Return to Article Details Dynamics of a model of nonlocal dispersion viral infection that includes humoral immunity, intracellular delay, and virus-to-cell and cell-to-cell transmissions

Dynamics of a model of nonlocal dispersion viral infection that includes humoral immunity, intracellular delay, and virus-to-cell and cell-to-cell transmissionsThanks:  Corresponding author email: djilali.salih@yahoo.fr, s.djilali@univ-chlef.dz
1Laboratory of Mathematics Modeling and Applications, University of Adrar
National Road No. 06, 01000, Adrar, Algeria.
2Department of Mathematics, Faculty of Exact Sciences and informatics,
Hassiba Benbouali University, Chlef, Algeria

Boubakr Lamouri1, Ahmed Boudaoui1 and Salih Djilali2
Date: November 11, 2025; accepted: June 30, 2026; published online: August 31, 2026.
Abstract.

This paper investigates a delayed spatiotemporal viral infection model with two types of transmissions, namely, virus-to-cell transmission, and cell-to-cell transmission, and a special focus on the cell-mediated immunity. First, we establish that the model system has a global attractor and that the model admits a unique positive solution which is globally defined. The basic reproduction numbers of immunity 1 and infection 0 are then determined. The immunity-related reproduction number 1 is defined only when 0>1, which serves as a threshold parameter. Indeed, it is obtained that the infection-free steady state is globally asymptotically stable when 0<1, the immunity-free infected steady state is globally asymptotically stable when 11<0, and the infected-immune steady state is globally asymptotically stable when 1>1. Finally, numerical simulation is performed in order to validate our theoretical results.

Key words and phrases: 
Global attractor, cell-to-cell transmission, nonlocal diffusion, basic reproduction number, uniform persistence.
2005 Mathematics Subject Classification
92B, 35B, 58Z

1. Introduction

Viral dynamics is a well-established branch of applied mathematics that aims to describe the temporal evolution of infected cells and viral load through mathematical modeling. During viral infection, the specific immune response plays a crucial role in controlling disease progression. This response includes both cell-mediated immunity, driven by cytotoxic T lymphocytes (CTLs), and humoral immunity, mediated by antibodies produced by B cells. Since the pioneering work of Nowak and Bangham [1], a wide range of mathematical models have been developed to investigate immune responses against infected cells [9][15].

To better understand the interplay between virus dynamics and immune response, we focus on a transmission mechanism that incorporates both virus-to-cell and cell-to-cell infection pathways. In this context, Jiazhe and Rui [2] proposed the following basic viral dynamic model:

(1) S(ι) =λβS(ι)I(ι)αS(ι)V(ι)βSS(ι),
V(ι) =βeμτVS(ιτV)I(ιτV)+αeμτVS(ιτV)V(ιτV)βVV(ι),
I(ι) =β1V(ι)γ1I(ι)R(ι)βII(ι),
R(ι) =ζI(ι)R(ι)βRR(ι).

Here, S, I, V, and R represent the densities of uninfected cells, infected cells, free virus particles, and B cells, respectively. The parameters describe biological processes such as infection rates, natural death rates, immune response activation, and viral production. The delay τV accounts for the intracellular period between viral entry and the release of new virions, while eμτV represents the survival probability during this latent phase.

It is well known that viruses can spread over relatively long distances within biological tissues, for instance through blood circulation or intercellular transport mechanisms. Therefore, the classical diffusion operator, typically modeled by the Laplacian, may not adequately capture such long-range movements, as it only describes local interactions based on infinitesimal displacements. In contrast, nonlocal diffusion operators provide a more realistic framework by incorporating spatial interactions over a finite range.

Specifically, nonlocal diffusion is often modeled by an integral operator of the form

ΩJ(ηϑ)(Ψ(ι,ϑ)Ψ(ι,η))𝑑ϑ,

where J() is a dispersal kernel describing the probability of movement from location ϑ to η. Unlike classical diffusion, this operator accounts for long-range dispersal and allows individuals (or particles) to move directly between distant locations. From a biological perspective, this is particularly relevant for viral propagation, where transport mechanisms are not purely local.

Another important distinction concerns boundary conditions. In classical diffusion models, no-flux (Neumann) boundary conditions are imposed through local derivatives. However, in the nonlocal framework, the dispersal process inherently involves interactions across the entire domain Ω. As a result, boundary effects are implicitly incorporated into the integral operator, provided that the kernel J is properly defined (for instance, with compact support or normalization conditions). In this work, we complement the nonlocal formulation with homogeneous Neumann-type conditions to ensure consistency with biological assumptions of no net outward flux.

Moreover, to enhance biological realism, we incorporate time delays representing the latent period between infection and viral production, as well as the activation delay of the immune response. Such delays have been widely studied in the literature [17, 22, 24], mainly in the context of virus-to-cell transmission. However, their combined effect with cell-to-cell transmission and nonlocal dispersal remains less understood.

Motivated by the above considerations, we propose the following delayed viral dynamic model with nonlocal diffusion:

(2) Sι= λ1(η)βS(η)Sβ(η)SIα(η)SV,
Vι= β(η)S(ιτv,η)I(ιτv,η)eμτV
+α(η)S(ιτv,η)V(ιτv,η)eμτVβV(η)V,
Iι= KIΩJ(ηϵ)(I(ι,ϵ)I(ι,η))𝑑ϵ+β1(η)VβI(η)Iγ1(η)IR,
Rι= KRΩJ(ηϵ)(R(ι,ϵ)R(ι,η))𝑑ϵ
+ζ(η)I(ιτR,η)R(ιτR,η)βR(η)R.

with ι>0 and ηΩ. The associated initial and boundary conditions are given by

(3) S(l,η)= S0(l,η),I(l,η)=I0(l,η),V(l,η)=V0(l,η),R(l,η)=R0(l,η),
(l,η)[τ,0]×Ω¯

and

In=Rn=0,ηΩ,ι>0,

where KR and KI are the diffusion coefficients. τR is the period of time needed for antigenic stimulation generating antibody response, Ω¯n is a bounded set with smooth boundary (represents the position).

Let τ=max{τv,τR}. Every parameter should be positive, space-dependent, and Hölder continuous on Ω¯.

The paper is structured as follows. We will examine the well-posedness of (2) and (3) in Section 2. In Section 3, we utilize Kuratowski’s measure of non-compactness to demonstrate the existence of a global compact attractor for the semiflow. The purpose of Section 4 is to determine the basic reproduction numbers of immunity 1 and infection 0. Keep in mind that for 1 to be defined, 0 must be greater than 1. Section 5: A triple dynamics is the primary outcome. In Section 6, when 11<0, the immunity-free infected steady state is globally asymptotically stable, when 1>1, the infected-immune steady state is globally asymptotically stable. These are the general terms used to describe the infection-free steady state and immunity-free infected steady state. Section 7 is configured to demonstrate the use of numerical simulation.

2. Preliminaries

We make the following assumptions on the system parameters (2)–(3):

  • (P0)

    𝐉(η) is Lipschitz continuous on Ω¯.

  • (P1)

    𝐉 is symmetric, non-negative and normalized: 𝐉(z)=𝐉(z) on n, Ω𝐉(z)𝑑z=1, and 𝐉(z)>0 on Ω¯.

Let 𝕏=C([τ,0]×Ω¯,),X=𝕏4, equipped with the supremum norm . Denote by 𝕏+ and X+ the associated positive cones. Throughout this work, all model parameters are assumed to be nonnegative, bounded, and continuous on Ω¯. For notational convenience, given any function C(Ω¯,), we define +:=supηΩ¯(η),:=infηΩ¯(η) , with {λ1,β,α,μ,βV1,βV2,βV3,βV4}.

