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 ℜ1 and infection ℜ0 are then determined. The immunity-related reproduction number ℜ1 is defined only when ℜ0>1, which serves as a threshold parameter. Indeed, it is obtained that the infection-free steady state is globally asymptotically stable when ℜ0<1, the immunity-free infected steady state is globally asymptotically stable when ℜ1≤1<ℜ0, and the infected-immune steady state is globally asymptotically stable when ℜ1>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) =λ−β⁢S⁢(ι)⁢I⁢(ι)−α⁢S⁢(ι)⁢V⁢(ι)−βS⁢S⁢(ι),
V′⁢(ι) =β⁢e−μ⁢τV⁢S⁢(ι−τV)⁢I⁢(ι−τV)+α⁢e−μ⁢τV⁢S⁢(ι−τV)⁢V⁢(ι−τV)−βV⁢V⁢(ι),
I′⁢(ι) =β1⁢V⁢(ι)−γ1⁢I⁢(ι)⁢R⁢(ι)−βI⁢I⁢(ι),
R′⁢(ι) =ζ⁢I⁢(ι)⁢R⁢(ι)−βR⁢R⁢(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 e−μ⁢τV 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⁡(η−ϑ)⁢(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) ∂S∂ι= λ1⁢(η)−βS⁢(η)⁢S−β⁡(η)⁢S⁢I−α⁡(η)⁢S⁢V,
∂V∂ι= β⁡(η)⁢S⁢(ι−τv,η)⁢I⁢(ι−τv,η)⁢e−μ⁢τV
+α⁡(η)⁢S⁢(ι−τv,η)⁢V⁢(ι−τv,η)⁢e−μ⁢τV−βV⁢(η)⁢V,
∂I∂ι= KI⁢∫ΩJ⁡(η−ϵ)⁢(I⁡(ι,ϵ)−I⁡(ι,η))⁢𝑑ϵ+β1⁢(η)⁢V−βI⁢(η)⁢I−γ1⁢(η)⁢I⁢R,
∂R∂ι= KR⁢∫ΩJ⁡(η−ϵ)⁢(R⁡(ι,ϵ)−R⁡(ι,η))⁢𝑑ϵ
+ζ⁡(η)⁢I⁢(ι−τR,η)⁢R⁢(ι−τR,η)−βR⁢(η)⁢R.

with ι>0 and η∈Ω. 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

∂I∂n=∂R∂n=0,η∈∂Ω,ι>0,

where KR and KI are the diffusion coefficients. τR is the period of time needed for antigenic stimulation generating antibody response, Ω¯⊂ℝn 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 ℜ1 and infection ℜ0. Keep in mind that for ℜ1 to be defined, ℜ0 must be greater than 1. Section 5: A triple dynamics is the primary outcome. In Section 6, when ℜ1≤1<ℜ0, the immunity-free infected steady state is globally asymptotically stable, when ℜ1>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 ℝn, ∫Ω𝐉⁡(z)⁢𝑑z=1, and 𝐉⁡(z)>0 on Ω¯.

Let 𝕏=C⁡([−τ,0]×Ω¯,ℝ),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 ℋ∈C⁡(Ω¯,ℝ), we define ℋ+:=supη∈Ω¯ℋ⁡(η),ℋ−:=infη∈Ω¯ℋ⁡(η) , with ℋ∈{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) d⁢𝕌d⁢ι⁢(ι)=A⁡[𝕌]⁢(ι)+F⁡[𝕌]⁢(ι)⁢ ,⁢𝕌⁢(ι)∈X+,

with

A⁢U=(A1⁢𝕌1A2⁢𝕌2A3⁢𝕌3A4⁢𝕌4)=(−βS⁢(v)⁢S⁢(ι,⋅)−βV⁢(η)⁢V⁢(ι,⋅)KI⁢∫ΩJ⁡(η−y)⁢(I⁡(ι,y)−I⁡(ι,η))⁢dy−βI⁢(η)⁢I⁢(ι,⋅)KR⁢∫ΩJ⁡(η−y)⁢(R⁡(ι,y)−R⁡(ι,η))⁢dy−βR⁢(η)⁢R⁢(ι,⋅)),

and

F⁢𝕌=(λ1⁢(η)−β⁡(η)⁢S⁢(ι,⋅)⁢I⁢(ι,⋅)−α⁡(η)⁢S⁢(ι,⋅)⁢V⁢(ι,⋅)βS(ι−τv,.)I(ι−τv,⋅)e−μ⁢τV+αS(ι−τv,⋅)V(ι−τv,⋅)e−μ⁢τVβ1⁢(η)⁢V⁢(ι,⋅)−γ1⁢(η)⁢I⁢(ι,η)⁢R⁢(ι,⋅)ζ⁡(η)⁢I⁢(ι−τR,⋅)⁢R⁢(ι−τR,⋅)).

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

AI⁢[Φ]⁢(⋅):=KI⁢∫ΩJ⁡(η−y)⁢(Φ⁡(y)−Φ⁡(⋅))⁢𝑑y−βI⁢(⋅)⁢Φ⁢(⋅)⁢; ⁢Φ∈C⁡(Ω¯).

Some properties on A are given by the following lemma.

Lemma .

A generates a strictly contractive and uniformly continuous semigroup {eA⁢ι}ι≥0.
Additionally, eA⁢ι⁢X+⊂X+ ∀ ι≥0.

Proof.

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

A1⁢𝕌=(−βS⁢S⁢(ι,⋅)−βVV(ι,⋅) −βI⁢I⁢(ι,⋅)−βR⁢R⁢(ι,⋅)),A2⁢𝕌=(00 KI⁢∫ΩJ⁡(η−ϵ)⁢(I⁡(ι,ϵ)−I⁡(ι,⋅))⁢𝑑ϵKR⁢∫ΩJ⁡(η−ϵ)⁢(R⁡(ι,ϵ)−R⁡(ι,⋅))⁢𝑑ϵ),

for 𝕌∈X. Therefore, a uniformly continuous and strictly contractive semigroup {eA⁢ι}ι≥0 on X is generated by A1. We may deduce that A2 is a Lipschitz function since J is bounded. A generates a C0 -semigroup {eA⁢ι}ι≥0, 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 limι→τ0−⁡⁢sup‖𝕌⁡(ι,𝕌0)‖=∞.
Moreover, 𝕌⁡(ι,𝕌0)∈X+ for ι∈(0,τ0) if 𝕌0∈X+.

Proof.

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

(5) 𝕌⁡(ι)=𝕌0⁢eA⁢ι+∫0ιeA⁡(ι−s)⁢F⁢(𝕌⁡(s))⁢𝑑s.

The function F:X→X 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 supι→τ0−‖𝕌⁡(ι,𝕌0)‖=∞.

Now, we show S⁡(ι,η)>0 for ι∈(0,τ0) and η∈Ω¯. We suppose that ∃ τ∼∈(0,τ0)and η0∈Ω 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 supι→τ0−‖𝕌⁡(ι,𝕌0)‖=∞., then we deduce that τ1∈(0,τ0) and S(τ∼,η0)=0 with ∂S⁡(τ∼,η)∂ι≤0. 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), η∈Ω¯.

We put Ahr⁢(𝕌)=𝕌⁡(ι,⋅)+hr⁢𝕌⁢(ι,⋅) for a sufficiently large positive hr. For 𝕌∈C⁡((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:

d⁢𝕌d⁢ι(τ)=Ahr[𝕌](ι)+Fhr[𝕌](ι). 

with Ahr⁢𝕌⁢(ι,⋅)=𝕌⁡(ι,⋅)−hr⁢𝕌⁢(ι,⋅). Then,

U(ι)=U0(η)eAhr⁢ι+∫0ιeAhr⁢(ι−s)Fhr(U(s))ds. 

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

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 d⁢Sd⁢t⁢(ι,η)≤λ⁡(η)−βS⁢S⁢(ι,η), Sˇ is a subsolution of

(6) {∂Sˇ∂ι(ι,η)=λ1(η)−βSSˇ(ι,η),η∈Ω,ι>0,Sˇ(0,η)=maxs∈[−τ,0]S0(s,η),η∈Ω.

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

lim supι→∞S⁡(ι,η)≤lim supι→∞Sˇ⁢(ι,η)=Sˇf⁢(η)=λ1⁢(η)βs⁢(η)⁢ uniformly for ⁢η∈Ω.

Consequently, ∃K>0 that depends on 𝕌0 such that

‖S‖≤K⁢,⁢ι>0.

