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Analysis of a heterogeneous SEIRS patch model with asymmetric mobility kernel


  • In this paper, we establish a spatial heterogeneous SEIRS patch model with asymmetric mobility kernel. The basic reproduction ratio R0 is defined, and threshold-type results on global dynamics are investigated in terms of R0. In certain cases, the monotonicity of R0 with respect to the heterogeneous diffusion coefficients is established, but this is not true in all cases. Finally, when the diffusion rate of susceptible individuals approaches zero, the long-term behavior of the endemic equilibrium is explored. In contrast to most prior studies, which focused primarily on the mobility of susceptible and symptomatic infected individuals, our findings indicate the significance of the mobility of exposed and recovered persons in disease dynamics.

    Citation: Shuangshuang Yin, Jianhong Wu, Pengfei Song. Analysis of a heterogeneous SEIRS patch model with asymmetric mobility kernel[J]. Mathematical Biosciences and Engineering, 2023, 20(7): 13434-13456. doi: 10.3934/mbe.2023599

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  • In this paper, we establish a spatial heterogeneous SEIRS patch model with asymmetric mobility kernel. The basic reproduction ratio R0 is defined, and threshold-type results on global dynamics are investigated in terms of R0. In certain cases, the monotonicity of R0 with respect to the heterogeneous diffusion coefficients is established, but this is not true in all cases. Finally, when the diffusion rate of susceptible individuals approaches zero, the long-term behavior of the endemic equilibrium is explored. In contrast to most prior studies, which focused primarily on the mobility of susceptible and symptomatic infected individuals, our findings indicate the significance of the mobility of exposed and recovered persons in disease dynamics.



    There is mounting evidence in epidemiology that spatial-temporal heterogeneity and human mobility have a substantial impact on the spread of infectious diseases [1,2,3,4]. Different population, social contact rates, individual response, control measures, and medical resources (sickbeds and physicians per thousand people) at different regions all contribute to spatial heterogeneity. Many reaction diffusion or patch models have been presented in recent years to examine the role of diffusion and spatial heterogeneity in disease transmission [3,5,6,7,8,9,10,11,12,13,14,15]. Among these works, Allen et al. [5] developed a susceptible-infected-susceptible (SIS) patch system that collaborated with spatial heterogeneity in the following way:

    {d¯Sjdt=dSkΩ(Ljk¯SkLkj¯Sj)βj¯Sj¯Ij¯Sj+¯Ij+γj¯Ij,jΩ,d¯Ijdt=dIkΩ(Ljk¯IkLkj¯Ij)+βj¯Sj¯Ij¯Sj+¯Ijγj¯Ij,jΩ, (1.1)

    where Ω={1,2,3,,n} with n2 denoting the patch numbers, ¯Sj(t) and ¯Ij(t) represent the number of susceptible and infected individuals, respectively, in patch j at any given time t. The transmission rate of the disease in patch j is denoted by βj, while the recovery rate is denoted by γj. Furthermore, dS and dI are the diffusion rates of the susceptible and infected populations, respectively. Matrix (Ljk)n×n is symmetric with Ljk representing the degree of movement from patch k into patch j.

    In [5], a spatial SIS reaction-diffusion model is studied, with the focus on the existence, uniqueness and particularly the asymptotic profile of the steady states. More works related to system (1.1) can be seen in [6] (e.g., asymptotic behaviors of the endemic equilibrium (EE) as the diffusion rate of the infected individuals (dI) approaches to zero) and [16,17] (e.g., asymmetric matrix (Ljk)n×n).

    However, these models neglected the mobility of exposed persons and did not incorporate the class of exposed individuals. Exposed individuals, without exhibiting symptoms, can be seen in various epidemic diseases, such as malaria, HIV/AIDS, and SARS-COV-2. The disease can spread spatial if exposed individuals with no symptoms travel, making it more difficult to control [18,19]. As a result, it appears critical to incorporate the exposed individuals subclass and examine the effects of exposed individuals' mobility on infection disease spread, which is mathematically linked to the basic reproduction number's dependence on the diffusion rates of exposed individuals. Earlier studies have investigated discrete-space multi-patch models concerning this issue [20,21]. The exposed and recovery classes will be extended to the SIS patch model, and the associated SEIRS patch model will be analyzed in this study.

    We consider a system of differential equations:

    {dSdt=dSLSdiag{βiSiSi+Ei+Ii+Ri}I+αR,iΩ,t>0,dEdt=dELE+diag{βiSiSi+Ei+Ii+Ri}IσE,iΩ,t>0,dIdt=dILI+σEdiag{γi}I,iΩ,t>0,dRdt=dRLR+diag{γi}IαR,iΩ,t>0, (1.2)

    where S, E, I, and R represent the number of individuals in the susceptible, exposed, infectious, and recovered compartments, respectively. The parameters β, σ, γ, and α denote the rates of transmission, latent period, recovery, and loss of immunity, respectively.

    Here Ω={1,2,3,,n} with n2 denoting the patch numbers, Si(t), Ei(t), Ii(t) and Ri(t) denote the population of susceptible, exposed, infected and recovered individuals at time t and patch i, respectively. Furthermore, dS, dE, dI and dR represent the diffusion coefficients associated with susceptible, exposed, infected and recovered individuals, respectively. We assume that β:=(βi)n×1 and γ:=(γi)n×1 depends on the environment, could be spatial heterogeneous and are assumed to be positive throughout this paper. Given that the latent period 1/σ which represents the time takes for an infection to become detectable after exposure and the rate of loss of immunity α which describes the proportion of individuals who lose their immunity to a disease over time, are inherent properties of the individual and not influenced by the external environment, it is reasonable to consider σ and α as constants in this study.

    L=(ij) is an n×n cooperative, irreducible matrix with ii=jiij referred as Laplacian matrix, where ij, denoting the degree of movement from patch k into patch j, is independent of t. Here an n×n matrix is said to be cooperative if all of its off-diagonal entries are non-negative, and to be irreducible if its index set {1,2,,n} cannot be split into two complementary sets (without common indices) {p1,p2,,pr} and {q1,q2,,qs} (r+s=n) such that apiqj=0 for all 1ir,1js. An n×n matrix is called quasi-positive (also called Metzler matrix) if lij0 for all ij. Here we mention that L can be asymmetric. Moreover, to match the real human mobility patterns and transportation phenomena such as gravity [22,23] or radiation mobility model [24], we assume that L can be asymmetric. We also assume that the following initial conditions are satisfied throughout the study.

