1 Introduction
A typical mechanism of mosquito-borne diseases involves a mosquito feeding on the blood of an infected host and ingesting the viruses or parasites, which can then be transferred to the next host bitten by the mosquito. Common mosquito-borne diseases include malaria, dengue, West Nile virus, chikungunya, yellow fever, and Zika, which are directly responsible for more than a million deaths per year. Mathematical models serve as valuable tools for understanding the transmission dynamics of mosquito-borne diseases. Most existing models overlook the compartment of male mosquitoes, focusing solely on female mosquitoes, and typically assume that either the total number of female mosquitoes or their recruitment rate is a constant or a given function. However, the life cycle of mosquitoes is quite complicated, as their reproduction is influenced by various stages of metamorphosis, along with pair formation and mating behaviors [Reference Bai and Zhao3, Reference Hadeler, Waldstatter and Wörz-Busekros8, Reference Qiu, Wei, Shan and Zhu18].
The main purpose of this article is to study mosquito growth by incorporating metamorphic stages (including aquatic and adult stages), pair formation, and the spatial movement of adult mosquitoes in a time-periodic environment. Let
$\Omega $
be a spatial habitat with smooth boundary
$\partial \Omega $
, and we consider a closed environment with the Neumann boundary conditions for subpopulations on the boundary. Let
$u_A(x,t)$
be the population size of the aquatic mosquitoes, and
$u_M(x,t)$
(resp.
$u_F(x,t)$
) be the population size of matured male mosquitoes (resp. matured female mosquitoes). The reproductive rate at location x and time t of adult female mosquitoes is denoted by
$\rho (x,t)$
. According to the law of harmonic mean [Reference Bai and Zhao3, Reference Hadeler, Waldstatter and Wörz-Busekros8, Reference Qiu, Wei, Shan and Zhu18], we set
and the mating function takes the following form:
which satisfies preservation of positivity, homogeneity, and monotonicity. Note that the aquatic mosquito population is constrained by the environmental carrying capacity
$K(x,t)$
, as female mosquitoes lay their eggs in clusters on the surface of stagnant water. As in [Reference Qiu, Wei, Shan and Zhu18], the growth rate of the aquatic mosquito population can be described by
This motivates us to consider the following reaction–diffusion system describing the growth of mosquitoes with pair formation in a time-varying environment:
where
$\Delta $
is the usual Laplacian operator and
$\nu $
is the unit outward normal vector to the boundary
$\partial \Omega $
; D is the diffusion coefficient of matured mosquitoes,
$d(x,t)$
(resp.
$\hat {d}(x,t)$
) represents the natural death rate of aquatic (resp. matured) mosquitoes;
$\gamma (x,t)$
is the maturation rate; and
$\theta $
is the probability of the aquatic mosquitoes maturing into male. With the seasonality, we further assume that there exists a
$T>0$
such that
$w(x,t+T)=w(x,t),\, \forall \, x\in \Omega $
,
$t\in \mathbb {R}$
, and there exists
$\kappa \in (0,1)$
such that
$w \in C^{\kappa ,\frac {\kappa }{2}} (\overline {\Omega } \times [0,T])$
for
$w=\rho ,\ K,\ d,\ \gamma ,\ \hat {d}$
, and all these functions are strictly positive. We should point out that a similar reaction–diffusion model of mosquito growth was studied in [Reference Bai and Zhao3] under the assumption that each of three equations contains a diffusion term. Biologically, aquatic mosquitoes typically stay within specific regions, and their movements can be ignored. Consequently, the first equation in (1.1) does not include a diffusion term, which leads to a lack of compactness for the solution maps of system (1.1).
In order to investigate the influence of the spatial heterogeneity on the spreading of the invading mosquitoes, we also modify system (1.1) into the following reaction–diffusion system in a time–space periodic habitat:
where the coefficient functions
$\rho (x,t),\, K(x,t),\, d(x,t),\, \gamma (x,t),$
and
$\hat {d}(x,t)$
are positive L-periodic in x and T-periodic in t, and of class
$C^{\kappa ,\frac {\kappa }{2}}([0,L]\times [0,T])$
for appropriate
$L>0$
,
$T>0,$
and
$\kappa \in (0,1)$
. We will study the spreading properties and time–space periodic traveling wave solutions for system (1.2). Since the linearization of
$H(u_M,u_F)$
at
$(0,0)$
exhibits a singularity, we cannot employ the linear operators approach to estimate the spreading speed. To overcome this difficulty, we need to study how to use an associated homogeneous system to determine the spreading speed.
In addition to the evolution dynamics of systems (1.1) and (1.2), we further quantitatively analyze the effects of the model parameters on basic reproduction ratios and spreading speeds. Notably, when the diffusion coefficient is sufficiently large, the spreading speed can either tend toward infinity or approach zero, which depends on the limiting behavior of the basic reproduction ratio in the case of a sufficiently large diffusion coefficient. To the best of our knowledge, this is the first analytic work to relate the limiting behavior of the spreading speed to that of the basic reproduction ratio.