Next, we let AI be the generator of the uniformly continuous semigroup {T1⁢(ι)}ι≥0

I⁡(ι,⋅) =T1⁢(ι)⁢I0+∫0ιT1⁢(ι−σ)⁢(β1⁢V⁢(σ,⋅)−γ1⁢(x)⁢I⁢(σ,⋅)⁢R⁢(σ,⋅))⁢𝑑σ,
≤T1⁢(ι)⁢I0+∫0ιT1⁢(ι−σ)⁢β1⁢V⁢(σ,⋅)⁢𝑑σ.

By taking the norm on both sides, we obtain

‖I⁡(ι,⋅)‖≤e−λ⁢ι⁢‖I0‖+‖β1‖⁢∫0ιe−λ⁡(ι−σ)⁢‖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,⋅)=e−βV⁢ι⁢V0+∫0ιe−βV⁢(ι−σ)⁢(e−μ⁢τV⁢S⁢(σ−τv,⋅)⁢(β⁢I⁢(σ−τv,⋅)+α⁢V⁢(σ−τv,⋅))⁢𝑑σCLOSE.

Therefore,

∥V(ι;.)∥≤
≤e−βV⁢ι⁢‖V0‖
+∫0ιe−βV⁢(ι−σ)e−μ⁢τV∥S(σ−τv,⋅)∥(∥β∥∥I(σ−τv,⋅)∥+∥α∥∥V(σ−τv,⋅)∥)dσ,
≤e−βV⁢ι⁢‖V0‖+K⁢e−μ⁢τV⁢∫0ιe−βV⁢(ι−σ)⁢(‖β‖⁢‖I⁡(σ−τv,⋅)‖+‖α‖⁢‖V⁡(σ−τv,⋅)‖)⁢𝑑σ,
≤e−βV⁢ι⁢‖V0‖+K⁢β∗⁢e−μ⁢τV⁢∫0ιe−βV⁢(ι−σ)⁢(‖I⁡(σ−τv,⋅)‖+‖V⁡(σ−τv,⋅)‖)⁢𝑑σ,

where β∗=max⁡{‖β‖,‖α‖}. Using the substitution σ∗=σ−τv and dropping tilde for simplicity, thus,

‖V⁡(ι,⋅)‖≤
≤e−βV⁢ι⁢‖V0‖+K⁢β∗⁢e−μ⁢τV⁢∫τvι−τve−βV⁢(ι−τv−σ)⁢(‖I⁡(σ,⋅)‖+‖V⁡(σ,⋅)‖)⁢𝑑σ
≤e−βV⁢ι⁢‖V0‖+K⁢β∗⁢e−μ⁢τV⁢∫τvι−τve−βV⁢(ι−τv−σ)⁢‖V⁡(σ,⋅)‖⁢𝑑σ
+Kβ∗e−μ⁢τV∫τvι−τve−βV⁢(ι−τv−σ)∥I(σ,⋅)∥dσ.

Thus,

‖V⁡(ι,⋅)‖≤
≤e−βV⁢ι⁢‖V0‖+K⁢β∗⁢e−μ⁢τV⁢∫τvι−τve−βV⁢(ι−τv−σ)⁢‖V⁡(σ,⋅)‖⁢𝑑σ
+Kβ∗e−μ⁢τV∫τvι−τve−βV⁢(ι−τv−σ){e−λ⁢σ∥I0∥+∥β1∥0σe−λ⁡(σ−s)∥V(s,⋅)∥ds}dσ
≤e−βV⁢ι⁢‖V0‖+K⁢β∗⁢e−μ⁢τV⁢∫ιvι−τve−βV⁢(τ−τv−σ)⁢‖V⁡(σ,⋅)‖⁢𝑑σ
+K⁢β∗⁢e−μ⁢τV|I0|∫−τvι−τve−βV⁢(ι−τv−σ)⁢e−λ⁢σ⁢𝑑σ
+|β1|K⁢β∗⁢e−μ⁢τV⁢∫−τvι−τve−βV⁢(ι−τv−σ)⁢∫−τvσe−λ⁡(σ−s)⁢‖V⁡(s,⋅)‖⁢𝑑s⁢𝑑σ
≤e−βV⁢ι⁢‖V0‖+K⁢β∗⁢e−μ⁢τV⁢∫−τvι−τve−βV⁢(τ−τv−σ)⁢‖V⁡(σ,⋅)‖⁢𝑑σ
+K⁢β∗⁢e−μ⁢τV|I0|∫−τvι−τve−λ⁢σ⁢𝑑σ
+∥β1∥Mβ∗e−μ⁢τV∫−τvι−τve−βV⁢(ι−τv−σ)∫−τvσe−λ⁡(σ−s)∥V(s,.)∥dsdσ.

Consequently,

‖V⁡(ι,⋅)‖≤
≤e−βV⁢ι⁢‖V0‖+K⁢β∗⁢e−μ⁢τV⁢∫−τvι−τve−βV⁢(τ−τv−σ)⁢‖V⁡(σ,⋅)‖⁢𝑑σ
+K⁢β∗⁢e−μ⁢τV⁢‖I0‖⁢eλ⁢τv−e−λ⁡(ι−τv)λ
+|β1|K⁢β∗⁢e−μ⁢τV⁢e−βV⁢(ι−τv)⁢∫−τvι−τveλ⁢s⁢‖V⁡(s,⋅)‖⁢∫sι−τve(βV−λ)⁢σ⁢𝑑σ⁢𝑑s
≤e−βV⁢τ⁢‖V0‖+K⁢β∗⁢e−μ⁢τV⁢∫−τvι−τve−βV⁢(τ−τv−σ)⁢‖V⁡(σ,⋅)‖⁢𝑑σ
+K⁢β∗⁢e−μ⁢τV⁢‖I0‖⁢eλ⁢τv−e−λ⁡(ι−τv)λ
+|β1|K⁢β∗⁢e−μ⁢τV⁢e−ϖ⁡(τ−τv)λ−βV⁢∫−τvι−τveϖ⁢s⁢‖V⁡(s,⋅)‖⁢𝑑s
≤‖V0‖+K⁢β∗⁢e−μ⁢τV⁢∫−τvι−τve−ϖ⁡(τ−τv−σ)⁢‖V⁡(σ,⋅)‖⁢𝑑σ
+K⁢β∗⁢e−μ⁢τV⁢‖I0‖⁢eλ⁢τvλ
+‖β1‖⁢K⁢β∗⁢e−μ⁢τVλ−βV∫−τvι−τve−ϖ⁡(ι−τv−s)∥V(s,⋅)∥ds
≤C1+C2⁢∫−τvι−τv‖V⁡(s,⋅)‖⁢𝑑s,

with

ϖ =min⁡{βV−,λ2},C1=|V0|+K⁢β∗⁢e−μ⁢τV⁢‖I0‖⁢eλ⁢τVλ,
C2 =max⁡{K⁢β∗⁢e−μ⁢τV,‖β1‖⁢K⁢β∗⁢e−μ⁢τVλ−βV}.

Gronwall’s inequality implies

‖V‖ ≤C1⁢eC2⁢ι⁢, ⁢ι≥0.

The last equation of (2) implies

R(ι,.)=e−βR⁢ιR0+∫0ιζ(x)e−βR⁢(ι−σ)I(σ−τR,x)R(ι−τR,x)dσ.

Therefore, σ∗⁣∗=σ−τR

∥R(ι,.)∥≤e−βR⁢ι∥R0∥+ζ∗∫−τRι−τRe−βR⁢(ι−σ)∥I(σ,⋅)∥∥R(σ,⋅)∥dσ.

It follows

(7) ‖R⁡(ι,⋅)‖ ≤C4⁢eC5⁢ι⁢, ⁢ι≥0,

with

‖I⁡(ι,⋅)‖ ≤‖I0‖+C1⁢‖β1‖⁢eC2⁢ιλ=C3,
‖I⁡(ι,⋅)‖ ≤‖I0‖⁢e−λ⁢ι+‖β1‖⁢∫0ιe−λ⁡(ι−σ)⁢‖V⁡(σ,⋅)‖⁢𝑑σ
≤‖I0‖⁢e−λ⁢ι+C1⁢‖β1‖⁢∫0ιe−λ⁡(ι−σ)⁢eC2⁢ι⁢𝑑σ
≤‖I0‖+C1⁢‖β1‖⁢eC2⁢ιλ=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)}ι≥0. 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⁢ precompact⇔κ⁡(P)=0.

To achieve this goal, we apply several lemmas.

Lemma .

The semiflow {Ξ⁡(ι)}ι≥0 is point dissipative.

Proof.

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

∂H⁡(ι,η)∂ι =λ1⁢(η)−(1−e−μ⁢τV)⁢β⁢(η)⁢S⁢(ι,η)⁢I⁢(ι,η)
−(1−e−μ⁢τV)⁢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∈ℝ+, with g1⁢(η)=H⁡(0,η)+λ1⁢(η)ω. Therefore, the existence of ξ1 independent of the initial conditions verifying

limι→∞⁢sup‖H⁡(ι,η)‖≤ξ1.