    (A1) Si(0),Ei(0),Ii(0),Ri(0)0 for iΩ and ni=1Ii(0)>0.

    It is easy to verify that βiSiIiSi+Ii+Ei+Ri is a Lipschitz continuous function of Si and Ii, therefore we define the quantity to be zero in cases where Si=0 or Ii=0. According to [34, Chapter I], it can be stated that the system (1.2) has a unique classical solution S,E,I,RC1((0,);Rn), and Si(t),Ei(t),Ii(t), Ri(t)>0 for iΩ and t>0.

    The total population at time t is defined as

    N(t)=ni=1(Si(t)+Ei(t)+Ii(t)+Ri(t)),

    and let the total number of individuals is a fixed positive constant at the initial time t=0, denoted by N0. By the system (1.2), we conclude that the total population size is constant, i.e.,

    N(t)=N0  for any  t0. (1.3)

    Let u,vRn. We write uv if uivi for any iΩ; u>v if uivi for any iΩ, and there exists j such that uj>vj; uv if ui>vi for any iΩ. We say u is non-negative, positive and strongly positive if u0,u>0,u0, respectively.

    Our primary focus will be on equilibrium solutions of the system (1.2), i.e., non-negative solutions of the following system:

    {dSL˜Sdiag{βi˜Si˜Si+˜Ii+˜Ei+˜Ri}˜I+α˜R=0,dEL˜E+diag{βi˜Si˜Si+˜Ii+˜Ei+˜Ri}˜Iσ˜E=0,dIL˜I+σ˜Ediag{γi}˜I=0,dRL˜R+diag{γi}˜Iα˜R=0,ni=1(˜Si+˜Ei+˜Ii+˜Ri)=N0, (1.4)

    where the population of susceptible, exposed, infected and recovered individuals at equilibrium are represented by the variables ˜S, ˜E, ˜I, and ˜R, respectively. A solution of (1.4) in which ˜Ii=0 for every iΩ is a disease-free equilibrium (DFE), while a solution in which ˜Ii>0 for some iΩ is referred to as an endemic equilibrium (EE). It is straightforward to observe that DFE is unique, given by E0=(˜N,0,0,0), where ˜N is the unique positive eigenvector corresponds to the eigenvalue of 0 in the following eigenvalue problem:

    Lϕ=0,ni=1ϕ=N0. (1.5)

    Moreover, for any EE, ˜S,˜E,˜I,˜R0.

    The purpose of this research is to examine how changes in human mobility and environment heterogeneity affect the persistence or eradication of infectious diseases. Our research will investigate the threshold-type dynamics of system (1.2), the characteristics of the basic production ratio, and when the diffusion rate of the susceptible individuals approaches zero, the asymptotic behaviors of the endemic equilibria of (1.4).

    The basic reproduction number is a fundamental concept in the study of infectious disease transmission. It is defined as the expected number of secondary infections generated by a single infected individual in a completely susceptible population [25,26]. More importantly, it is frequently used to determine the behaviour of various epidemic models' thresholds. Specifically, when the basic reproduction number is less than unity, the disease is expected to fade out, while a basic reproduction number greater than unity indicates the potential for the disease to establish itself within the population. The next-generation operator method has been widely utilized for the computation of fundamental reproduction number, see [25,27,28] and for related investigations, see [3,28,29].

    By [28,29], the basic reproduction number, denoted as R0, can be expressed as R0=1μ0 (see Lemma 2). Here, μ0 represents the unique positive eigenvalue associated with a positive eigenvector of the following problem:

    {dELφE+σφE=μdiag{βi}φI,iΩ,dILφI+diag{γi}φIσφE=0,iΩ. (1.6)

    In terms of R0, we state the dynamics for system (1.2) as follows:

    Theorem 1.1. (i) If R01, then DFE is globally asymptotically stable, that is E(t),I(t),R(t)0 as t, and S˜N as t;

    (ii) If R0>1, then there exists a positive constant ϵ0 such that any positive solution of (1.2) satisfies

    lim inft(S(t),E(t),I(t),R(t))(˜N,0,0,0)>ϵ0. (1.7)

    Furthermore, system described by (1.2) has at least one endemic equilibrium.

    It is not hard to see that the value of the basic reproduction number gives an indication on the persistence or extinction of infectious diseases. Therefore, we focus on the dependence of R0 on dE,dI, dS and dR. It is easy to know that R0 is independent of dS and dR. The dependence of R0 on dE,dI can have important implications for disease control because it explores how changes in human mobility affect the persistence or extinction of infectious diseases. [16] demonstrates that R0 for model (1.1) is decreasing in dI. However, for the SEIRS system (1.2), the monotonicity of R0 with respect to dI is more subtle due to the presence of mobility of exposed individuals.

    To start with, we give the asymptotic properties of R0 when dE,dI tend to 0 or infinity in Theorem 1.2. Define β=(βi)n×1,γ=(γi)n×1 and set

    B=diag{βi},Y=diag{γi},Z=diag{βi/γi}.

    Moreover, ξ is defined as the positive eigenvector of L that is unique, satisfying iΩξi=1.

    Theorem 1.2. (i) Fix dI>0. Then R0ρ((dIL+Y)1B) as dE0, and R0B(dIL+Y)1ξ1ξ1 as dE.

    (ii) Fix dE>0. Then R0ρ(σ(dEL+σIn×n)1Z) as dI0 and R0ni=1ξiβini=1ξiγi as dI.

    (iii) As dE,dI0, then R0max{βiγi,iΩ};

    (iv) As dE and dI0, then R0ni=1ξiβiγi.

    The following result is concerned with the monotonicity of R0 with respect to dE,dI:

    Theorem 1.3. If L is symmetric and either β or γ is a multiple of vector 1, then R0 exhibits a monotone decreasing behaviour with respect to the parameters dE and dI. Furthermore, strict monotonicity is observed if and only if one of them is not multiple of vector 1.