We now provide a few standard results taking into account the model’s well-posedness (2)–(3). Model (2)–(3) can be rewritten in the following abstract form

(4) d𝕌dι(ι)=A[𝕌](ι)+F[𝕌](ι) ,𝕌(ι)X+,

with

AU=(A1𝕌1A2𝕌2A3𝕌3A4𝕌4)=(βS(v)S(ι,)βV(η)V(ι,)KIΩJ(ηy)(I(ι,y)I(ι,η))dyβI(η)I(ι,)KRΩJ(ηy)(R(ι,y)R(ι,η))dyβR(η)R(ι,)),

and

F𝕌=(λ1(η)β(η)S(ι,)I(ι,)α(η)S(ι,)V(ι,)βS(ιτv,.)I(ιτv,)eμτV+αS(ιτv,)V(ιτv,)eμτVβ1(η)V(ι,)γ1(η)I(ι,η)R(ι,)ζ(η)I(ιτR,)R(ιτR,)).

The following linear operator should be defined on C(Ω¯).

AI[Φ]():=KIΩJ(ηy)(Φ(y)Φ())𝑑yβI()Φ()ΦC(Ω¯).

Some properties on A are given by the following lemma.

Lemma .

A generates a strictly contractive and uniformly continuous semigroup {eAι}ι0.
Additionally, eAιX+X+ ι0.

Proof.

It is possible to break down operator A as A=A1+A2, with

A1𝕌=(βSS(ι,)βVV(ι,) βII(ι,)βRR(ι,)),A2𝕌=(00 KIΩJ(ηϵ)(I(ι,ϵ)I(ι,))𝑑ϵKRΩJ(ηϵ)(R(ι,ϵ)R(ι,))𝑑ϵ),

for 𝕌X. Therefore, a uniformly continuous and strictly contractive semigroup {eAι}ι0 on X is generated by A1. We may deduce that A2 is a Lipschitz function since J is bounded. A generates a C0 -semigroup {eAι}ι0, by Theorem 1.2 in [4] and Corollary VI.1.11 in [5].

Theorem 1.

Suppose that 𝕌0=(φ1(b,η),φ2(b,η),φ3(b,η))X. Hence, there exists τ0>0 such that (2)-(3) has unique positive solution 𝕌(,𝕌0)X, such that either τ0= or limιτ0sup𝕌(ι,𝕌0)=.
Moreover, 𝕌(ι,𝕌0)X+ for ι(0,τ0) if 𝕌0X+.

Proof.

The solution of system (2)-(3) can be written as follows

(5) 𝕌(ι)=𝕌0eAι+0ιeA(ιs)F(𝕌(s))𝑑s.

The function F:XX is locally Lipschitz on X. As established by Lemma 3.1 in [6], Proposition 4.16 in [7] guarantees that system (2) has a unique mild solution, denoted by 𝕌(.,𝕌0), where τ0 verifies either τ0= or lim supιτ0𝕌(ι,𝕌0)=.

Now, we show S(ι,η)>0 for ι(0,τ0) and ηΩ¯. We suppose that τ(0,τ0)and η0Ω verifying that S(τ,η0)=0 and S(ι,η0)>0 for ι<τ. Next, we can specify

τ1=inf{ι(0,τ0);S(ι,η0)=0}.

Since τ0 satisfies τ0= or lim supιτ0𝕌(ι,𝕌0)=., then we deduce that τ1(0,τ0) and S(τ,η0)=0 with S(τ,η)ι0. The first equation of (2) yields

S(τ1,η0)ι=λ1(η0)>0,

that contradicts the assumptions. Hence, S(ι,η)>0 for ι(0,τ0), ηΩ¯.

We put Ahr(𝕌)=𝕌(ι,)+hr𝕌(ι,) for a sufficiently large positive hr. For 𝕌C((0,τ0),X)ß(0,1hr), obviously Ahr(𝕌) is positive. In conjunction with the open ball ß(0,1hr) in X, with radius 1hr and center 0. Hence, (2) can be reworded as this:

d𝕌dι(τ)=Ahr[𝕌](ι)+Fhr[𝕌](ι). 

with Ahr𝕌(ι,)=𝕌(ι,)hr𝕌(ι,). Then,

U(ι)=U0(η)eAhrι+0ιeAhr(ιs)Fhr(U(s))ds. 

Hence, 𝕌(ι,𝕌0)X+ for ι(0,τ0) and 𝕌0X+. ∎

Theorem 2.

The solution of (2)–(3) is globally defined.

Proof.

Let 𝕌=(S,I,V,R) be the solution associated to 𝕌0=(S0,I0,V0,R0). Since dSdt(ι,η)λ(η)βSS(ι,η), Sˇ is a subsolution of

(6) {Sˇι(ι,η)=λ1(η)βSSˇ(ι,η),ηΩ,ι>0,Sˇ(0,η)=maxs[τ,0]S0(s,η),ηΩ.

The unique positive steady state of (6), represented by Sˇf, is globally attractive and well known. By the comparison theorem, we get

lim supιS(ι,η)lim supιSˇ(ι,η)=Sˇf(η)=λ1(η)βs(η) uniformly for ηΩ.

Consequently, K>0 that depends on 𝕌0 such that

SK,ι>0.

Next, we let AI be the generator of the uniformly continuous semigroup {T1(ι)}ι0

I(ι,) =T1(ι)I0+0ιT1(ισ)(β1V(σ,)γ1(x)I(σ,)R(σ,))𝑑σ,
T1(ι)I0+0ιT1(ισ)β1V(σ,)𝑑σ.

By taking the norm on both sides, we obtain

I(ι,)eλιI0+β10ιeλ(ισ)V(σ,)𝑑σ.

We assume that the eigenvalue for the operator AI is λ. According to Proposition 1 in [8], c>0 fulfills λ>c>0. The second equation of (2)–(3) suggests

V(ι,)=eβVιV0+0ιeβV(ισ)(eμτVS(στv,)(βI(στv,)+αV(στv,))𝑑σCLOSE.

Therefore,

V(ι;.)
eβVιV0
+0ιeβV(ισ)eμτVS(στv,)(βI(στv,)+αV(στv,))dσ,
eβVιV0+KeμτV0ιeβV(ισ)(βI(στv,)+αV(στv,))𝑑σ,
eβVιV0+KβeμτV0ιeβV(ισ)(I(στv,)+V(στv,))𝑑σ,

where β=max{β,α}. Using the substitution σ=στv and dropping tilde for simplicity, thus,

V(ι,)
eβVιV0+KβeμτVτvιτveβV(ιτvσ)(I(σ,)+V(σ,))𝑑σ
eβVιV0+KβeμτVτvιτveβV(ιτvσ)V(σ,)𝑑σ
+KβeμτVτvιτveβV(ιτvσ)I(σ,)dσ.

Thus,

V(ι,)
eβVιV0+KβeμτVτvιτveβV(ιτvσ)V(σ,)𝑑σ
+KβeμτVτvιτveβV(ιτvσ){eλσI0+β10σeλ(σs)V(s,)ds}dσ
eβVιV0+KβeμτVιvιτveβV(ττvσ)V(σ,)𝑑σ
+KβeμτV|I0|τvιτveβV(ιτvσ)eλσ𝑑σ
+|β1|KβeμτVτvιτveβV(ιτvσ)τvσeλ(σs)V(s,)𝑑s𝑑σ
eβVιV0+KβeμτVτvιτveβV(ττvσ)V(σ,)𝑑σ
+KβeμτV|I0|τvιτveλσ𝑑σ
+β1MβeμτVτvιτveβV(ιτvσ)τvσeλ(σs)V(s,.)dsdσ.