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

∂∂ι⁢∫ΩI⁡(ι,η)⁢𝑑η= KI⁢∫Ω∫ΩJ⁡(η−ϵ)⁢(I⁡(ι,ϵ)−I⁡(ι,η))⁢𝑑ϵ⁢𝑑η
+∫Ωβ1(η)V(ι,η)dη−∫ΩβI(η)I(ι,η)dη
−∫Ωγ1(η)I(ι,η)R(ι,η)dη,
≤ ∫Ωβ1⁢(η)⁢V⁢(ι,η)⁢𝑑η−βI−⁢∫ΩI⁡(ι,η)⁢𝑑η,

which gives

∂∂ι⁢∫ΩI⁡(ι,η)⁢𝑑η+βI−⁢∫ΩI⁡(ι,η)⁢𝑑η≤∫Ωβ1⁢(η)⁢V⁢(ι,η)⁢𝑑η.

Let us multiply by eβI−⁢ι,

eβI−⁢ι⁢∂∂ι⁢∫ΩI⁡(ι,η)⁢𝑑η+βI−⁢eβI−⁢ι⁢∫ΩI⁡(ι,η)⁢𝑑η ≤ eβI−⁢ι⁢∫Ωβ1⁢(η)⁢V⁢(ι,η)⁢𝑑η,
dd⁢ι⁢eβI−⁢ι⁢∫ΩI⁡(ι,η)⁢𝑑η ≤ eβI−⁢ι⁢∫Ωβ1⁢(η)⁢V⁢(ι,η)⁢𝑑η,

Integrating by parts, we find

∫0ι∂∂s⁢eβI−⁢s⁢∫ΩI⁡(s,η)⁢𝑑η⁢𝑑s ≤∫0ιeβI−⁢s⁢∫Ωβ1⁢(η)⁢V⁢(s,η)⁢𝑑η⁢𝑑s,
eβI−⁢ι⁢∫ΩI⁡(ι,η)⁢𝑑η+∫ΩI⁡(0,η)⁢𝑑η ≤∫0ιeβI−⁢s⁢∫Ωβ1⁢(η)⁢V⁢(s,η)⁢𝑑η⁢𝑑s,
∫ΩI⁡(ι,η)⁢𝑑η ≤e−βI−⁢ι⁢∫ΩI⁡(0,η)⁢𝑑η
+1βI−(1−e−βI−⁢ι)∫Ωβ1(η)V(s,η)dη:=D1.

On the other hand, we have

∂∂ι⁢I⁢(ι,η)⁢d⁢η = KI⁢∫ΩJ⁡(η−ϵ)⁢(I⁡(ι,ϵ)−I⁡(ι,η))⁢𝑑ϵ+β1⁢(η)⁢V⁢(ι,η)−βI⁢(η)⁢I⁢(ι,η)
≤ KI⁢(supη∈Ω⁢J)⁢∫ΩI⁡(ι,ϵ)⁢𝑑ϵ+β1⁢(η)⁢V⁢(ι,η)−βI⁢(η)⁢I⁢(ι,η).

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

d⁢I⁢(ι,⋅)d⁢ι≤KI⁢(supη∈Ω⁢J)⁢D1+β1+⁢ξ1−(KI+βI−)⁢I⁢(ι,⋅).

Then,

I⁡(ι,⋅)≤I0⁢(ι,⋅)⁢e−(KI+βI−)⁢ι+KI⁢(supη∈Ω⁢J)⁢D1+β1+⁢ξ1KI+βI−⁢(1−e−(KI+βI−)⁢ι).

After that, ∃ξ2 independent of the initial condition satisfying

lim supι→∞‖I⁡(ι,η)‖≤ξ2.

We integrate the final equation of (2) on Ω

∂∂ι⁢∫ΩR⁡(ι,η)⁢𝑑x= KR⁢∫Ω∫ΩJ⁡(η−ϵ)⁢(R⁡(ι,ϵ)−R⁡(ι,η))⁢𝑑ϵ⁢𝑑η
+∫Ωζ(η)V(ι−τR,η)I(ι−τR,η)dη−∫ΩβR(η)R(ι,η)dη
≤ ⁢∫Ωζ⁡(η)⁢R⁢(ι−τR,η)⁢I⁢(ι−τR,η)⁢𝑑η−βR−⁢∫ΩR⁡(ι,η)⁢𝑑η.

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

lim supι→∞‖R⁡(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 0≤k⁡(ι)<1, and a bounded set θ, such that {Ξ(σ)θ,0≤σ≤ι} 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),ι≥0 for (2)–(3) is asymptotically compact for all Ψ∈X+, in that for all sequences ιk→∞. Consequently, ∃ιk⁢i to satisfy 𝕌⁡(ιk⁢i,⋅,𝕌0) converges in X+ as i →∞.

Proof.

Let us assume that 𝕌0∈X+, there exists η>0 verifying

∣S(ι,η;𝕌0)∣≤ϖ,∣I(ι,η;𝕌0)∣≤ϖ,for all η∈Ω-,ι≥0.

It is sufficient to demonstrate that 𝕌⁢(ι,η,𝕌0)n≥1 is equicontinuous in η∈Ω¯, 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,η2∈Ω¯ as long as |η1−η2|<ρ. It is evident that h⁡(ι,η)=λ⁡(η)+β⁡(η)⁢S⁢(ι,η)⁢I⁢(ι,η)⁢e−μ⁢τV+α⁡(η)⁢S⁢(ι,η)⁢V⁢(ι,η)⁢e−μ⁢τV is uniformly continuous in η∈Ω¯ uniformly for ι≥0. 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))]
≤ 4⁢η⁢(h⁡(ι,η1)−h⁡(ι,η2))−βS⁢(η1)⁢(S⁡(ι,η1)−S⁡(ι,η2))⁢2
+∣S⁡(ι,η2)⋅(βS⁢(η1)−βS⁢(η2))∣
≤ (4⁢ϖ⁢ε2+2⁢ε)−βS−⁢(η1)⁢(S⁡(ι,η1)−S⁡(ι,η2))2,

for all ι> −τV and η1,η2∈Ω¯ verifying |η1−η2|<ρ, we have

[S⁡(ι,η1)−S⁡(ι,η2)]2≤e−βS−⁢ι⁢[[S⁡(0,η1)−S⁡(0,η2)]2]+(4⁢ϖ⁢ε2+2⁢ε)⁢∫0ιe−2⁢βS−⁢(ι−σ)⁢𝑑σ.

Then

[S⁡(ι,η1)−S⁡(ι,η2)]2≤[S⁡(0,η1)−S⁡(0,η2)]2+(4⁢ϖ⁢ε2+2⁢ε)⁢∫0ιe−2⁢βS−⁢(ι−σ)⁢𝑑σ.

Given that η∈Ω¯, the initial condition’s uniform continuity suggests that ∣S0⁢(0,η1)−S0⁢(0,η2)∣<ε2 for all η1,η2∈Ω¯ verifying that |η1−η2|<ρ1. Therefore, for any |η1−η2|< m⁢i⁢n⁢{ρ,ρ1}, with η1,η2∈Ω- , we have

[S⁡(ι,η1)−S⁡(ι,η2)]2≤ε24+(4⁢ϖ⁢ε2+2⁢ε)2⁢βS−.

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

∂I∂ι⁢(ι,η) =KI⁢∫ΩJ⁡(η−ϵ)⁢(I⁡(ι,ϵ)−I⁡(ι,η))⁢𝑑ϵ−βI⁢(η)⁢I⁢(ι,η),
∂R∂ι⁢(ι,η) =KR⁢∫ΩJ⁡(η−ϵ)⁢(R⁡(ι,ϵ)−R⁡(ι,η))⁢𝑑ϵ−βR⁢(η)⁢R⁢(ι,η),
I⁡(s,η) =I0(s,η),R(s,η)=R0(s,η), s∈[−τ,0], η∈Ω-.