    If both β and γ are not multiples of vector 1, Theorem 1.3 may be not valid. Refer to Theorem 3.3.

    Theorem 1.4. Let ij=1,ij,ii=(n1) and

    sign{βiβj}=sign{γjγi} (1.8)

    holds for any i,jΩ. Then R0 is nonincreasing in dE,dI.

    Under the conditions in Theorems 1.3 and 1.4, the nonincreasing behaviour of R0 with respect to dE and dI seems counter-intuitive. If no traffic control or transportation restrictions policies are adopted during the disease spread, large diffusion rate also means accessing more health resources (β is a multiple of vector 1; condition (1.8)) or escaping from high risk areas (γ is a multiple of vector 1; condition (1.8)).

    Finally, to better understand the influence of the suspected individuals movement on the spread of disease, we demonstrate that the asymptotic behavior of the endemic equilibrium as the diffusion rate of susceptible individuals approaches zero. By Theorem 1.1, when R0>1, there exist at least one endemic equilibrium. To this end, let's examine this linear eigenvalue problem

    dRLϕ+αdiag{1γiβi}ϕ=λϕ. (1.9)

    And here represent the minimum eigenvalue of (1.9) as Λ1(dRL+αdiag{1γiβi}) ([33, Theorems 1 and 2]).

    Theorem 1.5. Assume R0>1 and Λ1(dRL+αdiag{1γiβi})<0. Then

    (i) There exist positive constants C1,C2, independent of dS, such that for sufficiently small dS,

    C1˜EidS,˜IidS,˜RidSC2,iΩ;

    (ii) As dS approaches zero, with respect to a given sequence,

    ˜S˜S=N0(ξM)ni=1(ξM),

    where MRn satisfying 0Mξ and |{iΩ:Mi=ξi}|(0,n).

    In particular, Theorem 1.5 demonstrates that as dS approaches zero, the variables ˜E, ˜I, and ˜R converge uniformly to zero in Ω. Additionally, ˜S is positive and there exists a non-empty subset {iΩ:˜Si=0}. From a biological standpoint, limiting the diffusion rate of susceptible individuals can effectively reduce the number of exposed and infected individuals in a population. We point out that if Λ1(dRLαdiag{1γiβi})>0, Theorem 1.5 may be not valid.

    Most of the results of SEIRS patch model (1.2) match the contents of SEIRS reaction diffusion model in [19], which deals with the continuous-space case to the discrete case. Compared with [19], the main improvement of this paper is as follows: investigation of threshold dynamics and asymptotic behaviours of R0 with respect to dE and dI are extended to asymmetric mobility kernel such as gravity [22,23] or radiation mobility model [24]. In [19, Theorem 1.2], under the condition that γ is a constant function, the proof of the monotonicity of R0 with respect to dE,dI is complicated and difficult for readers to access the core ideas. For SEIRS patch model in this paper, the proof is shortened (see Theorem 3.2) and can give more intuitiveness to understand the proof of [19, Theorem 1.2]. Moreover, though the results in Theorem 1.4 is similar in spirit to [19, Theorem 1.3], the proof for patch model is novel and totally different from continuous-space case.

    This paper is organized as follows. In Section 2, we establish the well-posedness of the system (1.2), introduce the basic reproduction number R0, and analyze the system's dynamics in terms of R0. Section 3 is dedicated to exploring the asymptotic stability and directional monotonicity of R0 in relation to dE and dI. Finally, Section 4 focuses on the behavior of the endemic equilibrium as dS approaches zero.

    To start with, we have the following uniform bound result, which can be directly derived from (1.3).

    Lemma 2.1. For some positive constants C1, which are independent of initial values, and T>0, the solution (S,E,I,R)C1((0,);R4n) of the system (1.2) satisfies

    |Si(t)|+|Ei(t)|+|Ii(t)|+|Ri(t)|C1,iΩ,t>T. (2.1)

    We now make use of the theory developed in [28] to derive R0 of system (1.2).

    Lemma 2.2. (i). R0=ρ(FV1)=ρ(σB(dEL+σIn×n)1(dIL+Y)1).

    (ii). The eigenvalue problem (1.6) has a positive eigenvalue that is unique, denoted by μ0, along with a corresponding positive eigenvector. Additionally, the basic reproduction number of system (1.2), denoted by R0, satisfies

    R0=1μ0. (2.2)

    Proof. (i) The linearization of system (1.2) at E0 is given by

    {dˉSdt=dSLˉSBˉI+αˉR,t>0,dˉEdt=dELˉE+BˉIσˉE,t>0,dˉIdt=dILˉI+σˉEYˉI,t>0,dˉRdt=dRLˉR+YˉIαˉR,t>0. (2.3)

    Note that E and I represent the infected population in (1.2). Furthermore, here F and V in [28] is denoted as

    F=(0B00)2n×2n,V=L+(σIn×n0σIn×nY), (2.4)

    where L=diag{dEL,dIL}. Part (i) follow immediately from [28]. We only need to prove part (ii). By Perron-Frobenius theorem [33, Theorems 1 and 2], it is possible to establish the existence and uniqueness of a positive eigenvalue μ0 with respect to a positive eigenvector φ within the system described in Eq (1.6), i.e.,

    Vφ=μ0Fφ.

    Note that there exists a positive eigenvector ϕ such that

    FV1ϕ=R0ϕ,

    which implies

    Vϕ=1R0Fϕ.

    By the uniqueness of μ0, we obtain R0=1μ0. This completes the proof.

    We now demonstrate that the stability of E0 can be determined by the basic reproduction number R0.

    Lemma 2.3. The disease-free equilibrium E0 is locally asymptotically stable if R0<1, unstable if R0>1.