Consequently,

V(ι,)
eβVιV0+KβeμτVτvιτveβV(ττvσ)V(σ,)𝑑σ
+KβeμτVI0eλτveλ(ιτv)λ
+|β1|KβeμτVeβV(ιτv)τvιτveλsV(s,)sιτve(βVλ)σ𝑑σ𝑑s
eβVτV0+KβeμτVτvιτveβV(ττvσ)V(σ,)𝑑σ
+KβeμτVI0eλτveλ(ιτv)λ
+|β1|KβeμτVeϖ(ττv)λβVτvιτveϖsV(s,)𝑑s
V0+KβeμτVτvιτveϖ(ττvσ)V(σ,)𝑑σ
+KβeμτVI0eλτvλ
+β1KβeμτVλβVτvιτveϖ(ιτvs)V(s,)ds
C1+C2τvιτvV(s,)𝑑s,

with

ϖ =min{βV,λ2},C1=|V0|+KβeμτVI0eλτVλ,
C2 =max{KβeμτV,β1KβeμτVλβV}.

Gronwall’s inequality implies

V C1eC2ιι0.

The last equation of (2) implies

R(ι,.)=eβRιR0+0ιζ(x)eβR(ισ)I(στR,x)R(ιτR,x)dσ.

Therefore, σ=στR

R(ι,.)eβRιR0+ζτRιτReβR(ισ)I(σ,)R(σ,)dσ.

It follows

(7) R(ι,) C4eC5ιι0,

with

I(ι,) I0+C1β1eC2ιλ=C3,
I(ι,) I0eλι+β10ιeλ(ισ)V(σ,)𝑑σ
I0eλι+C1β10ιeλ(ισ)eC2ι𝑑σ
I0+C1β1eC2ιλ=C3.

Therefore, the solution is globally defined, that is, τ0=+. ∎

3. Compactness

We now examine the solution map’s compactness. The semiflow produced by system (2)–(3) is denoted by {Ξ(ι)}ι0. Ξ(ι) is not compact. we may demonstrate the asymptotic compactness of forward orbits in order to demonstrate that Ξ(ι) has a global compact attractor. Thus, using Kuratowski measure κ , which is defined as follows,

κ(𝐁)=inf{r: has a finite cover of diameter<r}.

Then

𝐁 precompactκ(P)=0.

To achieve this goal, we apply several lemmas.

Lemma .

The semiflow {Ξ(ι)}ι0 is point dissipative.

Proof.

We let H(ι,η)=S(ι,η)+V(ι+τv,η); hence,

H(ι,η)ι =λ1(η)(1eμτV)β(η)S(ι,η)I(ι,η)
(1eμτV)α(η)S(ι,η)V(ι,η)βS(η)S(ι,η)βV(η)V(ι+τv,η)
 λ1(η)βS(η)S(ι,η)βV(η)V(ι+τv,η)
 λ1(η)ωH(ι,η)

with ω=min{βS,βV}. Therefore, we get H(ι,η)g1(η), ι+, with g1(η)=H(0,η)+λ1(η)ω. Therefore, the existence of ξ1 independent of the initial conditions verifying

limιsupH(ι,η)ξ1.

On Ω,we integrate the third equation of (2).

ιΩI(ι,η)𝑑η= KIΩΩJ(ηϵ)(I(ι,ϵ)I(ι,η))𝑑ϵ𝑑η
+Ωβ1(η)V(ι,η)dηΩβI(η)I(ι,η)dη
Ωγ1(η)I(ι,η)R(ι,η)dη,
Ωβ1(η)V(ι,η)𝑑ηβIΩI(ι,η)𝑑η,

which gives

ιΩI(ι,η)𝑑η+βIΩI(ι,η)𝑑ηΩβ1(η)V(ι,η)𝑑η.

Let us multiply by eβIι,

eβIιιΩI(ι,η)𝑑η+βIeβIιΩI(ι,η)𝑑η eβIιΩβ1(η)V(ι,η)𝑑η,
ddιeβIιΩI(ι,η)𝑑η eβIιΩβ1(η)V(ι,η)𝑑η,

Integrating by parts, we find

0ιseβIsΩI(s,η)𝑑η𝑑s 0ιeβIsΩβ1(η)V(s,η)𝑑η𝑑s,
eβIιΩI(ι,η)𝑑η+ΩI(0,η)𝑑η 0ιeβIsΩβ1(η)V(s,η)𝑑η𝑑s,
ΩI(ι,η)𝑑η eβIιΩI(0,η)𝑑η
+1βI(1eβIι)Ωβ1(η)V(s,η)dη:=D1.

On the other hand, we have

ιI(ι,η)dη = KIΩJ(ηϵ)(I(ι,ϵ)I(ι,η))𝑑ϵ+β1(η)V(ι,η)βI(η)I(ι,η)
KI(supηΩJ)ΩI(ι,ϵ)𝑑ϵ+β1(η)V(ι,η)βI(η)I(ι,η).

By applying this inequality, the third equation in (2) suggests

dI(ι,)dιKI(supηΩJ)D1+β1+ξ1(KI+βI)I(ι,).

Then,

I(ι,)I0(ι,)e(KI+βI)ι+KI(supηΩJ)D1+β1+ξ1KI+βI(1e(KI+βI)ι).

After that, ξ2 independent of the initial condition satisfying

lim supιI(ι,η)ξ2.

We integrate the final equation of (2) on Ω

ιΩR(ι,η)𝑑x= KRΩΩJ(ηϵ)(R(ι,ϵ)R(ι,η))𝑑ϵ𝑑η
+Ωζ(η)V(ιτR,η)I(ιτR,η)dηΩβR(η)R(ι,η)dη
Ωζ(η)R(ιτR,η)I(ιτR,η)𝑑ηβRΩR(ι,η)𝑑η.

By similar reasoning, there is a ξ3 independent of the initial condition satisfying

lim supιR(ι,η)ξ3.

Therefore, the semiflow Ξ(ι) is point dissipative. ∎

We now demonstrate that Ξ(ι) is a κ contraction; that is, for all ι>0, there exists a continuous function 0k(ι)<1, and a bounded set θ, such that {Ξ(σ)θ,0σι} is bounded and κ(Ξ(σ)θ)k(ι)κ(θ). First, we demonstrate the asymptotic compactness of S and V.

Lemma .

The forward orbit δ+(Ψ):𝕌(ι,,𝕌0),ι0 for (2)–(3) is asymptotically compact for all ΨX+, in that for all sequences ιk. Consequently, ιki to satisfy 𝕌(ιki,,𝕌0) converges in X+ as i .

Proof.

Let us assume that 𝕌0X+, there exists η>0 verifying

S(ι,η;𝕌0)ϖ,I(ι,η;𝕌0)ϖ,for all ηΩ-,ι0.

It is sufficient to demonstrate that 𝕌(ι,η,𝕌0)n1 is equicontinuous in ηΩ¯, ι>0 using the Arzelà–Ascoli theorem. In other words, for all ρ>0, there exists ε>0 guaranteeing |h(ι,η1)h(ι,η2)|<ε2 for all η1,η2Ω¯ as long as |η1η2|<ρ. It is evident that h(ι,η)=λ(η)+β(η)S(ι,η)I(ι,η)eμτV+α(η)S(ι,η)V(ι,η)eμτV is uniformly continuous in ηΩ¯ uniformly for ι0. Clearly,

ι[S(ι,η1)S(ι,η2)]2=
= 2(S(ι,η1)S(ι,η2))[h(ι,η1)h(ι,η2)βS(η1)(S(ι,η1)
S(ι,η2))S(ι,η2)(βS(η1)βS(η2))]
4η(h(ι,η1)h(ι,η2))βS(η1)(S(ι,η1)S(ι,η2))2
+S(ι,η2)(βS(η1)βS(η2))
(4ϖε2+2ε)βS(η1)(S(ι,η1)S(ι,η2))2,

for all ι> τV and η1,η2Ω¯ verifying |η1η2|<ρ, we have

[S(ι,η1)S(ι,η2)]2eβSι[[S(0,η1)S(0,η2)]2]+(4ϖε2+2ε)0ιe2βS(ισ)𝑑σ.