Therefore,

I⁡(ι,⋅)=T1⁢(ι)⁢I0+∫0ιT1⁢(ι−σ)⁢(β1⁢(⋅)⁢V⁢(σ,⋅,𝕌0)−γ1⁢(⋅)⁢I⁢(σ,⋅,𝕌0)⁢R⁢(σ,⋅,𝕌0))⁢𝑑σ,

and

R⁡(ι,⋅)=T2⁢(ι)⁢R0+ζ⁡(⋅)⁢∫0ιT2⁢(v−σ)⁢V⁢(σ−τR,⋅,𝕌0)⁢I⁢(σ−τR,⋅,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=−infΨ0∈L2⁢(Ω)‖Ψ0‖L2⁢(Ω)=1{∫Ω∫ΩJ(η−ε)(Ψ0(ε)−Ψ0(η))2dεdη+∫ΩβI(η)Ψ02(η)dη},

and

μ1=−infΨ1∈L2⁢(Ω)‖Ψ1‖L2⁢(Ω)=1{∫Ω∫ΩJ(η−ε)(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),ι≥0

and

Φ2⁢(ι)⁢𝕌0= (S⁡(⋅,⋅,𝕌0),V⁡(⋅,⋅,𝕌0)CLOSE,
∫0ιT1⁢(ι−σ)⁢(β1⁢(⋅)⁢V⁢(σ,⋅,𝕌0)−γ1⁢(⋅)⁢I⁢(σ,⋅,𝕌0)⁢R⁢(σ,⋅,𝕌0))⁢𝑑σ,
OPENζ⁡(⋅)⁢∫0ιT2⁢(τ−σ)⁢V⁢(σ−τR,⋅,𝕌0)⁢I⁢(σ−τR,⋅,𝕌0)⁢𝑑σ),ι≥0

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 𝐀⊂X+ and ι≥0. Additionally,

‖Φ1⁢(ι)⁢𝕌0‖ ≤ ‖eLV⁢ι⁢𝕌0+eLR⁢ι⁢𝕌0‖
≤ eμ0⁢ι⁢‖𝕌0‖+eμ1⁢ι⁢‖𝕌0‖
≤ eμ∗⁢ι⁢‖𝕌0‖,ι>0,

with μ∗=max⁡{μ0,μ1} Hence,

‖Φ1⁢(ι)‖≤eμ∗⁢ι.

Hence, for ι>0

κ⁡(Φ⁡(ι)⁢𝐀)≤κ⁡(Φ1⁢(ι)⁢𝐀)+κ⁡(Φ2⁢(ι)⁢𝐀)≤eμ∗⁢ι⁢κ⁢(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 ℂ.

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)⁢e−μ⁢τV+α⁡(η)⁢S⁢(η)⁢V⁢(η)⁢e−μ⁢τV−βV⁢(η)⁢V⁢(η),
0= KI⁢∫ΩJ⁡(η−ϵ)⁢(I⁡(ϵ)−I⁡(η))⁢𝑑ϵ+β1⁢(η)⁢V⁢(η)−βI⁢(η)⁢I⁢(η)−γ1⁢(η)⁢I⁢(η)⁢R⁢(η),
0= KR⁢∫ΩJ⁡(η−ϵ)⁢(R⁡(ϵ)−R⁡(η))⁢𝑑ϵ+ζ⁡(η)⁢I⁢(η)⁢R⁢(η)−βR⁢(η)⁢R⁢(η),η∈Ω-

Clearly, (2) has a unique VFSS H0=(λ1⁢(η)βS⁢(η),0,0,0),η∈Ω¯ . We linearize (2) in order to get the basic reproduction number ℜ0 corresponding to (2) at H0, we have

(9) ∂V∂ι= β⁡(η)⁢λ1⁢(η)βS⁢(η)⁢I⁢(ι−τv,η)⁢e−μ⁢τV+α⁡(η)⁢λ1⁢(η)βS⁢(η)⁢V⁢(ι−τv,η)⁢e−μ⁢τV−βV⁢(η)⁢V⁢(ι,η),
∂I∂ι= KI⁢∫ΩJ⁡(η−ϵ)⁢(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,η∈Ω-.

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

{∂∂ι(VI)=𝐀(VI), η∈Ω,ι≥0,∂I∂υ=0, η∈∂Ω, ι≥0,

we define the following linear operators ℬ,ℱ:ℤ→ℤ⁡(ℤ:=C⁡(Ω,ℝ2)) ,

(10) ℬ=(−βV⁢(η)0β1⁢(η)AI), ℱ=(β⁡(η)⁢λ1⁢(η)βS⁢(η)⁢e−μ⁢τVα⁡(η)⁢λ1⁢(η)βS⁢(η)⁢e−μ⁢τV00),

and the linear operator 𝐀:=ℬ+ℱ

We designate 𝐓ℬ⁢(ι) as the semigroup that ℬ generates. Therefore, we can define ℜ0 as the spectral radius of 𝒦, which is

ℜ0=𝔯⁡(𝒦).

Here, the next generation operator, 𝒦:=−ℱ⁢ℬ−1, can be written as

𝒦⁡(Θ)⁢(η)=ℱ⁡(η)⁢∫0∞𝐓B⁢(ι)⁢(Θ)⁢(η)⁢𝑑ι,Θ∈C⁡(Ω-,ℝ2),η∈Ω.-

To derive a variational expression for the basic reproduction number ℜ0, we obtain the following result.

Lemma .

ℜ0=𝔯⁡(−ℱ⁢ℬ−1)=r⁡(dV+dI) with dV=β⁢λ1βS⁢βV⁢e−μ⁢τV and dI=−α⁢β1⁢e−μ⁢τV⁢λ1⁢(x)βS⁢βV⁢AI−1.

Proof.

Let ϕ=ℱ⁢Ψ and Ψ=−ℬ−1⁢Θ. Clearly,

{βVΨ1=Θ1 βI⁢Ψ1−AI⁢Ψ2=Θ2.

Solving this system, we get

Ψ1 = Θ1βV,
Ψ2 = AI−1⁢(βI⁢Ψ1−Θ2).

Consequently,

ϕ1 = d1⁢Ψ1+d2⁢Ψ2,
ϕ2 = 0,

with d1=β⁢λ1βS⁢βV⁢e−μ⁢τV+α⁢e−μ⁢τV⁢λ1⁢β1βS⁢βV⁢AI−1, and d2=−AI−1⁢α⁢λ1βS⁢e−μ⁢τV. Hence, we can write

−ℱ⁢ℬ−1⁢(Θ1Θ2)=(d1⁢Ψ1+d2⁢Ψ20).

Iteratively, we get

(−ℱ⁢ℬ−1)n⁢(Θ1Θ2)=(d1n⁢Ψ1+d1n−1⁢d2⁢Ψ20).

Hence ‖dIn‖≤|(−ℱ⁢ℬ−1)n≤|d1n−1|(‖d1‖+‖d2‖). By applying the squeeze theorem and Gelfand’s formula, we obtain r⁡(−ℱ⁢ℬ−1)=𝔯⁡(d1). Since d1=dV+dI, the proof is achieved. From Section 4, we get

(11) ℜ0=𝔯⁡(dV+dI),

where dV=β⁢λ1βS⁢βV is the cell-to-cell transmission’s next generation operator, and dI=α⁢λ1⁢β1βS⁢βV⁢AI−1 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) ℜ0V=𝔯⁡(dV)=maxη∈Ω-⁢β⁡(η)⁢λ1⁢(η)⁢e−μ⁢τVβS⁢(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) ℜ0I=𝔯⁡(dI)=supΘ∈L2,Θ≠0⁢∫Ωα⁡(η)⁢λ1⁢(η)⁢β⁢(η)⁢e−μ⁢τVβS⁢(η)⁢βV⁢(η)⁢Θ2⁢(η)⁢𝑑ηKI⁢∫Ω∫Ω12⁢J⁢(η−ϵ)⁢(Θ⁡(ϵ)−Θ⁡(η))2⁢𝑑ϵ⁢𝑑η+∫ΩβI⁢(η)⁢Θ2⁢(η)⁢𝑑η.

Clearly, ℜ0≤ ℜ0V +ℜ0I . The equality holds if β⁡(η)⁢λ1⁢(η)βS⁢(η)⁢βV⁢(η) is space independent.
The basic reproduction number ℜ1 for the antibody response is determined by

ℜ1=supΘ∈L2,Θ≠0⁢∫Ωζ⁡(η)⁢I∗⁣∗⁢(η)⁢Θ2⁢(η)⁢𝑑ηKR⁢∫Ω∫Ω12⁢J⁢(η−ϵ)⁢(Θ⁡(ϵ)−Θ⁡(η))2⁢𝑑ϵ⁢𝑑η+∫ΩβR⁢(η)⁢Θ2⁢(η)⁢𝑑η.

∎

It is evident that ℜ1 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.

ℜ1=ζ⁡(η)⁢I∗⁣∗⁢(η)βR⁢(η).
Lemma .

ℜ0−1 and δ⁡(𝐀) have the same sign.