    Proof. The proof of Lemma 2.3 can follow from [28]. For readers' convenience, we also give some details here. To prove the locally asymptotically stability if R0<1, we only need to prove inf{Reλ, λΛ}>0, where Λ is the spectrum of the following eigenvalue problem

    {dSLϕSBϕI+αϕR+λϕS=0,dELϕE+BϕIσϕE+λϕE=0,dILϕI+σϕEYϕI+λϕI=0,dRLϕR+YϕIαϕR+λϕR=0. (2.5)

    Note that Λ=Λ{(ϕE,ϕI)0}Λ{(ϕE,ϕI)=0}. Obviously, inf{Reλ,λΛ{(ϕE,ϕI)=0}}>0. Moreover, it can be shown that Λ(ϕE,ϕI)0 is a subset of the set of σ(LF+V), where L,F,V are defined in (2.4) and σ(LF+V) denotes the eigenvalues of the matrix LF+V. The infimum of the real parts of the eigenvalues of LF+V is positive, here denoted by λ1, i.e., inf{Reλ,λσ(LF+V)}=λ1>0. Moreover, λ1 represents the principal eigenvalue of

    {dELϕEBϕI+σϕE=λϕE,dILϕI+YϕIσϕE=λϕI. (2.6)

    Consider the left eigenvector corresponding to principal eigenvalue of (2.6), i.e.,

    {dELTϕE+σϕEσϕI=λ1ϕE,dILTϕIBϕE+YϕI=λ1ϕI. (2.7)

    Multiply (1.6)'s first equation by (ϕE)T and (2.7)'s first equation by φTE. Subtracting the resulting equations yields

    λ1φTEϕE=1R0φTIBϕEσ(ϕI)TφE. (2.8)

    Moreover, multiply (1.6)'s second equation by (ϕI)T and (2.7)'s second equation by φTI. Subtracting the resulting equations yields

    λ1φTIϕI=φTIBϕEσ(ϕI)TφE. (2.9)

    Adding two Eqs (2.8) and (2.9) yields

    λ1(φTEϕE+φTIϕI)=1R0R0φTIBϕE.

    Then we have sign(1R0)=sign(λ1) for the positive of φE,ϕE,φI,ϕI,β. We obtain inf{Reλ,λΛ{(ϕE,ϕI)0}}>0. Therefore, if R0<1, E0 is locally asymptotically stable.

    To establish the linear instability of E0 when R0>1, it is necessary to demonstrate the existence of a non-trivial solution to (2.5) with Reλ<0. To this end, let λ=λ1<0, where λ1 denotes the principal eigenvalue of (2.6) and select (ϕE,ϕI) as the eigenvector of (2.6) associated with λ1 and solving for ϕS and ϕR in (2.5). Therefore, E0 is unstable if R0>1.

    We now demonstrate that the disease will be extinct if R0<1, i.e., the E0 is globally asymptotically stable, and we show that when R0>1 at least one EE exists and that both ˜S and ˜E are positive.

    Proof of Theorem 1.1. (i). In order to analyze the behavior of the system (1.2), we make use of LaSalle's invariance principle (Theorem 1 in [30]) and construct Lyapunov function. We consider the ordered space X=R4n equipped with the supremum norm, and observe that X has a nonempty interior, which we denote by int(P), where the cone P is composed of all functions in X that are nonnegative. Set

    X0={u=(us,ue,ui,ur)X|jΩ(us,j+ue,j+ui,j+ur,j)=N0}

    and U=PX0. It can be shown that system (1.2) and (1.3) defines a dynamic system on U. Moreover, for any initial condition (s0,e0,i0,r0)U, the unique solution of the system is denoted by Φt(s0,e0,i0,r0)=(S,E,I,R) for any t>0. It's worth noting that Φt is compact, and for each u0U, the orbit of u0 under the dynamical system generated by (1.2) has compact closure in U.

    Define

    L(u)=uTeϕE+uTiϕI

    for uU, where (ϕE,ϕI) is the eigenvector corresponding to the principal eigenvalue λ1 of (2.7), i.e., left eigenvector of (2.6). Next, we show that L(u) serves as a Lyapunov function for (1.2). Considering an arbitrary solution of the system (1.2) coupled with (1.3), we obtain:

    ddtL(u(t))=(ϕE)TEt+(ϕI)TIt=(ϕE)T(dELE+diag{βiSiSi+Ii+Ei+Ri}IσE)+(ϕI)T(dILI+σEγI)=ITdiag{βi(Ei+Ii+Ri)Si+Ii+Ei+Ri}ϕEλ1(ETϕE+ITϕI). (2.10)

    Following the same process as in the proof of Lemma 2.3, R01 yields that λ10. Furthermore, S,E,I,R0, and β,ϕE,ϕI>0. Consequently, we can assert that ddtL(u(t))0, which indicates that L(u) functions as a Lyapunov function for the system (1.2).

    Next define

    ˙L(u0):=ddtL(u(t))|t=0  and   M={u0U|˙L(u0)=0},

    where u is the unique solution of (1.2) with u0=(s0,e0,i0,r0)U as initial condition. By (2.10), we have M={u0=(s0,e0,i0,r0)U|i0=0} if λ1=0, and M={u0=(s0,e0,i0,r0)U|e0=i0=0} if λ1>0. Using (1.2), we can infer that when λ10, the maximal invariant set in M can be characterized by:

    ˆM:={u0=(s0,e0,i0,r0)U|e0=i0=0}.

    Consequently, we can utilize the LaSalle invariant principle (stated in Theorem 1 in [30]) to obtain:

    (E(t),I(t))(0,0),  as  t.

    By combining the above equation with (1.2), we can infer that R(t)0 as t. Consequently, utilizing (1.3), we obtain ni=1S(t)˜N as t, where ˜N is defined in (1.5).

    We omit the proof of Theorem 1.1 (ii) because it is standard.

    In terms of the basic reproduction number R0, we established the threshold dynamics of system (1.2) in the previous section. In this part, we will examine the asymptotic characteristics and monotonicity of the basic reproduction number in relation to the parameters dE and dI. The aim is to explore the influence of population movement on the long-term behavior of infectious diseases, particularly persistence or eradication. In this section, we will establish the proofs of Theorems 1.2 and 1.4. Theorem 1.3 is a consequence of Theorems 3.1 and 3.2.

    For further conveniences, we give some preliminary results here. To start with, we give some definitions. Let u,vRn. We define the relation between u and v as follows: uv if uivi for iΩ; u>v if uivi for iΩ, and j such that uj>vj; uv if ui>vi for iΩ. We say that u is non-negative, positive and strongly positive if u0,u>0,u0, respectively. Moreover, we define the vector norm

    u1=ni=1|ui|,u=max1in{|ui|}.