Then

[S(ι,η1)S(ι,η2)]2[S(0,η1)S(0,η2)]2+(4ϖε2+2ε)0ιe2βS(ισ)𝑑σ.

Given that ηΩ¯, the initial condition’s uniform continuity suggests that S0(0,η1)S0(0,η2)<ε2 for all η1,η2Ω¯ verifying that |η1η2|<ρ1. Therefore, for any |η1η2|< min{ρ,ρ1}, with η1,η2Ω- , we have

[S(ι,η1)S(ι,η2)]2ε24+(4ϖε2+2ε)2βS.

Since S is equicontinuous, we can infer that ε is arbitrary. We demonstrate that I, V and R are equicontinuous by using a similar logic. ∎

Lemma .

The semiflow Ξ(ι) is a κ-contraction.

Proof.

Clearly, I(ι,η,𝕌0) and R(ι,η,𝕌0) satisfy

Iι(ι,η) =KIΩJ(ηϵ)(I(ι,ϵ)I(ι,η))𝑑ϵβI(η)I(ι,η),
Rι(ι,η) =KRΩJ(ηϵ)(R(ι,ϵ)R(ι,η))𝑑ϵβR(η)R(ι,η),
I(s,η) =I0(s,η),R(s,η)=R0(s,η), s[τ,0], ηΩ-.

Therefore,

I(ι,)=T1(ι)I0+0ιT1(ισ)(β1()V(σ,,𝕌0)γ1()I(σ,,𝕌0)R(σ,,𝕌0))𝑑σ,

and

R(ι,)=T2(ι)R0+ζ()0ιT2(vσ)V(στR,,𝕌0)I(στR,,𝕌0)𝑑σ,

according to Lemma 2.1 in [10], the operators LV and LR have strictly positive eigenfunctions, which correspond to strictly negative eigenvalues μ0 and μ1, respectively. These eigenvalues are defined by

μ0=infΨ0L2(Ω)Ψ0L2(Ω)=1{ΩΩJ(ηε)(Ψ0(ε)Ψ0(η))2dεdη+ΩβI(η)Ψ02(η)dη},

and

μ1=infΨ1L2(Ω)Ψ1L2(Ω)=1{ΩΩJ(ηε)(Ψ1(ε)Ψ1(η))2dεdη+ΩβR(η)Ψ12(η)dη}.

Now, we decompose the semiflow into Φ(ι)=Φ1(ι)+Φ2(ι), where

Φ1(ι)U0=(0,0,T1(ι)I0,T2(ι)R0),ι0

and

Φ2(ι)𝕌0= (S(,,𝕌0),V(,,𝕌0)CLOSE,
0ιT1(ισ)(β1()V(σ,,𝕌0)γ1()I(σ,,𝕌0)R(σ,,𝕌0))𝑑σ,
OPENζ()0ιT2(τσ)V(στR,,𝕌0)I(στR,,𝕌0)𝑑σ),ι0

We infer that S,V,I, and R are asymptotically compact from Section 3. Consequently, Φ2 is compact. Thus, we have κ(Φ2(ι)𝐀)=0 for a bounded set 𝐀X+ and ι0. Additionally,

Φ1(ι)𝕌0 eLVι𝕌0+eLRι𝕌0
eμ0ι𝕌0+eμ1ι𝕌0
eμι𝕌0,ι>0,

with μ=max{μ0,μ1} Hence,

Φ1(ι)eμι.

Hence, for ι>0

κ(Φ(ι)𝐀)κ(Φ1(ι)𝐀)+κ(Φ2(ι)𝐀)eμικ(𝐀).

The conclusion that follows is that Ξ(ι) is a κ- contraction. ∎

We infer from  Section 3 that the semiflow Ξ(ι) is point dissipative. Moreover,  Section 3 shows that Ξ(ι) is a κ-contraction. By Lemma 2.3.5 in [11], every κ-contraction semiflow is asymptotically smooth. Furthermore, Theorem 2.6 in [12] implies that Ξ(ι) possesses a global compact attractor that attracts all bounded subsets of X+. We denote this attractor by .

4. Basic reproduction number and steady-state

A steady-state (S(η),V(η),I(η),R(η)) of (2) is the solution of the following system

(8) 0= λ1(η)βS(η)S(η)β(η)S(η)I(η)α(η)S(η)V(η),
0= β(η)S(η)I(η)eμτV+α(η)S(η)V(η)eμτVβV(η)V(η),
0= KIΩJ(ηϵ)(I(ϵ)I(η))𝑑ϵ+β1(η)V(η)βI(η)I(η)γ1(η)I(η)R(η),
0= KRΩJ(ηϵ)(R(ϵ)R(η))𝑑ϵ+ζ(η)I(η)R(η)βR(η)R(η),ηΩ-

Clearly, (2) has a unique VFSS H0=(λ1(η)βS(η),0,0,0),ηΩ¯ . We linearize (2) in order to get the basic reproduction number 0 corresponding to (2) at H0, we have

(9) Vι= β(η)λ1(η)βS(η)I(ιτv,η)eμτV+α(η)λ1(η)βS(η)V(ιτv,η)eμτVβV(η)V(ι,η),
Iι= KIΩJ(ηϵ)(I(ι,ϵ)I(ι,η))𝑑ϵ+β1(η)V(ι,η)
βI(η)I(ι,η)γ1(η)I(ι,η)R(ι,η),ι>0,ηΩ-.

We only take into consideration the subsystem because S and R are absent from the middle two equations

{ι(VI)=𝐀(VI), ηΩ,ι0,Iυ=0, ηΩ, ι0,

we define the following linear operators ,:(:=C(Ω,2)) ,

(10) =(βV(η)0β1(η)AI), =(β(η)λ1(η)βS(η)eμτVα(η)λ1(η)βS(η)eμτV00),

and the linear operator 𝐀:=+

We designate 𝐓(ι) as the semigroup that generates. Therefore, we can define 0 as the spectral radius of 𝒦, which is

0=𝔯(𝒦).

Here, the next generation operator, 𝒦:=1, can be written as

𝒦(Θ)(η)=(η)0𝐓B(ι)(Θ)(η)𝑑ι,ΘC(Ω-,2),ηΩ.-

To derive a variational expression for the basic reproduction number 0, we obtain the following result.

Lemma .

0=𝔯(1)=r(dV+dI) with dV=βλ1βSβVeμτV and dI=αβ1eμτVλ1(x)βSβVAI1.

Proof.

Let ϕ=Ψ and Ψ=1Θ. Clearly,

{βVΨ1=Θ1 βIΨ1AIΨ2=Θ2.

Solving this system, we get

Ψ1 = Θ1βV,
Ψ2 = AI1(βIΨ1Θ2).

Consequently,

ϕ1 = d1Ψ1+d2Ψ2,
ϕ2 = 0,

with d1=βλ1βSβVeμτV+αeμτVλ1β1βSβVAI1, and d2=AI1αλ1βSeμτV. Hence, we can write

1(Θ1Θ2)=(d1Ψ1+d2Ψ20).

Iteratively, we get

(1)n(Θ1Θ2)=(d1nΨ1+d1n1d2Ψ20).