5. Extinction scenario of virus

Here, we show that for ℜ0<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 ℜ0<1, and unstable if ℜ0>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)⁢e−λ⁢τV⁢e−μ⁢τV+α⁡(η)⁢λ1⁢(η)βS⁢(η)⁢Θ2⁢(η)⁢e−λ⁢τV⁢e−μ⁢τV−βV⁢(η)⁢Θ2⁢(η)
=λ⁢Θ2⁢(η),
KI⁢∫ΩJ⁡(η−ϵ)⁢(Θ3⁢(ι,ϵ)−Θ3⁢(ι,η))⁢𝑑ϵ+β1⁢(η)⁢Θ2⁢(η)−βI⁢(η)⁢Θ3⁢(η)
=λ⁢Θ3⁢(η),

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

(15) β⁡(η)⁢λ1⁢(η)βS⁢(η)⁢Θ3⁢(η)⁢e−μ⁢τV+α⁡(η)⁢λ1⁢(η)βS⁢(η)⁢Θ2⁢(η)⁢e−μ⁢τV−βV⁢(η)⁢ϕ2⁢(η)= λ⁢Θ2⁢(η),
KI⁢∫ΩJ⁡(η−ϵ)⁢(Θ3⁢(ι,ϵ)−Θ3⁢(ι,η))⁢𝑑ϵ+β1⁢(η)⁢Θ2⁢(η)−βI⁢(η)⁢Θ3⁢(η)= λ⁢Θ3⁢(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 ω0≤μ0<0. As a result, H0stability (or instability) is determined by δ⁡(A). Section 4 gives us that ℜ0−1 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 ℜ1≤1<ℜ0, 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 ℜ0 <1, then there exists a ρ>0 that satisfies ρ<1. It may be written as

(16) ρ=supΘ∈L2,Θ≠0{ 1KI⁢∫Ω∫Ω12⁢J⁢(η−ϵ)⁢(Θ⁡(η)−Θ⁡(ϵ))⁢Θ⁢(η)⁢𝑑ϵ⁢𝑑η+∫ΩβI⁢(η)⁢Θ2⁢(η)⁢𝑑η×
×∫Ωα⁡(η)⁢λ1⁢(η)⁢βI⁢(η)⁢e−μ⁢τVβS⁢(η)⁢βV⁢(η)−β∗⁢e−μ⁢τV⁢λ⁢(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)⁢e−μ⁢τV)⁢Θ10⁢(η)+β1⁢(η)⁢Θ20⁢(η),η∈Ω-,
0= 1ρ⁢β⁢(η)⁢e−μ⁢τV⁢λ1⁢(η)βS⁢(η)⁢Θ10⁢(η)+KI⁢∫ΩJ⁡(−ϵ)⁢(Θ20⁢(ι,ϵ)−Θ20⁢(ι,η))⁢𝑑ϵ−βI⁢(η)⁢Θ20⁢(η),
η∈Ω-.
Proof.

Notice that for ℜ0 <1, we obtain that ℜ⁡(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)⁢e−μ⁢τV0β1⁢(η)AI)+(0β⁡(η)⁢e−μ⁢τV⁢λ1⁢(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:=−𝖥𝖡−1 and applying Theorem 3.5 in [13], we conclude that if ℛ0<1, then δ⁡(𝐀)<0 and, consequently, ρ<1. We then consider an eigenvalue ϱ of 𝐀 that solves the following problem

OPENϱ⁢Θ⁢(η)=A⁡[Θ]⁢(η),η∈Ω,Θ=(Θ1⁢(η),Θ2⁢(η))T∈C⁡(CLOSE-⁢Ω,ℝ2).

Next, we claim that the problem

(19) 𝖡⁡[Θ]⁢(η)=ϱ⁢𝖥⁢[Θ]⁢(η),η∈Ω-,

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

ρ=𝔯⁡(−𝖥𝖡−1)=𝔯⁡(−𝖡−1⁢𝖥)=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

(γ⁢I−𝖡)−1⁢Θ=∫0∞e−ϵ⁢ι⁢T𝖡⁢(ι)⁢𝑑ι,Θ∈𝔇⁡(𝖡).

Assume that γ=0. Next,

−𝖡−1⁢Θ=∫0∞TB⁢(ι)⁢𝑑ι,Θ∈𝔇⁡(𝖡).

Additionally, we get

𝖥⁡(−𝖡−1)⁢Θ=𝐊⁡[Θ]⁢(η).

Now, we set Θ =−𝖡⁢Θ-, then we get

𝖥⁢Θ-=−𝐊⁡(η)⁢𝖡⁢Θ-.

Therefore,

−𝖡⁢Θ-=𝖥⁢Θ-𝐊⁡(η),η∈Ω-.

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∗=(Θ1∗CLOSE, OPENΘ2∗),η∈Ω- is the corresponding positive eigenfunction , we have

(20) 0 =−(βV⁢(η)−α⁡(η)⁢λ1⁢(η)βS⁢(η)⁢e−μ⁢τV)⁢Θ1∗⁢(η)+β1⁢(η)⁢Θ2∗⁢(η),η∈Ω-,
0 =ϱ∗⁢β⁢(η)⁢e−μ⁢τV⁢λ1⁢(η)βS⁢(η)⁢Θ1∗⁢(η)+KI⁢∫ΩJ⁡(η−ϵ)⁢(Θ2∗⁢(ι,ϵ)−Θ2∗⁢(ι,η))⁢𝑑ϵ−βI⁢(η)⁢Θ2∗⁢(η),
η∈Ω-.

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

Θ1∗(η)=β1⁢(η)⁢βS⁢(η)βV⁢(η)⁢βS⁢(η)−α⁡(η)⁢λ1⁢(η)⁢e−μ⁢τVΘ2∗(η), η∈Ω-.

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

(21) 0= ϱ∗⁢β⁢(η)⁢e−μ⁢τV⁢λ1⁢(η)βS⁢(η).β1⁢(η)⁢βS⁢(η)βV⁢(η)⁢βS⁢(η)−α⁡(η)⁢λ1⁢(η)⁢e−μ⁢τV⁢Θ2∗⁢(η)
+KI∫ΩJ(η−ϵ)(Θ2∗(ι,ϵ)−Θ2∗(ι,η))dϵ−βI(η)Θ2∗(η),
η∈Ω-,

hence,

KI⁢∫ΩJ⁡(η−ϵ)⁢(Θ2∗⁢(ι,ϵ)−Θ2∗⁢(ι,η))⁢𝑑ϵ−βI⁢(η)⁢Θ2∗⁢(v)
+ϱ∗⁢β⁡(η)⁢e−μ⁢τV⁢λ1⁢(η)⁢β1⁢(η)βV⁢(η)⁢βS⁢(η)−α⁡(η)⁢λ1⁢(η)⁢e−μ⁢τV⁢Θ2∗⁢(η)=0.

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

(22) ρ=supΘ∈L2,Θ≠0{ 1KI⁢∫Ω∫Ω12⁢J⁢(η−ϵ)⁢(Θ⁡(η)−Θ⁡(ϵ))⁢Θ⁢(η)⁢𝑑ϵ⁢𝑑η+∫ΩβI⁢(η)⁢Θ2⁢(η)⁢𝑑η×
×∫Ωα⁡(η)⁢λ1⁢(η)⁢β1⁢(η)⁢e−μ⁢τVβS⁢(η)⁢βV⁢(η)−β∗⁢e−μ⁢τV⁢λ1⁢(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)⁢e−μ⁢τV)⁢Θ10⁢(η)+β1⁢(η)⁢Θ20⁢(η) =0,η∈Ω-
1ρ⁢β⁢(η)⁢e−μ⁢τV⁢λ1⁢(η)βS⁢(η)⁢Θ10⁢(η)+KI⁢∫ΩJ⁡(η−ϵ)⁢(Θ20⁢(ι,ϵ)CLOSE
OPEN−Θ20⁢(ι,η))⁢d⁢ϵ−βI⁢(η)⁢Θ20⁢(η) =0,η∈Ω-

∎

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 ℜ0 be defined by (22) and (23). Then H0 is globally stable in ℂ if ℜ0<1.

Proof.

It has been proved that lim supι→∞S⁡(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 ℜ0<1, then for ρθ small enough there exist positive functions (Θ1θ⁢(η),Θ2θ⁢(η)) satisfying the following system:

(24) 0= −(βV⁢(η)−α⁡(η)⁢e−μ⁢τV⁢(λ1⁢(η)βS⁢(v)+θ))⁢Θ1θ⁢(η)+β1⁢(η)⁢Θ2θ⁢(η),
0= 1ρθ⁢β⁢(η)⁢e−μ⁢τV⁢(λ1⁢(η)βS⁢(η)+θ)⁢Θ1θ⁢(η)
+KI∫ΩJ(η−ϵ)(Θ2θ(ϵ)−Θ2θ(η))dϵ−βI(η)Θ2θ(η),η∈Ω-.