    Consider an n×n matrix, denoted as A, with the set of eigenvalues σ(A). Let r(A) to be the spectral radius of A, i.e.,

    r(A)=max{|λ|:λσ(A)}.

    Denote s(A) be the spectral bound of A, i.e.,

    s(A)=max{Reλ:λσ(A)}.

    Recall that a principal eigenvalue of the matrix A means a real and simple eigenvalue with positive eigenvectors.

    Now we have the following preliminary results.

    Lemma 3.1. L admits a unique principal eigenvalue 0 with a unique positive eigenvector ξ satisfying iΩξi=1. Moreover, for any vector φ, we have φTLφ0.

    Throughout this paper, we define ξ to be the positive eigenvector that is unique satisfying iΩξi=1.

    Lemma 3.2 ([17]). Consider an n×n quasi-positive irreducible matrix, denoted as A=(aij)n×n. Let Q be a diagonal matrix, denoted as Q=diag{qi}. Then, we have the subsequent results:

    (i). If s(A)<0, then s(μA+Q) is strictly decreasing in μR+. Furthermore,

    limμ0s(μA+Q)=max{qi}

    and

    limμs(μA+Q)=;

    (ii). If s(A)=0, then s(μA+Q) is strictly decreasing provided that Q is not a multiple of In×n. Furthermore,

    limμ0s(μA+Q)=max{qi}

    and

    limμs(μA+Q)=iΩqiξi,

    where ξ refers to the positive eigenvector of s(A) that is unique and satisfies the condition iΩξi=1. (It is worth noting that if every row of matrix A has a sum of zero, then ξ denotes the left positive eigenvector of A.)

    Lemma 3.3. For any dE>0,dI>0,

    min{βiγi,iΩ}R0max{βiγi,iΩ}. (3.1)

    Proof. According to Lemma 2.2, the reciprocal of the basic reproduction number, denoted as 1R0, represents principal eigenvalue that is unique of the eigenvalue problem expressed in Eq (1.6). Therefore, it follows that:

    {dELφE+σφE=1R0BφI,dILφI+YφIσφE=0,. (3.2)

    It follows from the addition of two equations of (3.2) that

    dELφEdILφI+YφI=1R0BφI, (3.3)

    which yields

    ni=1γi(R0βiγi)φI,i=0.

    Since γi and φI are positive, we obtain (3.1).

    Lemma 3.3 showed an estimate of R0 and implied that if βiγiConstant, then R0 is constant, which is independent of dE,dI.

    Proof of Theorem 1.2. We only need to prove (i) here because (ii) can be shown through similar arguments. The case dE0 can be derived by the continuity of eigenvalue with respect to dE [32, Proposition 2.1]. Now we explore the case dE. By Lemma 3.3, if required, we can pass to a sequence such that R0˜R0 as dE. Without loss of generality, we may assume φE+φI=1. Passing to a sequence if necessary, φE˜φE,φI˜φI in Rn+ as dE. As shown in Lemma 3.2, ˜φE is a multiple of ξ and ˜φI is a solution of a certain equation

    dIL˜φI+Y˜φIσ˜φE=0.

    Summarizing the first equation of (3.2) yields that R0B(dIL+Y)1ξ1ξ1. This ends the proof of (i).

    (iii) The statement can be derived by the continuity of eigenvalue with respect to dE [32, Proposition 2.1].

    (iv) The statement can be directly deduced from Lemma 3.2 and statement (ii).

    Now, we show some cases that R0 has monotonicity with respect to dE,dI results as follows.

    Theorem 3.1. If β is a multiple of vector 1 and L is symmetric, then R0 is monotone decreasing in dE,dI. Moreover, the strict monotonicity holds if γi is not a multiple of vector 1.

    Proof. Firstly, we demonstrate that R0 is monotone decreasing with respect to dE. By [32, Proposition 2.1], we know that R0 and the corresponding eigenvectors (φE,φI),(φE,φI) are both differentiable functions of dE,dI. By direct differentiating both sides of the equations in (3.2) by dE,dI, we obtain

    {dELφELφE+σφE=1R0BφIR0R20BφI,dILφI+YφIσφE=0, (3.4)

    and

    {dELφE+σφE=1R0BφIR0R20BφI,dILφILφI+YφIσφE=0, (3.5)

    respectively. Since no confusion occurs in subsequent proofs, here the prime notation is utilized to signify differentiation in relation to either E or I, for the purpose of convenience. The initial equation of (3.2) is multiplied by φE and the initial equation of (3.4) is multiplied by φE. The resulting equations are then subtracted to obtain the desired outcome,

    R0R20φTIBφE=φTELφE+1R0(φ,TIBφEφTIBφE).

    Analogously, the second equation of (3.2) is multiplied by φI and the second equation of (3.4) is multiplied by φI. The resulting equations are then subtracted to obtain the desired outcome, (It should be noted that, since σ is constant, it can be treated as a constant throughout the procedure)

    (φ,TIφEφTIφE)=0.

    If β is a multiple of vector 1, we can see that

    βR0R20φTIφE=φTELφE.

    By Lemma 3.1, φTELφE0. We obtain R00 as φI,φE are positive. In addition, the equality is only possible if φE is a multiple of vector 1. This fact together with the first equation of (3.2) leads to the conclusion that φI must be a multiple of vector 1. The obtained expression, along with the second equation of (3.2), implies that the parameter γ must be a multiple of the vector 1. Consequently, it follows that the quantity R0 exhibits a monotone decreasing behavior with respect to the variable dE, with strict monotonicity being guaranteed only if the parameter γ is not a multiple of vector 1.

    Next we show the monotonicity of R0 with respect to dI. The initial equation of (3.2) is multiplied by φE, and initial equation of (3.5) is multiplied by φE. The resulting equations are subtracted to obtain the desired outcome,

    R0R20φTIBφE=φ,TIBφEφTIBφE.

    Analogously, the second equation of (3.2) is multiplied by φI, and the second equation of (3.5) is multiplied by φI. The resulting equations are subtracted to obtain the desired outcome,

    φTILφI+σ(φ,TIφEφTIφE)=0.