The rest of this article is organized as follows. In Section 2, we prove the continuity of the cone spectral radius with respect to perturbations of the cone norm, without assuming compactness. In Section 3, we overcome the non-compactness of associated solution maps to establish the threshold dynamics of system (1.1) in a bounded habitat in terms of the basic reproduction ratio, which is defined by a time-periodic homogeneous evolution system without compactness. In Section 4, we study the propagation properties and traveling waves of system (1.2) and quantitatively investigate the spreading speed in the autonomous case.
2 Preliminaries
Let E be a Banach space with the norm
$\Vert \cdot \Vert $
. A closed convex set D is called a cone if
$\lambda D \subseteq D$
for all
$\lambda \geq 0$
and
$D \cap (-D) =\{ 0\}$
. A cone D is called solid if it contains an interior point (i.e.,
$\mathrm {Int} (D) \neq \emptyset $
), and normal if there exists
$c> 0$
such that
We write
$y\geq _{D}x$
if
$y - x \in D$
;
$y>_{D}x$
if
$y - x \in D$
and
$y - x\neq 0$
; and
$y \gg _{D} x$
if
$y - x \in \mathrm {Int} (D)$
.
Next, we recall some basic definitions (see, e.g., [Reference Chang4, Reference Nussbaum17, Reference Thieme19]). An operator A is called positive on D if
$A(D)\subseteq D$
; homogeneous (of degree one) if
$A(\alpha x) = \alpha A x$
for all
$\alpha \geq 0$
,
$x\in D$
; D-order-preserving if
$Ax \geq _{D} A y$
whenever
$x \geq _{D} y \geq _{D} 0$
; and D-strongly-order-preserving if
$Ax \gg _{D} A y$
whenever
$x>_{D} y \geq _{D} 0$
. Let
$A: D \rightarrow D$
be a homogeneous map, define
If the supremum exists, then A is said to be bounded and
$\Vert A \Vert _{D}$
is called the cone norm of A. Furthermore,
is called the cone spectral radius on D.
In the following, we extend the elementary results in [Reference Wang, Zhang and Zhao20, Lemma 2.4] to the case of non-compact homogeneous maps.
Lemma 2.1 Let E be a Banach space with a solid and normal cone D. Assume that C is a cone with
$C \subseteq D$
. Let
$L:C \rightarrow C$
be a bounded, continuous, homogeneous, and D-order-preserving map such that
$r_{C}(L)>0$
and
$Lu =r_{C}(L) u$
for some
$u \in \mathrm {Int}(D) \cap C$
, and let
$\{L_n\}_{n \geq 1}$
be a sequence of bounded, continuous, homogeneous, and D-order-preserving maps from C to C. If
$\lim \limits _{n \rightarrow \infty }\Vert L -L_{n} \Vert _{C}=0$
, then
$\lim _{n \rightarrow \infty } r_{C} (L_n) = r_{C} (L)$
.
Proof Define
Since D is normal and
$u \in \mathrm {Int}(D)$
,
$\Vert \cdot \Vert _u$
is an equivalent norm on E (see, e.g., [Reference Amann2, Theorem 2.4]). Let
$u_n:= L_n u$
for all
$n \geq 1$
. It is easy to see that
Then there exists a sequence
$\epsilon _n$
with
$\epsilon _n \rightarrow 0^+$
as
$n \rightarrow \infty $
and
$\Vert u_n -r_{C} (L) u \Vert _{u} \leq \epsilon _n r_{C} (L)$
for all
$n \geq 1$
. Therefore, for each
$n \geq 1$
,
An easy computation yields that
In view of [Reference Wang, Zhang and Zhao20, Lemma 2.1 and Corollary 2.7], it follows that
Letting
$n \rightarrow \infty $
, we then obtain
which implies that
$\lim _{n \rightarrow \infty } r_{C} (L_n) = r_{C} (L)$
.
Proposition 2.1 Let E be a Banach space with a solid cone D. Assume that C is a cone with
$C \subseteq D$
. Let
$L:C \rightarrow C$
be a bounded, continuous, homogeneous, and D-order-preserving map such that
$Lu =r_{C}(L) u$
for some
$u \in \mathrm {Int}(D) \cap C$
. If
$r_{C}(L)=0$
, then
$Lv=0$
for all
$v \in C$
.
Proof Let
$v\in C$
be given. Since
$u\in \mathrm {Int}(D)$
and
$v\in C\subseteq D$
, there exists
$\delta =\delta (v)>0$
such that
$u\ge _D \delta v$
. As L is D-order-preserving and homogeneous, we have
$Lu\ge _D L(\delta v)=\delta Lv$
. It follows from
$Lu=r_C(L)u=0$
that
$Lv\le _D0$
. On the other hand,
$Lv\in C\subseteq D$
. Hence,
$Lv\in D\cap (-D)=\{0\}$
, and therefore
$Lv=0$
.
As a consequence of [Reference Mallet-Paret and Nussbaum15, Corollary 4.6], we have the following observation.