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)= e−μ⁢τV⁢∫ΩΘ1θ⁢(η)⁢β⁢(η)⁢(λ1⁢(η)βS⁢(η)+θ)⁢∫0τVI⁡(τ−s,η)⁢𝑑s
+Θ1θ(η)α(η)(λ1⁢(η)βS⁢(η)+θ)∫0τVV(ι−s,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)∂ι⁢𝑑η
= ∫Ω{Θ1θ(η)[β(η)S(ι−τv,η)I(ι−τv,η)e−μ⁢τV
+α(η)S(ι−τv,η)V(ι−τv,η)e−μ⁢τV−βV(η)V(ι,η)]}dη
+∫Ω{Θ2θ(η)[KI∫ΩJ(η−ϵ)(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(ι−τv,η)e−μ⁢τV
+α(η)(λ1⁢(η)βS⁢(η)+θ)V(ι−τv,η)e−μ⁢τV−βV(η)V(ι,η)]}dη
+∫Ω{Θ2θ(η)[KI∫ΩJ(η−ϵ)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⁡(η−ϵ)⁢𝑑ϵ. Using (24), we get

ϑ′⁢(ι)≤
≤ ∫Ω{Θ1θ(η)[β(η)(λ1⁢(η)βS⁢(η)+θ)I(ι−τv,η)e−μ⁢τV
+α(η)(λ1⁢(η)βS⁢(η)+θ)V(ι−τv,η)e−μ⁢τV]}dη
+∫Ω{Θ2θ(η)[KI∫ΩJ(η−ϵ)I(ι,ϵ)dϵ+β1(η)V(ι,η)]}dη
−∫ΩV(ι,η){α(η)λ1⁢(η)βS⁢(η)e−μ⁢τVϕ1θ(η)+β1(η)Θ2θ(η)}dη
−∫ΩI(ι,η){1ρθβ(η)e−μ⁢τV(λ1⁢(η)βS⁢(η)+θ)Θ1θ(η)+KI∫ΩJ(η−ϵ)Θ2θ(ϵ)dϵ}dη.

Using the assumptions on J, we have

∫Ω∫ΩJ⁡(η−ϵ)⁢Θ2θ⁢(ϵ)⁢I⁢(ι,η)⁢𝑑ϵ⁢𝑑η=∫Ω∫ΩJ⁡(η−ϵ)⁢Θ2θ⁢I⁢(ι,ϵ)⁢𝑑η⁢𝑑ϵ.

Therefore, some simplification gives

ϑ′⁢(ι) ≤ ∫ΩΘ1θ⁢(η)⁢β⁢(η)⁢(λ1⁢(η)βS⁢(η)+θ)⁢e−μ⁢τV⁢[−I⁡(ι,η)ρθ+I⁡(ι−τv,η)]⁢𝑑η
+∫ΩΘ1θ(η)α(η)(λ1⁢(η)βS⁢(η)+θ)e−μ⁢τV[V(ι−τv,η)−V(ι,η)].

ϑτV′⁢(ι) is given by

ϑτV′⁢(τ)= e−μ⁢τV⁢∫ΩΘ1θ⁢(η)⁢β⁢(η)⁢(λ1⁢(η)βS⁢(η)+θ)⁢∂∂ι⁢∫0τVI⁡(ι−s,η)⁢𝑑s
+Θ1θ(η)α(η)(λ1⁢(η)βS⁢(η)+θ)∂∂ι∫0τVV(ι−s,η)dsdη
= e−μ⁢τV⁢∫ΩΘ1θ⁢(η)⁢β⁢(η)⁢(λ1⁢(η)βS⁢(η)+θ)⁢∫0τV∂∂t⁢I⁢(ι−s,η)⁢𝑑s
+Θ1θ(η)α(η)(λ1⁢(η)βS⁢(η)+θ)∫0τV∂∂ιV(ι−s,η)dsdη
= e−μ⁢τV⁢∫ΩΘ1θ⁢(η)⁢β⁢(η)⁢(λ1⁢(η)βS⁢(η)+θ)⁢∫0τV(−∂∂s)⁢I⁢(ι−s,η)⁢𝑑s
+Θ1θ(η)α(η)(λ1⁢(η)βS⁢(η)+θ)∫0τV(−∂∂s)V(ι−s,η)dsdη
= −{e−μ⁢τV∫ΩΘ1θ(η)β(η)(λ1⁢(η)βS⁢(η)+θ)I(ι−s,η)∣0τV
+Θ1θ(η)α(η)(λ1⁢(η)βS⁢(η)+θ)V(ι−s,η)∣0τVdη}
= e−μ⁢τV⁢∫ΩΘ1θ⁢(η)⁢β⁢(η)⁢(λ1⁢(η)βS⁢(η)+θ)⁢(I⁡(ι,η)−I⁡(ι−τV,η))
+Θ1θ⁢(η)⁢α⁢(η)⁢(λ1⁢(η)βS⁢(η)+θ)⁢(V⁡(ι,η)−V⁡(ι−τV,η))⁢d⁢η

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

E′⁢(ι)≤e−μ⁢τV⁢∫ΩΘ1θ⁢(η)⁢β⁢(η)⁢I⁢(ι,η)⁢(λ1⁢(η)βS⁢(η)+θ)⁢(1−1ρθ)⁢𝑑η≤0.

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 η∈Ω as ι⟶∞.
We further obtain that S⁡(ι,η)→ λ1⁢(η)βS⁢(η) uniformly for η∈Ω as ι→∞. To sum up, the theorem is proved. ∎

6. Uniform persistence

The following spaces should be defined:

ξ0 = {𝕌∈X+:V(⋅,η)>0,I(⋅,η)>0 ,η∈Ω¯};
∂ξ0 = {𝕌∈X+:V(⋅,η)≡0,I(⋅,η)≡0 ,η∈Ω¯};
M∂ = {𝕌0∈∂ξ0:Φ(ι,𝕌0)∈∂ξ0 for ι≥0}.

Then, we obtain the following results.

Theorem 5.

Assume that ℜ1≤1<ℜ0. 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)}ι≥0 restricted to ∂ξ0.

For 𝕌0∈∂ξ0, (2) becomes

(26) {d⁢Sd⁢ι(ι,η)=λ1(η)−βS(η)S(ι,η), V(⋅,η)=0,I(⋅,η)=0, ⁢ι>0,η∈Ω-.

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

Claim .

For any 𝕌0∈ξ0, we have 𝕌>0 for all η∈Ω- , and ι>0. We consider the problem

∂i⁡(ι,η)∂ι =KI⁢∫ΩJ⁡(η−ϵ)⁢(i⁡(ι,ϵ)−i⁡(ι,η))⁢𝑑ϵ−βI⁢(η)⁢i⁢(ι,η),ι>0,η∈Ω-.
i⁡(0,η) =I0⁢(η),η∈Ω.-

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, η∈Ω¯. This proof follows the approach of Proposition 2.2 in [19]. We define the operator Υ:ℂ⁡(Ω-)→ℂ⁡(Ω-)

Υ⁢i⁢(ι,η)=KI⁢∫ΩJ⁡(η−y)⁢(I⁡(ι,y)−I⁡(ι,η))⁢𝑑y,η∈Ω-.

Clearly, Υ is continuous, and {eΥ⁢ι}ι≥0 forms a uniformly continuous, positive semigroup on ℂ⁡(Ω¯) generated by Υ. , where for ι≥0, we get eΥ⁢ι =∑n=0∞ιn⁢Υnn!. Further,

eΥ⁢τ⁢(I0⁢(s,η))=∑n=0∞τn⁢Υn⁢(I0⁢(s,η))n!,ι>0,s∈[−τ,0].

Since i⁡(τ,η)≠0, we have Υn+1⁢(I0⁢(s,η))=J⁡(η−y)⁢Υn⁢I0⁢(s,υ)⁢d⁢υ,η∈Ω¯, s∈[−τ,0], and n≥1. The iteration process implies the existence of n0 such that Υn+1⁢(I⁡(0,η))>0,η∈Ω¯ and n≥n0. Hence,Υn⁢(I0⁢(s,η))>0,η∈Ω¯,s∈[−τ,0]. Consequently,

i⁡(ι,η) =e−(KI−βI⁢(η))eΥ⁢τ(I0(s,η))+∫0τ e−(KI−βI⁢(η))⁢(τ−σ) eΥ⁡(τ−σ)i(ι,η)dσ,
≥e−(KI−βI⁢(η))⁢eΥ⁢τ⁢(I0⁢(s,η))>0,

for ι>0,η∈Ω¯. 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 ℜ1≤1<ℜ0. In order to prove this assertion, we demonstrate that there exists a sufficiently small θ1>0 such that the semiflow {Ξ⁡(ι)}ι≥0 satisfies

lim supι→∞‖Ξ⁡(ι,𝕌0)−H0‖≥θ1.