    If β is a multiple of vector 1. We obtain

    σR0R20φTIφE=φTILφI.

    Similar to the previously mentioned arguments, it can be stated that the quantity R0 is exactly monotonically decreasing in behaviour with respect to the variable dI if and only if the parameter γ is not a multiple of vector 1.

    Theorem 3.2. If γ is a multiple of vector 1 and L is symmetric, then R0 is monotone decreasing function of dE,dI Furthermore, it is only when the parameter β is not a multiple of vector 1 that the monotonicity of R0 becomes strictly decreasing.

    Proof. Note from Lemma 2.2 that R0=ρ(σB(dEL+σIn×n)1(dIL+Y)1). Let γ=y1. Then R0 satisfies

    (dEL+σIn×n)(dIL+yIn×n)φ=1R0σBφ, (3.6)

    where φ is a strongly positive eigenvector.

    We only prove that R0 is monotone decreasing function of dE and the proof of monotonocity with respect to dI is similar. Differentiating (3.6) by dE yields that

    (dEL+σIn×n)(dIL+yIn×n)φL(dIL+yIn×n)φ=1R0σBφR0R20σBφ. (3.7)

    Multiplying (3.6) by φT and (3.7) by φ,T, and subtracting the resulting equations yield that

    R0R20σφTBφ=dI(Lφ)TLφ+φTLφ0.

    Therefore, R0 is monotone decreasing function of dE. Applying analogous arguments to that utilized in the proof of Theorem 3.1, it can be asserted that the quantity R0 is exactly monotonically decreasing behaviour with respect to dE if and only if the parameter β is not a multiple of vector 1.

    Now we prove Theorem 1.4.

    Proof of Theorem 1.4. Note that the sign conditions mean that for any i,jΩ, βiβj(γiγj) or βiβj(γiγj) holds true. We divide the proof of Theorem 1.4 into two steps.

    Step 1. First we show that in (1.6),

    sign{βiβj}=sign{φE,iφE,j}=sign{φI,iφI,j} (3.8)

    holds for any i,jΩ. Fix i,j, set ρE=φE,iφE,j,ρI=φI,iφI,j, and without loss generality, let βjβi,γjγi. Note from (1.6) that

    {(n1)dEφE,idEkiφE,k+σφE,i=μ0βiφI,i,(n1)dEφE,jdEkjφE,k+σφE,j=μ0βjφI,j,(n1)dIφI,idIkiφI,k+γiφI,iσφE,i=0,(n1)dIφI,jdIkjφI,k+γjφI,jσφE,j=0.

    Subtracting the last and first two equations yields that

    {σρE(ndI+γi)ρI=(γiγj)φI,j,(ndE+σ)ρE+μβiρI=μ(βjβi)φI,j. (3.9)

    Moreover, it follows from (1.6) that

    {σφE,i(ndI+γi)φI,i=dIjΩφI,i,(ndE+σ)φE,i+μβiφI,i=dEjΩφE,j. (3.10)

    Denote h0=[(γiγj)φI,j,μ(βjβi)φI,j]T,h1=[dIjΩφI,j,dEjΩφE,j]T, h2=[φE,i,φI,i]T and

    M=(σ(ndI+γi)(ndE+σ)μβi).

    Thus, (3.10) can be rewritten as Mh2=h10. By [31, Fact 6.11.13 (xii)], M is M-matrix and M1 is a positive matrix. Here a matrix A=(aij)n×n is called an M-matrix if aij0 for all ij and A=sIB with B having all off-diagonal elements negative and sr(B). Therefore, [ρE,ρI]T=M1h00, which implies (3.8).

    By similar arguments, we can also obtain that

    sign{βiβj}=sign{φE,iφE,j}=sign{φI,iφI,j} (3.11)

    Step 2. Moreover, the reciprocal of the basic reproduction number, denoted as 1R0, represents the principal eigenvalue that is unique of the eigenvalue problem expressed in Eq (1.6). Using the same notations F,V of (2.4), (1.6) can be written as

    Vφ=1R0Fφ,

    where φ=(φE,φI). Thus the adjoint problem of (1.6) can be written as

    VTφ=1R0FTφ,

    i.e.,

    {dELTφE+σφEσφI=0,dILTφI+YφI=1R0BφE, (3.12)

    where φ=(φE,φI) corresponds to a positive eigenvector. We can derive the following equation by multiplying the first equation of (3.4) by the transpose of φE, denoted by φ,TE, and multiplying the first equation of (3.12) by the transpose of the derivative of φE, denoted by (φE)T, then subtracting the resulting equations:

    R0R20φTIBφE=φ,TELφE+1R0(φI)TBφEσφ,TIφE. (3.13)

    The second equation of (3.4) is multiplied by φI, and the second equation of (3.12) is multiplied by φI, which yields

    1R0(φI)TBφEσφ,TIφE=0. (3.14)

    As a consequence of (3.13) and (3.14), we have

    R0R20φTIBφE=φ,TELφE=nni=1φE,iφE,i+ni,j=1φE,jφE,i=12(nni=1φE,iφE,i+nnj=1φE,jφE,jni,j=1φE,iφE,jni,j=1φE,jφE,i)=12ni,j=1(φE,iφE,j)(φE,iφE,j)0. (3.15)

    This combined with (3.15) suggests that R00. By similar arguments as the proof of Theorem 3.1, if and only if both the parameters β and γ are constant vectors the equality holds.

    We then prove that R0 decreases monotonically with respect to dI. Using similar arguments as before, we demonstrate that,

    R0R20φTIBφE=φ,TILφI=12ni,j=1(φI,iφI,j)(φI,iφI,j)0.

    and the remaining arguments are analogous to the previous ones.

    In previous subsections, we show that R0 is monotone decreasing in dE,dI in some cases. In this part, we will prove that R0 isn't always monotone decreasing with respect with dE,dI.

    Theorem 3.3. There exist d0E and d1I<d2I such that R0(d0E,d1I)<R0(d0E,d2I), if

    ni=1ξiβini=1ξiγi>ni=1ξiβiγi.