Hence dIn|(1)n|d1n1|(d1+d2). By applying the squeeze theorem and Gelfand’s formula, we obtain r(1)=𝔯(d1). Since d1=dV+dI, the proof is achieved. From Section 4, we get

(11) 0=𝔯(dV+dI),

where dV=βλ1βSβV is the cell-to-cell transmission’s next generation operator, and dI=αλ1β1βSβVAI1 is the next generation operator for virus-to-cell transmission. The basic reproduction number for cell-to-cell transmission only, when virus-to-cell transmission is not present, is

(12) 0V=𝔯(dV)=maxηΩ-β(η)λ1(η)eμτVβS(η)βV(η).

Furthermore, the basic reproduction number can also be stated by the following variational formula in the situation of virus-to-cell only:

(13) 0I=𝔯(dI)=supΘL2,Θ0Ωα(η)λ1(η)β(η)eμτVβS(η)βV(η)Θ2(η)𝑑ηKIΩΩ12J(ηϵ)(Θ(ϵ)Θ(η))2𝑑ϵ𝑑η+ΩβI(η)Θ2(η)𝑑η.

Clearly, 0 0V +0I . The equality holds if β(η)λ1(η)βS(η)βV(η) is space independent.
The basic reproduction number 1 for the antibody response is determined by

1=supΘL2,Θ0Ωζ(η)I(η)Θ2(η)𝑑ηKRΩΩ12J(ηϵ)(Θ(ϵ)Θ(η))2𝑑ϵ𝑑η+ΩβR(η)Θ2(η)𝑑η.

It is evident that 1 is increasing in ζ(η) and I(η), and decreasing in βR and KR. In particular, we obtain the spatially homogeneous model, where model (2)–(3) degenerates.

1=ζ(η)I(η)βR(η).
Lemma .

01 and δ(𝐀) have the same sign.

5. Extinction scenario of virus

Here, we show that for 0<1, the VFSS is globally stable. First, we use the following theorem to show that this steady state is locally stable:

Theorem 3.

H0 is locally asymptotically stable if 0<1, and unstable if 0>1.

Proof.

To begin with, we notice that 𝐀τV, the generator of the linearized system (9), and 𝐀, the generator of the corresponding linear system without delay (τV=0), are not identical. Moreover, the principal eigenvalue of the problem is S(𝐀τV).

(14) β(η)λ1(η)βS(η)Θ3(η)eλτVeμτV+α(η)λ1(η)βS(η)Θ2(η)eλτVeμτVβV(η)Θ2(η)
=λΘ2(η),
KIΩJ(ηϵ)(Θ3(ι,ϵ)Θ3(ι,η))𝑑ϵ+β1(η)Θ2(η)βI(η)Θ3(η)
=λΘ3(η),

with 𝐀τV as the generator of the semigroup PτV(ι) related to the delayed linear system (9), for all ηΩ¯ . The principal eigenvalue of the problem is δ(𝐀).

(15) β(η)λ1(η)βS(η)Θ3(η)eμτV+α(η)λ1(η)βS(η)Θ2(η)eμτVβV(η)ϕ2(η)= λΘ2(η),
KIΩJ(ηϵ)(Θ3(ι,ϵ)Θ3(ι,η))𝑑ϵ+β1(η)Θ2(η)βI(η)Θ3(η)= λΘ3(η).

We determine that δ(A) has the same sign as S(AτV) by applying Theorem 9.2.1 in [14] to the eigenvalue problem associated with the first equation of (14) and (15). As a result, there is a sign difference between the exponential growth rate associated with the semi-group linked to the delayed system (14), known as ω(PτV)), and the one associated with P(ι), the semi-group linked to the generator 𝐀. The exponential growth bound of P(ι) is now found and is defined as

ω(𝕋)=max{δ(𝐀),ω0(𝕋)},

utilizing eω0(𝕋):=κ(P(ι)) where κ represents the non-compactness measure. Finding the sign of ω(PτV) is necessary to demonstrate the local stability of H0. If ω(𝕋)<0, then H0 is locally asymptotically stable; if ω(𝕋)>0, then H0 is unstable. Since ω(𝕋) and ω(PτV) have the same sign, it is possible to infer the local asymptotic stability of VFSS by using Theorem 2.1 in [16] to determine the sign of ω(𝕋). Section 3 gives us ω0μ0<0. As a result, H0stability (or instability) is determined by δ(A). Section 4 gives us that 01 and δ(𝐀) have the same sign. Consequently, if R0<1, then H0 is locally asymptotically stable since δ(𝐀)<0 and ω(𝕋)<0. Nevertheless, if 11<0, δ(𝐀)>0. Consequently, ω(𝕋)=δ(𝐀)>0, and H0 becomes unstable. ∎

The following lemma is used to construct test functionals.

Lemma .

Given the assumption that 0 <1, then there exists a ρ>0 that satisfies ρ<1. It may be written as

(16) ρ=supΘL2,Θ0{ 1KIΩΩ12J(ηϵ)(Θ(η)Θ(ϵ))Θ(η)𝑑ϵ𝑑η+ΩβI(η)Θ2(η)𝑑η×
×Ωα(η)λ1(η)βI(η)eμτVβS(η)βV(η)βeμτVλ(η)Θ2(η)dη}.

Additionally, the following system is also satisfied by the positive functions Θ0=(Θ10(η),Θ20(η))T.

(17) 0= (βV(η)α(χ)λ1(η)βS(η)eμτV)Θ10(η)+β1(η)Θ20(η),ηΩ-,
0= 1ρβ(η)eμτVλ1(η)βS(η)Θ10(η)+KIΩJ(ϵ)(Θ20(ι,ϵ)Θ20(ι,η))𝑑ϵβI(η)Θ20(η),
ηΩ-.
Proof.

Notice that for 0 <1, we obtain that (η):=β(η)λ1(η)βS(η)βV(η) 1. As a consequence, we use a different decomposition from (𝐀:=𝖡+𝖥) of the operator 𝐀 as

(18) 𝐀:=(βV(η)α(η)λ1(η)βS(η)eμτV0β1(η)AI)+(0β(η)eμτVλ1(η)βS(η)00):=𝖡+𝖥

We assume that the semigroup generated by 𝖡 is 𝕋𝖡(ι). Clearly, S(𝖡)<0. We assume that ρ is the spectral radius of 𝐊, which is

ρ=𝔯(𝐊)

Using 𝐊:=𝖥𝖡1 and applying Theorem 3.5 in [13], we conclude that if 0<1, then δ(𝐀)<0 and, consequently, ρ<1. We then consider an eigenvalue ϱ of 𝐀 that solves the following problem

OPENϱΘ(η)=A[Θ](η),ηΩ,Θ=(Θ1(η),Θ2(η))TC(CLOSE-Ω,2).

Next, we claim that the problem

(19) 𝖡[Θ](η)=ϱ𝖥[Θ](η),ηΩ-,

has a unique principal eigenpair (η,Θ0) that fulfills

ρ=𝔯(𝖥𝖡1)=𝔯(𝖡1𝖥)=1ϱ.

We assume that 𝕋𝖡(ι) is the semigroup related to the generator 𝖡 in order to demonstrate this assertion. Theorem 3.5 in [13] suggests that since 𝖡 is a positive resolvent and T𝖡(ι) is a positive semigroup

(γI𝖡)1Θ=0eϵιT𝖡(ι)𝑑ι,Θ𝔇(𝖡).

Assume that γ=0. Next,

𝖡1Θ=0TB(ι)𝑑ι,Θ𝔇(𝖡).

Additionally, we get

𝖥(𝖡1)Θ=𝐊[Θ](η).

Now, we set Θ =𝖡Θ-, then we get

𝖥Θ-=𝐊(η)𝖡Θ-.

Therefore,

𝖡Θ-=𝖥Θ-𝐊(η),ηΩ-.