We use contradiction arguments to demonstrate this result. Suppose that lim supι→∞‖𝚵⁡(ι,𝕌0)−H0‖≤θ1. Hence, there exists ι1 such that 𝕌⁡(ι,𝕌0) satisfies

λ1⁢(η)βS⁢(η)−θ1≤S⁡(ι,η)≤λ1⁢(η)βS⁢(η)+θ1,0≤V⁡(ι,η)≤θ1,ι≥ι1,η∈Ω¯.

Since ℜ1≤1<ℜ0, we have Sθ⁢(A)>0. Then, there exists ε sufficiently small satisfying

χ⁢ϕ⁢(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,η∈Ω.

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⁢(ι−τv,η)⁢e−μ⁢τV
+α⁡(η)⁢(λ1⁢(η)βS⁢(η)+θ1)⁢V⁢(ι−τv,η)⁢e−μ⁢τV−βV⁢(η)⁢V⁢(ι,η),
∂I∂ι⁢(ι,η)= KI⁢∫ΩJ⁡(η−y)⁢(I⁡(ι,y)−I⁡(ι,η))⁢𝑑y+β1⁢(η)⁢V⁢(ι,η)−βI⁢(η)⁢I⁢(ι,η),
ι> ι1,η∈Ω.-

By comparison principle, we obtain

OPEN(V⁡(η),I⁡(η))T=yP⁢eyP⁢(ι−ι1CLOSE)⁢ΘP⁢(η).

There is a contradiction since yP>0.

Claim .

System (2) is strongly uniformly persistent in ξ0 for ℜ1≤1<ℜ0. For the orbit ϰ+⁢(𝕌0):=𝕌ι≥0⁢{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 𝕌0∈∂ξ0, then there is no cycle in ∂ξ0 from H0 to H0. We obtain Ws⁢(H0)∩ξ0=∅, with Ws⁢(E0)={𝕌0∈X:limι→∞⁢Ξ⁢(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 ℛ1≤1<ℛ0. 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)⁢e−μ⁢τV+α⁡(η)⁢S∗⁢(η)⁢V∗⁢(η)⁢e−μ⁢τV−βV⁢(η)⁢V∗⁢(η)η∈Ω-.
0= KI⁢∫ΩJ⁡(η−ϵ)⁢(I∗⁢(ϵ)−I∗⁢(η))⁢𝑑ϵ+β1⁢(η)⁢V∗⁢(η)−βI⁢(η)⁢I∗⁢(η).

7. Global behavior analysis

Theorem 6.

If ℜ1≤1< ℜ0, 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)+ϱ1⁢τV,

with

ϱ1⁢(ι)= ∫Ωζ1(η){e−μ⁢τVP∗(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

ϱ1⁢τV= ∫Ωe−μ⁢τV⁢ζ1⁢(η)⁢∫0τVβ⁡(η)⁢S∗⁢(η)⁢I∗⁢(η)⁢H⁢(S⁡(ι−ϑ)⁢I⁢(ι−ϑ)S∗⁢I∗)
+α⁡(η)⁢S∗⁢(η)⁢V∗⁢(η)⁢H⁢(S⁡(ι−ϑ)⁢V⁢(ι−ϑ)S∗⁢V∗)⁢d⁢ϑ⁢d⁢η,

where H⁡(t)=t−1−ln⁡(t), for t>0, ζ1⁢(η)=2⁢e−μ⁢τV⁢β1⁢(η)⁢V∗⁢(η)β⁡(η)⁢S∗⁢(η),ζ2⁢(x)=2⁢e−μ⁢τV⁢β⁡(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⁡(ι−τV,η),IτV=I⁡(ι−τV,η),
VτV =V⁡(ι−τV,η),S∗=S∗⁢(η),I∗=I∗⁢(η),V∗=V∗⁢(η).

First, we compute ϱ1′⁢(ι), which is

ϱ1′⁢(ι)=
= ∫Ωζ1⁢(η)⁢{e−μ⁢τV⁢(1−S∗S)⁢∂S∂ι+(1−V∗V)⁢∂V∂ι+ζ2⁢(η)⁢(1−I∗I)⁢∂I∂ι}⁢𝑑η
= ∫Ωζ1(η){e−μ⁢τV(1−S∗S)[λ1−βSS−βSI−αSV]
+(1−V∗V)⁢[β⁢SτV⁢IτV⁢e−μ⁢τV+α⁢SτV⁢VτV⁢e−μ⁢τV−βV⁢V]
+ζ2(η)(−I∗I)[KI∫ΩJ(η−ϵ)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)S∗I∗+α(η)S∗V∗, βV(η)V∗ =β(η)S∗I∗e−μ⁢τV+α(η)S∗V∗e−μ⁢τV, (βI(η)+KI)I∗ =KI∫ΩJ(η−ϵ)I∗(ϵ)dϵ+β1(η)V∗,⁢η∈Ω-.

Thus, after some simplification, ϱ1′⁢(ι) becomes

ϱ1′⁢(ι)=
= ∫Ωζ1(η){e−μ⁢τVβS(η)S∗(2−S∗S−SS∗)
+e−μ⁢τV⁢β⁢(η)⁢S∗⁢I∗⁢[2−S∗S−S⁢IS∗⁢I∗+II∗−V∗V−SτV⁢IτVS∗⁢I∗−SτV⁢IτV⁢V∗S∗⁢I∗⁢V]
+e−μ⁢τV⁢α⁢(η)⁢S∗⁢V∗⁢[2−S∗S−S⁢VS∗⁢V∗+SτV⁢VτVS∗⁢V∗−SτV⁢VτVS∗⁢V]
+KI⁢ζ2⁢(η)⁢[(1−I∗I)⁢KI⁢∫ΩJ⁡(η−ϵ)⁢I⁢(ι,ϵ)⁢𝑑ϵ+(1−II∗)⁢∫ΩJ⁡(η−ϵ)⁢I∗⁢(ϵ)⁢𝑑ϵ]
+ζ2(η)β1(η)V∗(1+VV∗−II∗−V∗⁢IV⁢I∗)}dη.

Notice that ζ2⁢(η)⁢β1⁢(η)⁢V∗= β⁡(η)⁢S∗⁢I∗⁢e−μ⁢τV and ζ1⁢(η)⁢ζ2⁢(η)=2⁢I∗. Then, we obtain

ϱ1′⁢(ι)= ∫Ωζ1(η){e−μ⁢τVβS(η)S∗(2−S∗S−SS∗)
+e−μ⁢τV⁢β⁢(η)⁢S∗⁢I∗⁢[3−S∗S−S⁢IS∗⁢I∗+SτV⁢IτVS∗⁢I∗−SτV⁢IτV⁢V∗S∗⁢I∗⁢V−V∗⁢IV⁢I∗]
+e−μ⁢τVα(η)S∗V∗[2−S∗S−S⁢VS∗⁢V∗+SτV⁢VτVS∗⁢V∗−SτV⁢VτVS∗⁢V]}dη
+2KI∫ΩI∗{(1−I∗I)∫ΩJ(η−ϵ)I∗(ϵ)dϵ
+(1−II∗)∫ΩJ(η−ϵ)I(ι,ϵ)dϵ}dη
= ∫Ωζ1(η){e−μ⁢τVβS(η)S∗(2−S∗S−SS∗)
+e−μ⁢τV⁢β⁢(η)⁢S∗⁢I∗⁢[3−S∗S−S⁢IS∗⁢I∗+SτV⁢IτVS∗⁢I∗−SτV⁢IτV⁢V∗S∗⁢I∗⁢V−V∗⁢IV⁢I∗]
+e−μ⁢τVα(η)S∗V∗[2−S∗S−S⁢VS∗⁢V∗−SτV⁢VτVS∗⁢V∗−SτV⁢VτVS∗⁢V]}dη
+KI∫Ω∫ΩJ(η−ϵ)I∗(ϵ)I∗(η){2−I∗⁢(η)I⁡(ι,ϵ)I⁡(ι,ϵ)I∗⁢(ϵ)−I∗⁢(ϵ)I⁡(ι,ϵ)I⁡(ι,ϵ)I∗⁢(η)}dηdϵ.

Alternatively, ϱ1⁢τV′ is computed as follows:

ϱ1⁢τV′= ∂∂ι⁢∫Ωe−μ⁢τV⁢ζ1⁢(η)⁢∫0τVβ⁡(η)⁢S∗⁢I∗⁢H⁢(Sϑ⁢IϑS∗⁢I∗)+α⁡(η)⁢S∗⁢V∗⁢H⁢(Sϑ⁢VϑS∗⁢V∗)⁢𝑑ϑ⁢𝑑η
= ∫Ωe−μ⁢τVζ1(η){β(η)S∗I∗[H(S⁢IS∗⁢I∗)−H(SτV⁢IτVS∗⁢I∗)]
+α(η)S∗V∗[H(S⁢VS∗⁢V∗)−H(SτV⁢VτVS∗⁢V∗)]}dη.