    Proof. Based on Theorem 1.2, it can be shown that for any fixed value of dE>0, R0ni=1ξiβini=1ξiγi as dI, and R0ni=1ξiβiγi as dI0,dE. For any arbitrarily small positive value of ϵ, there exists a sufficiently large constant C1(ϵ) such that for any 1d1I,dEC1(ϵ), we have

    R0(dE,d1I)(1+ϵ)ni=1ξiβiγi.

    Furthermore, there exists C2(ϵ,dE) such that for any d2IC2(ϵ,dE),

    R0(dE,d2I)(1ϵ)ni=1ξiβini=1ξiγi.

    Since

    ni=1ξiβini=1ξiγi>ni=1ξiβiγi,

    we can choose ϵ0 small enough such that

    (1ϵ0)ni=1ξiβini=1ξiγi>(1+ϵ0)ξiβiγi,

    and let d0E=C1(ϵ0),d1I=1C1(ϵ0),d2I=C2(ϵ0,d0E), we have R0(d0E,d1I)<R0(d0E,d2I).

    Theorem 3.4. Let ν0=ni=1ξiγini=1ξiβi and φ1,ϕ1 be the unique solutions of

    Lφ1=ν0BξYξ

    and

    Lϕ1=ν0(ni=1βiξi)ξYξ,

    respectively. If

    (γν0β)T(φ1ϕ1)>0, (3.16)

    there exist d0I and d1E<d2E such that R0(d1E,d0I)<R0(d2E,d0I).

    Proof. Consider the principal eigenvalues of the following two eigenvalue problems, which referred to as μ and ν, respectively,

    dILφ+Yφ=μBφ, (3.17)

    and

    dILϕ+Yϕ=ν(ni=1βiϕi)ξ (3.18)

    with ni=1φi=ni=1ϕi=1. Note from Theorem 1.2 that for any fixed dI>0, R01μ as dE0 and R01ν as dE. Now we take ϵ=1dI and perform regular expansions on (ϕ,ν) and (φ,μ) to obtain the following expressions.

    φi=φ0,i+ϵφ1,i+ϵ2φ2,i(ϵ),ϕi=ϕ0,i+ϵϕ1,i+ϵ2ϕ2,i(ϵ),μ=μ0+ϵμ1+ϵ2μ2(ϵ),ν=ν0+ϵν1+ϵ2ν2(ϵ). (3.19)

    Our objective is to establish that the inequality μ>ν holds true for small values of ϵ subject to the condition (3.16). Upon performing direct computation, we obtain the following expressions, φ0=ϕ0=ξ, μ0=ν0=ni=1ξiγini=1ξiβi and φ1,ϕ1 satisfy

    Lφ1=ν0BξYξ

    and

    Lϕ1=ν0(ni=1βiξi)ξYξ,

    respectively. Furthermore, we have

    ni=1γiφ1,i=μ0ni=1βiφ1,i+μ1ni=1βiξi (3.20)

    and

    ni=1γiϕ1,i=ν0ni=1βiϕ1,i+ν1ni=1βiξi. (3.21)

    Therefore, by condition (3.16), (3.20), (3.21) and μ0=ν0, we obtain

    (μ1ν1)ni=1βiξi=ni=1(γiν0,iβ)(φ1,iϕ1,i)>0.

    Thus μ>ν for large dI. Therefore, we can find d0I large, and d2E large, d1E small such that R0(d1E,d0I)<R0(d2E,d0I).

    Remark: We present an example in which condition (3.16) holds. Let Ω={1,2}, β1=1.01,β2=1.424 and γ1=1.01,γ2=2.01, by direct computation, we can see

    ni=1(γiν0βi)(φ1,iϕ1,i)=0.0043>0.

    Theorem 1.5 is a consequence of Lemma 4.1, Theorems 4.1 and 4.2. To analyze the behavior of endemic equilibria as dS tends towards zero, we shall examine the asymptotic properties of the system. Now we consider the alternative statements of the endemic equilibrium problem (1.4).

    Lemma 4.1. (˜S,˜E,˜I,˜R) satisfies (1.4) if and only if (S,E,I,R) satisfies the following system,

    {dELE+diag{βiSiSi+Ii+Ei+Ri}IσE=0,dILI+σEYI=0,dRLR+YIαR=0,dSS+dEE+dII+dRR=ξ. (4.1)

    Moreover, ˜S=κS,˜E=κE,˜I=κI,˜R=κR and

    κ=N0ni=1(Si+Ei+Ii+Ri).

    Proof. Note that

    L(dS˜S+dE˜E+dI˜I+dR˜R)=0,

    which implies that

    dS˜S+dE˜E+dI˜I+dR˜R=κξ,κR+,

    where ξ is the unique positive eigenvector of the unique principal eigenvalue 0 of L satisfying iΩξi=1. Set S=˜S/κ,E=˜E/κ,I=˜I/κ,R=˜R/κ. (4.1) can be obtained from (1.4).

    Now we investigate the asymptotic behavior of the endemic equilibria when dS0. Recall that Λ1(dRL+αdiag{1γiβi}) is the smallest eigenvalue of (1.9).

    Theorem 4.1. Suppose that R0>1. Under this assumption, we have

    (i) As dS0, EE,II,RR

    (ii) The set J+:={i|Mi=ξi,iΩ} is nonempty, where Mi:=dEEi+dIIi+dRRi;

    (iii) If further assume Λ1(dRL+αdiag{1γiβi})<0, then the set J:={i|Mi<ξi,iΩ} is not empty.

    Proof. (i). Note that Ei,Ii,Ri>0 for any iΩ,dS>0. dSS+dEE+dII+dRR=ξ yields that βiSiIiSi+Ei+Ii+Ri exhibit uniform boundedness for any dS>0. After extracting a subsequence if necessary, EE as dS0 where E0. Using analogous analysis, it can be shown that II,RR as dS0 where I,R0, which satisfy

    {dILI+σEYI=0,dRLR+YIαR=0. (4.2)

    Now we show that E0, i.e., E>0. It can be proved by contradiction. Assuming E=0, it follows from Eq (4.2) that I=R=0. Consequently, S almost everywhere as dS0. Thus βiSiSi+Ei+Ii+Riβi   as dS0. Define

    K=E1+I1+R1,ˆE=EK,ˆI=IK,ˆR=RK.