Given that (19) has a unique positive eigenvalue ϱ, then ϱ=1ρ with a positive eigenfunction. In (18), we obtain ϱ by substituting ϱ with Θ=(Θ1CLOSE, OPENΘ2),ηΩ- is the corresponding positive eigenfunction , we have

(20) 0 =(βV(η)α(η)λ1(η)βS(η)eμτV)Θ1(η)+β1(η)Θ2(η),ηΩ-,
0 =ϱβ(η)eμτVλ1(η)βS(η)Θ1(η)+KIΩJ(ηϵ)(Θ2(ι,ϵ)Θ2(ι,η))𝑑ϵβI(η)Θ2(η),
ηΩ-.

The first equation in (20) suggests that (η)<1

Θ1(η)=β1(η)βS(η)βV(η)βS(η)α(η)λ1(η)eμτVΘ2(η), ηΩ-.

Therefore, we enter it in the second equation in (20) to get

(21) 0= ϱβ(η)eμτVλ1(η)βS(η).β1(η)βS(η)βV(η)βS(η)α(η)λ1(η)eμτVΘ2(η)
+KIΩJ(ηϵ)(Θ2(ι,ϵ)Θ2(ι,η))dϵβI(η)Θ2(η),
ηΩ-,

hence,

KIΩJ(ηϵ)(Θ2(ι,ϵ)Θ2(ι,η))𝑑ϵβI(η)Θ2(v)
+ϱβ(η)eμτVλ1(η)β1(η)βV(η)βS(η)α(η)λ1(η)eμτVΘ2(η)=0.

By considering the decomposition (18) and taking into account that ϱ=1ρ, we deduce that

(22) ρ=supΘL2,Θ0{ 1KIΩΩ12J(ηϵ)(Θ(η)Θ(ϵ))Θ(η)𝑑ϵ𝑑η+ΩβI(η)Θ2(η)𝑑η×
×Ωα(η)λ1(η)β1(η)eμτVβS(η)βV(η)βeμτVλ1(η)Θ2(η)dη}.

Consider that Θ0=(Θ10(η),Θ20(η))T is the eigenfunction associated with the eigenvalue ρ. Consequently, ρ and Θ0 satisfy the following system

(23) (βV(η)α(η)λ1(η)βS(η)eμτV)Θ10(η)+β1(η)Θ20(η) =0,ηΩ-
1ρβ(η)eμτVλ1(η)βS(η)Θ10(η)+KIΩJ(ηϵ)(Θ20(ι,ϵ)CLOSE
OPENΘ20(ι,η))dϵβI(η)Θ20(η) =0,ηΩ-

Remark .

System (24) is used to establish the global stability of the VFSS.

The following theorem presents the principal result of this section.

Theorem 4.

Let 0 be defined by (22) and (23). Then H0 is globally stable in if 0<1.

Proof.

It has been proved that lim supιS(ι,η)λ1(η)βS(η).Therefore, there exists θ>0 sufficiently small such that S(ι,η)λ1(η)βS(η)+θ for ι sufficiently large. Furthermore, given the fact that ρ is defined by (22), if 0<1, then for ρθ small enough there exist positive functions (Θ1θ(η),Θ2θ(η)) satisfying the following system:

(24) 0= (βV(η)α(η)eμτV(λ1(η)βS(v)+θ))Θ1θ(η)+β1(η)Θ2θ(η),
0= 1ρθβ(η)eμτV(λ1(η)βS(η)+θ)Θ1θ(η)
+KIΩJ(ηϵ)(Θ2θ(ϵ)Θ2θ(η))dϵβI(η)Θ2θ(η),ηΩ-.

We then define the following Lyapunov function

(25) E(ι)=ϑ(ι)+ϑτV(ι),

with

ϑ(ι)=ΩΘ1θ(η)V(ι,η)+Θ2θ(η)V(ι,η)𝑑η,

and

ϑτV(ι)= eμτVΩΘ1θ(η)β(η)(λ1(η)βS(η)+θ)0τVI(τs,η)𝑑s
+Θ1θ(η)α(η)(λ1(η)βS(η)+θ)0τVV(ιs,η)dsdη,

where system (24) defines Θ1θ(η),Θ2θ(η). The derivative of ϑ(ι) is

ϑ(ι)=
= ΩΘ1θ(η)V(ι,η)τ+Θ2θ(η)V(ι,η)ι𝑑η
= Ω{Θ1θ(η)[β(η)S(ιτv,η)I(ιτv,η)eμτV
+α(η)S(ιτv,η)V(ιτv,η)eμτVβV(η)V(ι,η)]}dη
+Ω{Θ2θ(η)[KIΩJ(ηϵ)(I(ι,ϵ)I(ι,η))dϵ+β1(η)V(ι,η)βI(η)I(ι,η)]}dη.

Using the inequality S(ι,η)λ1(η)βS(η)+θ, we get

ϑ(ι) Ω{Θ1θ(η)[β(η)(λ1(η)βS(η)+θ)I(ιτv,η)eμτV
+α(η)(λ1(η)βS(η)+θ)V(ιτv,η)eμτVβV(η)V(ι,η)]}dη
+Ω{Θ2θ(η)[KIΩJ(ηϵ)I(ι,ϵ)dϵ+β1(η)V(ι,η)
βI(η)I(ι,η)(γ1(η)+KIJ1(η))I(ι,η)]}dη,

with J1(η)=ΩJ(ηϵ)𝑑ϵ. Using (24), we get

ϑ(ι)
Ω{Θ1θ(η)[β(η)(λ1(η)βS(η)+θ)I(ιτv,η)eμτV
+α(η)(λ1(η)βS(η)+θ)V(ιτv,η)eμτV]}dη
+Ω{Θ2θ(η)[KIΩJ(ηϵ)I(ι,ϵ)dϵ+β1(η)V(ι,η)]}dη
ΩV(ι,η){α(η)λ1(η)βS(η)eμτVϕ1θ(η)+β1(η)Θ2θ(η)}dη
ΩI(ι,η){1ρθβ(η)eμτV(λ1(η)βS(η)+θ)Θ1θ(η)+KIΩJ(ηϵ)Θ2θ(ϵ)dϵ}dη.

Using the assumptions on J, we have

ΩΩJ(ηϵ)Θ2θ(ϵ)I(ι,η)𝑑ϵ𝑑η=ΩΩJ(ηϵ)Θ2θI(ι,ϵ)𝑑η𝑑ϵ.

Therefore, some simplification gives

ϑ(ι) ΩΘ1θ(η)β(η)(λ1(η)βS(η)+θ)eμτV[I(ι,η)ρθ+I(ιτv,η)]𝑑η
+ΩΘ1θ(η)α(η)(λ1(η)βS(η)+θ)eμτV[V(ιτv,η)V(ι,η)].

ϑτV(ι) is given by

ϑτV(τ)= eμτVΩΘ1θ(η)β(η)(λ1(η)βS(η)+θ)ι0τVI(ιs,η)𝑑s
+Θ1θ(η)α(η)(λ1(η)βS(η)+θ)ι0τVV(ιs,η)dsdη
= eμτVΩΘ1θ(η)β(η)(λ1(η)βS(η)+θ)0τVtI(ιs,η)𝑑s
+Θ1θ(η)α(η)(λ1(η)βS(η)+θ)0τVιV(ιs,η)dsdη
= eμτVΩΘ1θ(η)β(η)(λ1(η)βS(η)+θ)0τV(s)I(ιs,η)𝑑s
+Θ1θ(η)α(η)(λ1(η)βS(η)+θ)0τV(s)V(ιs,η)dsdη
= {eμτVΩΘ1θ(η)β(η)(λ1(η)βS(η)+θ)I(ιs,η)0τV
+Θ1θ(η)α(η)(λ1(η)βS(η)+θ)V(ιs,η)0τVdη}
= eμτVΩΘ1θ(η)β(η)(λ1(η)βS(η)+θ)(I(ι,η)I(ιτV,η))
+Θ1θ(η)α(η)(λ1(η)βS(η)+θ)(V(ι,η)V(ιτV,η))dη

. By summing ϑ(ι) and ϑτV(ι), we obtain

E(ι)eμτVΩΘ1θ(η)β(η)I(ι,η)(λ1(η)βS(η)+θ)(11ρθ)𝑑η0.