Adding ϱ1′⁢(ι) and ϱ1⁢τV′, we find

E∗′⁢(ι)= ∫Ωζ1(η){e−μ⁢τVβS(η)S∗(2−S∗S−SS∗)
+e−μ⁢τV⁢β⁢(η)⁢S∗⁢I∗⁢[3−S∗S−S⁢IS∗⁢I∗−SτV⁢IτVS∗⁢I∗−SτV⁢IτV⁢V∗S∗⁢I∗⁢V−V∗⁢IV⁢I∗]
+e−μ⁢τVα(η)S∗V∗[2−S∗S−S⁢VS∗⁢V∗−SτV⁢VτVS∗⁢V∗−SτV⁢VτVS∗⁢V]}dη
+KI∫Ω∫ΩJ(η−ϵ)I∗(ϵ)I∗(η){2−I∗⁢(η)I⁡(ι,ϵ)I⁡(ι,ϵ)I∗⁢(ϵ)−I∗⁢(ϵ)I⁡(ι,ϵ)I⁡(ι,ϵ)I∗⁢(η)}dηdϵ
+∫Ωe−μ⁢τVζ1(η){β(η)S∗I∗[H(S⁢IS∗⁢I∗)−H(SτV⁢IτVS∗⁢I∗)]
+α(η)S∗V∗[H(S⁢VS∗⁢V∗)−H(SτV⁢VτVS∗⁢V∗)]}dη
= ∫Ωe−μ⁢τVζ1(η){βS(η)S∗(2−S∗S−SS∗)
+β⁡(η)⁢S∗⁢I∗⁢[−H⁡(S∗S)−H⁡(SτV⁢IτV⁢V∗S∗⁢I∗⁢V)−H⁡(I∗⁢VI⁢V∗)]
+α(η)S∗V∗[−H(S∗S)−H(SτV⁢VτVS∗⁢V)]}dη
+KI∫Ω∫ΩJ(η−ϵ)I∗(ϵ)I∗(η){2−I∗⁢(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)={2.2523⁢e1η2−1,−1<η<1,0,otherwise.

This ensures that

∫ℝJ⁡(η)⁢𝑑η≈1,

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

Refer to caption
Figure 1. Kernel function J.

We consider the following set of parameters:

μ =10−1, βS⁢(η) =15⋅10−1, βV⁢(η) =12⋅10−1, βI⁢(η) =13⋅10−1,
λ1⁢(η) =29⋅10−1, τV =98⋅10−1, β⁡(η) =β¯, α⁡(η) =1,
KI =5⋅10−2, ζ =98⋅10−1.
S0⁢(ι,η) =3⋅10−1+5⋅10−2⁢cos⁡(η),
V0⁢(ι,η) =41⋅10−2+10−2⁢cos⁡(η)+10−3⁢cos⁡(ι),
I0⁢(ι,η) =71⋅10−2+10−1⁢cos⁡(η)+10−2⁢cos⁡(3⁢ι).

First, we take β¯=47⋅10−2. Then ℛ0=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 ℛ0=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¯=23⋅10−2 and βR⁢(η)=187⋅10−1, where ℛ0=3.5874>1 and ℛ1=0.3956<1; see Figure 3.

Refer to caption
Refer to caption
Figure 3. When ℛ0=3.5874>1 and ℛ1=0.3956<1, the immunity-free infected steady state H1 is globally asymptotically stable.

Finally, for βR⁢(η)=53⋅10−1 and ℛ1=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 ℛ1=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 ℜ0 and the basic reproduction number of immunity ℜ1. By analyzing the characteristic equations and constructing Lyapunov functionals, we have obtained the following conclusions: if ℜ0<1, then the infection-free steady state H0 is globally asymptotically stable; if ℜ1⩽1<ℜ0, 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.

References

  • [1] M.A. Nowak, C.R.M. Bangham, Population dynamics of immune responses to persistent viruses, Science, 272 (1996), pp. 74–79.
  • [2] J. Lin, R. Xu, X. Tian, Threshold dynamics of an HIV-1 virus model with both virus-to-cell and cell-to-cell transmissions, intracellular delay, and humoral immunity, Appl. Math. Comput., 315 (2017), pp. 516–530.
  • [3] J. Garcia-Melian, J.D. Rossi, On the principal eigenvalue of some nonlocal diffusion problems, J. Differential Equations, 246 (2009), pp. 21–38.
  • [4] K.J. Engel and R. Nagel, One-Parameter Semigroups for Linear Evolution Equations, Graduate Texts in Mathematics, Springer, New York, 2000.
  • [5] A. Pazy, Semigroups of Linear Operators and Applications to Partial Differential Equations, Springer, New York, 1983.
  • [6] S. Han and C. Lei, Stability of equilibria of a diffusive SEIR epidemic model with nonlinear incidence, Appl. Math. Lett., 98 (2019), pp. 114–120.
  • [7] G. Webb, Theory of Nonlinear Age-Dependent Population Dynamics, CRC Press, 1985.
  • [8] F.Y. Yang and W.T. Li, Dynamics of a nonlocal dispersal SIS epidemic model, Commun. Pure Appl. Anal., 16 (2017), p. 781.
  • [9] K. Allali, S. Harroudi, D.F.M. Torres, Analysis and optimal control of an intracellular delayed HIV model with CTL immune response, J. Math. Comput. Sci., 12 (2018), pp. 111–127.
  • [10] G. Zhao and S. Ruan, Spatial and temporal dynamics of a nonlocal viral infection model, SIAM J. Appl. Math., 78 (2018) no. 4, pp. 1954–1980.
  • [11] J.K. Hale, Asymptotic Behavior of Dissipative Systems, Mathematical Surveys and Monographs, vol. 25, American Mathematical Society, Providence, RI, 1988.
  • [12] P. Magal and X.Q. Zhao, Global attractors and steady states for uniformly persistent dynamical systems, SIAM J. Math. Anal., 37 (2005) no. 1, pp. 251–275.
  • [13] H.R. Thieme, Spectral bound and reproduction number for infinite-dimensional population structure and time heterogeneity, SIAM J. Math. Anal., 70 (2009), pp. 188–211.
  • [14] X.Q. Zhao, Dynamical Systems in Population Biology, vol. 16, Springer, New York, 2017.
  • [15] A.M. Elaiw, A.A. Raezah, K. Hattaf, Stability of HIV-1 infection with saturated virus-target and infected-target incidences and CTL immune response, Int. J. Biomath., 10 (2017), 1750070.
  • [16] W. Desch and W. Schappacher, Linearized stability for nonlinear semigroups, in Differential Equations in Banach Spaces, Lecture Notes in Mathematics, A. Favini and E. Obrecht (eds.), Springer-Verlag, Berlin, Heidelberg, 1986, pp. 61–67.
  • [17] K. Hattaf and N. Yousfi, A generalized HBV model with diffusion and two delays, Comput. Math. Appl., 69 (2015), pp. 31–40.
  • [18] H.L. Smith and X.Q. Zhao, Robust persistence for semidynamical systems, Nonlinear Anal., 47 (2001) no. 9, pp. 6169–6179.
  • [19] W. Shen and A. Zhang, Spreading speeds for monostable equations with nonlocal dispersal in space periodic habitats, J. Differential Equations, 249 (2010) no. 4, pp. 747–795.
  • [20] W. Wang and X.Q. Zhao, Basic reproduction numbers for reaction–diffusion epidemic models, SIAM J. Appl. Dyn. Syst., 11 (2012), pp. 1652–1673.
  • [21] C. Qin, Y. Chen, X. Wang, Global dynamics of a delayed diffusive virus infection model with cell-mediated immunity and cell-to-cell transmission, Math. Biosci. Eng., 17 (2020) no. 5, pp. 4678–4705.
  • [22] Y. Yang, T. Zhang, J. Zhou, Global attractivity of a time-delayed viral infection model with spatial heterogeneity, Appl. Math. Lett., 116 (2021), 107035.
  • [23] S. Djilali, Threshold asymptotic dynamics for a spatial age-dependent cell-to-cell transmission model with nonlocal dispersal, Discrete Contin. Dyn. Syst., 28 (2023) no. 7, pp. 4108–4143.
  • [24] L. Liu, R. Xu, Z. Jin, Global dynamics of a spatial heterogeneous viral infection model with intracellular delay and nonlocal diffusion, Appl. Math. Model., 82 (2020), pp. 150–167.