    Observe that ˆE,ˆI and ˆR are strictly positive, and ˆE1+ˆI1+ˆR1=1. After extracting a subsequence if necessary, we obtain ˆE,ˆI,ˆR approaches ˆE,ˆI,ˆR respectively as dS0, where ˆEi,ˆIi,ˆRi0 for iΩ and

    ˆE+ˆI+ˆR=1. (4.3)

    It follows from βiSiSi+Ei+Ii+Riβi   as dS0 that ˆE is a solution of

    dELˆEσˆE+BˆI=0,

    which gives

    {dELˆEσˆE+BˆI=0,dILˆI+σˆEYˆI=0,dRLˆR+YˆIαˆR=0. (4.4)

    It can be deduced from Eq (4.3) that the values of ˆE,ˆI,ˆR are significantly greater than zero, which indicates that R0=1. This lead to a contradiction, implying E>0. Therefore, we obtain I,R0.

    To establish the claim that |J+|>0, we prove by contradiction. Suppose that |J+|=0, then as dS0, it follows that S, and thus βiSiSi+Ei+Ii+Riβi   as dS0. Therefore, E is a solution of

    dELEσE+BI=0,

    which yields

    {dELEσE+BI=0,dILI+σEYI=0,dRLR+YIαR=0. (4.5)

    Based on Eq (4.5) in conjunction with I0 and R0, it can be inferred that E0. Consequently, we have R0=1, which leads to a contradiction. Therefore, |J+|>0.

    Part (iii) can be shown through contradiction. Assume now that |J|=0. Denote hi:=βiSiIiSi+Ei+Ii+RiαRi and choose φRn subject to the condition that φ0. By multiplying the first three equations of (4.1) by φT and adding the results together, we obtain

    φTL(dEE+dII+dRR)+φTh=0. (4.6)

    As |J|=0, M=0. Thus, we obtain

    φTh0  as  dS0 (4.7)

    φRn such that φ0.

    Let ϕ0 be a positive left eigenvector of Λ1(dRL+αdiag{1γiβi}), i.e.

    dRLTϕ0+αdiag{1γiβi}ϕ0=Λ1ϕ0, (4.8)

    Since S,E,I,R0 on Ω and dRLR+YIαR=0, we obtain

    dRLR+αdiag{1γiβi}R>diag{γiβi}h,. (4.9)

    Multiplying (4.9) by ϕ0 and applying (4.8), we obtain

    Λ1ϕT0R>ni=1γihiβiϕ0,i.

    Let dS0, it follows from (4.7) that Λ1ϕT0R0. Given that ϕ0,R>0 on Ω, it follows that Λ1>0. This contradiction finishes the proof of (iii).

    Theorem 4.2. Assume R0>1 and Λ1(dRL+αdiag{1γiβi})<0. Then we have

    (i) As dS0,

    ξdSN0ni=1(ξiMi)  and  ˜S˜S=N0(ξM)ni=1(ξiMi);

    (ii) There exist positive constants C1,C2, independent of dS such that for sufficiently small dS,

    C1˜EidS,˜IidS,˜RidSC2,iΩ.

    Proof. (i). For further purposes, denote M:=dEE+dII+dRR. By (4.1), we have

    N0=ni=1(˜Si+˜Ei+˜Ii+˜Ri)=κdS(ni=1dS(Ei+Ii+Ri)+ni=1(ξiMi)).

    Given that S,E,I,R0 and dSS+dEE+dII+dRR=ξ, we can conclude that E,I and R exhibit uniform boundednes with respect to dS. Therefore,

    ni=1dS(Ei+Ii+Ri)0  as  dS0.

    By Theorem 4.1(i), (ii),

    ni=1(ξiMi)ni=1(ξiMi)>0   as  dS0.

    Therefore,

    κdSN0ni=1(ξiMi)   as  dS0. (4.10)

    Furthermore, (4.1) implies ˜S=κdS(ξM). Based on (4.10) and Theorem 4.1(i), we obtain

    ˜S˜S=N0(ξM)ni=1(ξiMi)

    as dS0.

    Now we proceed to the proof of (ii). By dSS+dEE+dII+dRR=ξ and ˜E=κdSdSE,  ˜I=κdSdSI,  ˜R=κdSdSR, we get

    0<˜E1dS,˜I1dS,˜R1dS<κdSmax{1dE,1dI,1dR}.

    Hence, (i) implies

    lim supdS0˜EdS,lim supdS0˜IdS,lim supdS0˜RdSN0ni=1(ξiMi)max{1dE,1dI,1dR}. (4.11)

    Now we prove

    min{˜Ei,˜Ii,˜Ri,iΩ}/dS0,  as  dS0 (4.12)

    by contradiction. Assume that min{˜Ei,˜Ii,˜Ri,iΩ}/dS=o(dS). By (1.4) and direct calculation, ni=1˜Edx,ni=1˜Ii,ni=1˜Ri=o(dS), which implies

    ni=1dE˜Ei+dI~Ii+dR˜RidS0  as  dS0. (4.13)

    Note that

    N0=ni=1κdSni=1dE˜Ei+dI˜Ii+dR˜RidS+ni=1(˜Ei+˜Ii+˜Ri).

    Let dS0, it follows from (i), (4.11) and (4.13) that

    N0=N0ni=1(ξiMi).

    As a result, we have |J|=0. This contradiction finishes the proof of (ii).

    The authors of this work receive financial support from various sources. S.Y. is partially funded by the National Natural Science Foundation of China (NSFC) grant number 12101487 and the China Scholarship Council under grant number 202006280442. J.W. is supported by the Canada Research Chair Program with reference number 230720. P.S. receives support from the National Natural Science Foundation of China (NSFC) grants 12101487 (PS) and 12220101001 (PS), the China Postdoctoral Science Foundation under grant number 2020M683445, and the Postdoctoral Fellowship of York University, Toronto, Canada. The authors also express their gratitude to the referees for their valuable assistance in reviewing the manuscript.

    The authors declare there is no conflict of interest.



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