Equality holds for I(ι,η)=0. Given this, the third equation of (2) implies that V(ι,η)=0. Therefore, (I(ι,η),V(ι,η))(0,0) uniformly for ηΩ as ι.
We further obtain that S(ι,η) λ1(η)βS(η) uniformly for ηΩ as ι. To sum up, the theorem is proved. ∎

6. Uniform persistence

The following spaces should be defined:

ξ0 = {𝕌X+:V(,η)>0,I(,η)>0 ,ηΩ¯};
ξ0 = {𝕌X+:V(,η)0,I(,η)0 ,ηΩ¯};
M = {𝕌0ξ0:Φ(ι,𝕌0)ξ0 for ι0}.

Then, we obtain the following results.

Theorem 5.

Assume that 11<0. Then the system (2)–(3) exhibits strong uniform persistence. Additionally, in ξ0, it possesses at least one positive steady state that remains uniformly persistent.

Proof.

To establish the uniform persistence of the system, it is sufficient to verify that all the assumptions of Theorem 3 in [18] are satisfied. ∎

The following claims provide the detailed proof:

Claim .

H0 is globally stable for semiflow {Ξ(ι)}ι0 restricted to ξ0.

For 𝕌0ξ0, (2) becomes

(26) {dSdι(ι,η)=λ1(η)βS(η)S(ι,η), V(,η)=0,I(,η)=0, ι>0,ηΩ-.

Therefore, S(ι,η) goes to λ1(η)βS(η) for all ηΩ-. Hence, ω(𝕌0)={(λ1(η)βS(η),0,0,0)}, where ω(ϕ) is the ω-limit set.

Claim .

For any 𝕌0ξ0, we have 𝕌>0 for all ηΩ- , and ι>0. We consider the problem

i(ι,η)ι =KIΩJ(ηϵ)(i(ι,ϵ)i(ι,η))𝑑ϵβI(η)i(ι,η),ι>0,ηΩ-.
i(0,η) =I0(η),ηΩ.-

Comparison principle implies that I(ι,η)i(ι,η). Now, it is sufficient to show that i(ι,η)>0 for all ι>0, ηΩ¯. This proof follows the approach of Proposition 2.2 in [19]. We define the operator Υ:(Ω-)(Ω-)

Υi(ι,η)=KIΩJ(ηy)(I(ι,y)I(ι,η))𝑑y,ηΩ-.

Clearly, Υ is continuous, and {eΥι}ι0 forms a uniformly continuous, positive semigroup on (Ω¯) generated by Υ. , where for ι0, we get eΥι =n=0ιnΥnn!. Further,

eΥτ(I0(s,η))=n=0τnΥn(I0(s,η))n!,ι>0,s[τ,0].

Since i(τ,η)0, we have Υn+1(I0(s,η))=J(ηy)ΥnI0(s,υ)dυ,ηΩ¯, s[τ,0], and n1. The iteration process implies the existence of n0 such that Υn+1(I(0,η))>0,ηΩ¯ and nn0. Hence,Υn(I0(s,η))>0,ηΩ¯,s[τ,0]. Consequently,

i(ι,η) =e(KIβI(η))eΥτ(I0(s,η))+0τ e(KIβI(η))(τσ) eΥ(τσ)i(ι,η)dσ,
e(KIβI(η))eΥτ(I0(s,η))>0,

for ι>0,ηΩ¯. The second equation in (2) yields 𝕌>0 from this inequality. The value ξ0 is therefore positively invariant.

Claim .

H0 is a uniform weak repeller for ξ0 for 11<0. In order to prove this assertion, we demonstrate that there exists a sufficiently small θ1>0 such that the semiflow {Ξ(ι)}ι0 satisfies

lim supιΞ(ι,𝕌0)H0θ1.

We use contradiction arguments to demonstrate this result. Suppose that lim supι𝚵(ι,𝕌0)H0θ1. Hence, there exists ι1 such that 𝕌(ι,𝕌0) satisfies

λ1(η)βS(η)θ1S(ι,η)λ1(η)βS(η)+θ1,0V(ι,η)θ1,ιι1,ηΩ¯.

Since 11<0, we have Sθ(A)>0. Then, there exists ε sufficiently small satisfying

χϕ(η) =(βV(η)α(η)(λ1(η)βS(η)+θ1)β(η)(λ1(η)βS(η)+θ1)β1(η)AI)Θ(η),Θ(η)
=(ϕ1(η),Θ2(η))T,ηΩ.

has a principal eigenpair (χp,Θp) with χp>0 and Θp=(Θ1p,Θ2p) . Hence, for ι>ι1,

V(ι,η)ι= β(η)(λ1(η)βS(η)+θ1)I(ιτv,η)eμτV
+α(η)(λ1(η)βS(η)+θ1)V(ιτv,η)eμτVβV(η)V(ι,η),
Iι(ι,η)= KIΩJ(ηy)(I(ι,y)I(ι,η))𝑑y+β1(η)V(ι,η)βI(η)I(ι,η),
ι> ι1,ηΩ.-

By comparison principle, we obtain

OPEN(V(η),I(η))T=yPeyP(ιι1CLOSE)ΘP(η).

There is a contradiction since yP>0.

Claim .

System (2) is strongly uniformly persistent in ξ0 for 11<0. For the orbit ϰ+(𝕌0):=𝕌ι0{Ξ(ι)}, let ϱ1(𝕌0) be the ϱ - limit set. Since Φ(ι,ξ0)ξ0 and Ξ(ι,ξ0)ξ0, then M=ξ0 and ϱ(𝕌0)=H0 for all 𝕌0 ξ0. Hence, H0 is isolated in X. Since ϱ(𝕌0)= H0 for all 𝕌0ξ0, then there is no cycle in ξ0 from H0 to H0. We obtain Ws(H0)ξ0=, with Ws(E0)={𝕌0X:limιΞ(ι,𝕌0)=H0}, based on the outcomes of Section 6. Since all the assumptions of Theorem 3 in [18] are satisfied, system (2) is strongly uniformly persistent for 11<0. Consequently, the assertion follows. Moreover, the endemic equilibrium E=(S,V,I) satisfies the following steady-state system:

(27) 0= λ1(η)βS(η)S(ι,η)β(η)S(ι,η)I(ι,η)α(η)S(ι,η)V(ι,η)
0= β(η)S(η)I(η)eμτV+α(η)S(η)V(η)eμτVβV(η)V(η)ηΩ-.
0= KIΩJ(ηϵ)(I(ϵ)I(η))𝑑ϵ+β1(η)V(η)βI(η)I(η).

7. Global behavior analysis

Theorem 6.

If 11< 0, then besides H0, there is also a unique immunity-free infected steady state H1=(S,V,I,0).

Proof.

