotone increasing, then one can see that there exists at least a coexistence solution of (4).

    Figure 1. Phase diagrams of Ib(x) and Im(x) showing the existence of a positive solution of (4) for small cross-diffusion (β1=0.001 and β2=0.002).

    In this paper, to understand the impact of a cross-diffusion and environmental heterogeneity on the dynamics of WNv, we consider coexistence states of a cross-diffusive WNv model in heterogenous environments under Dirichlet boundary condition. This problem without spatially-dependent coefficients is similar to that has been studied in [16]. It is worth mentioning that problem (1) is weakly-coupled parabolic system which only involves the random diffusion in a homogeneous environment. However, in addition to the dispersive force, the diffusions of birds and mosquitoes are also interacted by each other and reaction depends on spatial heterogeneity of the environment. Therefore, we introduce the cross-diffusion terms Δ[(d1+α1Ib+β1Ibγ1+Im)Ib], Δ[(d2+β2Imγ2+Ib+α2Im)Im] to model (1), which can better describe the interplay between birds and mosquitoes in diffusion.

    The main result of this paper is twofold. Firstly, we introduce a definition of RD0, which is known as the basic reproduction number of problem (4) (Theorem 2.2). In the case that all coefficients are constants, we provide an explicit formula for RD0 (Theorem 2.3). Secondly, the coexistence of problem (4) is investigated by using method of upper and lower solutions and its associated monotone iterative schemes (Theorem 3.2 and Fig. 1) under condition RD0>1 provided that β1 and β2 are sufficiently small, whereas if RD01, problem (4) has no coexistence solution (Theorem 3.3). Our results show that no existence exists for small WNv transmission probabilities (αm and αb), and small biting rate of mosquitoes on birds (βb) (Remark 2). Moreover, the coexistence solution of problem (4) is between the maximal and minimal solution (ˉIb,ˉIm) and (I_b,I_m), respectively, and the true solution can be obtained by constructing the monotone iterative sequences (Theorem 3.4). However, the uniqueness of coexistence solution is still unclear.

    We believe that the strongly-coupled problem (4) can produce much more complex dynamics of WNv than the weakly-coupled system (1). Such problems need further investigations. In fact, even for the corresponding parabolic problems with cross-diffusion, the existence of the solution is known only for some special cases, see [24,32] and references therein. To further investigate the effect of cross-diffusion in comparison to no cross-diffusion or small cross-diffusion, we come back to problem (3), Fig. 2 shows that the global solution of problem (3) exists and stabilizes to a positive steady-state for small cross-diffusion (β1=0.132 and β2=0.11), we can also see that the global solution of (3) exists for β10.132 and β20.11 by simulations. However, if we choose a little big cross-diffusion, for example, β1=0.133 and β2=0.11, we can see from Fig. 3 that the global solution of problem (3) does not exist. We leave it for future work.

    Figure 2. Phase diagrams of Ib(x,t) and Im(x,t) shows that the solution of (3) exists and stabilizes to a positive steady-state for small cross-diffusion (β1=0.132 and β2=0.11).
    Figure 3. Phase diagrams of Ib(x) and Im(x) shows that the global solution of (3) does not exist for big cross-diffusion (β1=0.133 and β2=0.11).

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