From (27), we see that system (27) has a unique endemic equilibrium H1=(S,V,I,0). Let us construct the following Lyapunov function,

E(ι)=ϱ1(ι)+ϱ1τV,

with

ϱ1(ι)= Ωζ1(η){eμτVP(η)H(S(ι,η)S(η))+V(η)H(V(ι,η)V(η))
+ζ2(η)I(η)H(I(ι,η)I(η))}dη,

and

ϱ1τV= ΩeμτVζ1(η)0τVβ(η)S(η)I(η)H(S(ιϑ)I(ιϑ)SI)
+α(η)S(η)V(η)H(S(ιϑ)V(ιϑ)SV)dϑdη,

where H(t)=t1ln(t), for t>0, ζ1(η)=2eμτVβ1(η)V(η)β(η)S(η),ζ2(x)=2eμτVβ(η)S(η)I(η)β1(η)V(η). For simplicity, we use the notations

S =S(ι,η),I=I(ι,η),V=V(ι,η),SτV=S(ιτV,η),IτV=I(ιτV,η),
VτV =V(ιτV,η),S=S(η),I=I(η),V=V(η).

First, we compute ϱ1(ι), which is

ϱ1(ι)=
= Ωζ1(η){eμτV(1SS)Sι+(1VV)Vι+ζ2(η)(1II)Iι}𝑑η
= Ωζ1(η){eμτV(1SS)[λ1βSSβSIαSV]
+(1VV)[βSτVIτVeμτV+αSτVVτVeμτVβVV]
+ζ2(η)(II)[KIΩJ(ηϵ)I(ι,ϵ)dϵ+β1(η)V(η)(βI(η)+KI)I(η) ]}dη.

H1=(S,V,I,0) satisfies

{λ1(η)=βS(η)S+β(η)SI+α(η)SV, βV(η)V =β(η)SIeμτV+α(η)SVeμτV, (βI(η)+KI)I =KIΩJ(ηϵ)I(ϵ)dϵ+β1(η)V,ηΩ-.

Thus, after some simplification, ϱ1(ι) becomes

ϱ1(ι)=
= Ωζ1(η){eμτVβS(η)S(2SSSS)
+eμτVβ(η)SI[2SSSISI+IIVVSτVIτVSISτVIτVVSIV]
+eμτVα(η)SV[2SSSVSV+SτVVτVSVSτVVτVSV]
+KIζ2(η)[(1II)KIΩJ(ηϵ)I(ι,ϵ)𝑑ϵ+(1II)ΩJ(ηϵ)I(ϵ)𝑑ϵ]
+ζ2(η)β1(η)V(1+VVIIVIVI)}dη.

Notice that ζ2(η)β1(η)V= β(η)SIeμτV and ζ1(η)ζ2(η)=2I. Then, we obtain

ϱ1(ι)= Ωζ1(η){eμτVβS(η)S(2SSSS)
+eμτVβ(η)SI[3SSSISI+SτVIτVSISτVIτVVSIVVIVI]
+eμτVα(η)SV[2SSSVSV+SτVVτVSVSτVVτVSV]}dη
+2KIΩI{(1II)ΩJ(ηϵ)I(ϵ)dϵ
+(1II)ΩJ(ηϵ)I(ι,ϵ)dϵ}dη
= Ωζ1(η){eμτVβS(η)S(2SSSS)
+eμτVβ(η)SI[3SSSISI+SτVIτVSISτVIτVVSIVVIVI]
+eμτVα(η)SV[2SSSVSVSτVVτVSVSτVVτVSV]}dη
+KIΩΩJ(ηϵ)I(ϵ)I(η){2I(η)I(ι,ϵ)I(ι,ϵ)I(ϵ)I(ϵ)I(ι,ϵ)I(ι,ϵ)I(η)}dηdϵ.

Alternatively, ϱ1τV is computed as follows:

ϱ1τV= ιΩeμτVζ1(η)0τVβ(η)SIH(SϑIϑSI)+α(η)SVH(SϑVϑSV)𝑑ϑ𝑑η
= ΩeμτVζ1(η){β(η)SI[H(SISI)H(SτVIτVSI)]
+α(η)SV[H(SVSV)H(SτVVτVSV)]}dη.

Adding ϱ1(ι) and ϱ1τV, we find

E(ι)= Ωζ1(η){eμτVβS(η)S(2SSSS)
+eμτVβ(η)SI[3SSSISISτVIτVSISτVIτVVSIVVIVI]
+eμτVα(η)SV[2SSSVSVSτVVτVSVSτVVτVSV]}dη
+KIΩΩJ(ηϵ)I(ϵ)I(η){2I(η)I(ι,ϵ)I(ι,ϵ)I(ϵ)I(ϵ)I(ι,ϵ)I(ι,ϵ)I(η)}dηdϵ
+ΩeμτVζ1(η){β(η)SI[H(SISI)H(SτVIτVSI)]
+α(η)SV[H(SVSV)H(SτVVτVSV)]}dη
= ΩeμτVζ1(η){βS(η)S(2SSSS)
+β(η)SI[H(SS)H(SτVIτVVSIV)H(IVIV)]
+α(η)SV[H(SS)H(SτVVτVSV)]}dη
+KIΩΩJ(ηϵ)I(ϵ)I(η){2I(η)I(ι,ϵ)I(ι,ϵ)I(ϵ)I(ϵ)I(ι,ϵ)I(ι,ϵ)I(η)}dηdϵ.

Thus, E(ι)0, and equality holds when S=S, V=V, I=I. As a result, the immunity-free infected steady state H1 is globally asymptotically stable. ∎

8. Graphical representation

Here, we validate our theoretical results by numerical simulation. We set Ω=(1,1) and

J(η)={2.2523e1η21,1<η<1,0,otherwise.

This ensures that

J(η)𝑑η1,

and that assumptions (P0) and (P1) are fulfilled.

Refer to caption
Figure 1. Kernel function J.

We consider the following set of parameters:

μ =101, βS(η) =15101, βV(η) =12101, βI(η) =13101,
λ1(η) =29101, τV =98101, β(η) =β¯, α(η) =1,
KI =5102, ζ =98101.
S0(ι,η) =3101+5102cos(η),
V0(ι,η) =41102+102cos(η)+103cos(ι),
I0(ι,η) =71102+101cos(η)+102cos(3ι).

First, we take β¯=47102. Then 0=0.7614<1, and the infection-free steady state H0 is globally asymptotically stable; see Figure 2.

Refer to caption
Refer to caption
Figure 2. When 0=0.7614<1, the infection-free steady state H0 is globally asymptotically stable.

Next, the immunity-free infected steady state H1 is globally asymptotically stable for β¯=23102 and βR(η)=187101, where 0=3.5874>1 and 1=0.3956<1; see Figure 3.

Refer to caption
Refer to caption
Figure 3. When 0=3.5874>1 and 1=0.3956<1, the immunity-free infected steady state H1 is globally asymptotically stable.

Finally, for βR(η)=53101 and 1=2.1894>1, the infected-immune steady state H2 is globally asymptotically stable; see Figure 4.

Refer to caption
Refer to caption
Figure 4. When 1=2.1894>1, the infected-immune steady state H2 is globally asymptotically stable.

9. Conclusions

This paper presents and examines a model of nonlocal dispersal viral infection that includes humoral immunity, intracellular delay, and transmissions from virus to cell as well as cell to cell.

We have proved that the global dynamics of system (2)–(3) is determined by the basic reproduction number of infection 0 and the basic reproduction number of immunity 1. By analyzing the characteristic equations and constructing Lyapunov functionals, we have obtained the following conclusions: if 0<1, then the infection-free steady state H0 is globally asymptotically stable; if 11<0, then the immunity-free infected steady state H1 is globally asymptotically stable. We note that constructing a Lyapunov functional for a delayed nonlocal diffusive problem is not straightforward.

Additionally, using a set of assertions that confirm every assumption of Theorem 3 in [18], we demonstrated the system’s uniform persistence. As a result, the semiflow Ξ(ι) is uniformly persistent, and the system admits at least one positive steady state. Finally, we supported our findings using graphical representations.

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