Lemma 2.2 Let E be a Banach space with a total cone D, and
$\alpha $
be the Kuratowski measure of noncompactness in E. Assume that C is a cone with
$C \subseteq D$
. Let
$L:C \rightarrow C$
be a bounded, continuous, homogeneous, and D-order-preserving map. If
$r_{C}(L)\geq 1$
and there exists
$k\in (0,1)$
such that
$\alpha (LB) \leq k \alpha (B)$
for any bounded subset B of C, then
$r_{C}(L)$
is an eigenvalue of L with an eigenvector in
$C \setminus \{ 0 \}$
.
Proof For readers’ convenience, we provide the idea how to choose r and
$\mu $
. In the case of
$r_{C}(L)>1$
, we can choose
$r=r_{C}(L)>1$
and
$\mu =1$
; and in the case of
$r_{C}(L)=1$
, we can choose
$r=r_{C}(L)=1$
and
$\mu \in (k,1)$
.
3 Global stability in a bounded habitat
In this section, we study the global dynamics of system (1.1) in a bounded habitat. In order to establish a threshold-type result, we proceed with three sections.
3.1 A nonlinear eigenvalue problem
We consider the following homogeneous and partially degenerate reaction–diffusion system:
Let
$X=C(\overline {\Omega },\mathbb {R}^3)$
,
$P=C(\overline {\Omega },\mathbb {R}^3_+)$
, and
$\{U(t,s): t \geq s\}$
be the T-periodic evolution family on X associated with (3.1). We substitute
$u_N(x,t)=e^{-\lambda t}w_N(x,t)$
,
$N=A,M,F$
, into (3.1), to obtain the associated nonlinear eigenvalue problem:
Define
$p(x,t):=-d(x,t)-\gamma (x,t)$
,
$\forall (x,t) \in \overline {\Omega } \times \mathbb {R}$
,
$\tilde {p}(x)=T^{-1}\int _{0}^{T} p(x,t) \mathrm {d} t$
,
$\forall x\in \overline {\Omega }$
and
$\eta = -\max _{x \in \Omega } \tilde {p} (x)$
. By using the arguments similar to those in [Reference Hsu, Wang and Zhao9], we have the following observation.
Lemma 3.1
$\alpha (U(T,0)B) \leq e^{-\eta T}\alpha (B)$
for any bounded
$B \subseteq P$
.
Motivated by [Reference Liang, Zhang and Zhao11, Theorem 2.16 and Lemma 3.3], we are able to prove the following result for the nonlinear eigenvalue problem (3.2).
Lemma 3.2
$U(T,0)$
admits the principal eigenvalue with an eigenvector
$\boldsymbol {\phi }^*=(\phi _A^*,\phi _M^*,\phi _F^*)^T \in \mathrm {Int}(P)$
provided that one of the following two conditions holds true:
-
(i) $r_{P}(U(T,0)) \geq 1$
. -
(ii) There exists a nonempty $C^2$
subdomain
$\Omega _0 \subseteq \Omega $
such that
$\tilde {p}(x)=\max _{x \in \Omega }\tilde {p}(x)$
for all
$x \in \Omega _0$
,
Proof (i) In view of Lemma 2.2,
$U(T,0)$
admits the principal eigenvalue
$r_{P}(U(T,0))$
corresponding to an eigenvector
$\boldsymbol {\phi }^*=(\phi _A^*,\phi _M^*,\phi _F^*)^T \in P\setminus \{ 0\}$
. It suffices to show that
$\boldsymbol {\phi }^* \in \mathrm {Int}(P)$
. Write
$\lambda ^*=-\frac {\ln r_{P}(U(T,0))}{T} \leq 0$
and
$\boldsymbol {w}^*(x,t)=[e^{\lambda ^*t}U(t,0)\boldsymbol {\phi }^*](x)$
,
$\forall (x,t) \in \overline {\Omega } \times \mathbb {R}$
. It is easy to see that
$\boldsymbol {w}^*=(w_A^*,w_M^*,w_F^*)^T$
is T-periodic in
$t \in \mathbb {R}$
.
We first show that
$\phi _M^*(x)>0$
,
$\forall x \in \overline {\Omega }$
. Suppose that
$\phi _M^*(x_M)=0$
for some
$x_M \in \overline {\Omega }$
. We claim that
$w_M^*(x,t) =0$
,
$\forall (x,t) \in \overline {\Omega } \times \mathbb {R}$
. Otherwise,
$w_M^*(x_0,t_0)>0$
for some
$x_0 \in \overline {\Omega }$
and
$t_0 \in [0,T)$
, it then follows from
$[e^{\lambda ^*T}U(T,0)\boldsymbol {\phi }^*](x)=\boldsymbol {\phi }^*(x)$
and the strong maximum principle on the second equation of (3.2) that
$\phi _M^*(x)>0$
,
$\forall x \in \overline {\Omega }$
, which contradicts
$\phi _M^*(x_M)=0$
. Thus,
$w_M^*(x,t) =0$
,
$\forall (x,t) \in \overline {\Omega } \times \mathbb {R}$
. Thanks to the second equation of (3.2) again with
$\lambda $
replaced by
$\lambda ^*$
, we have
$w_A^*(x,t)=0$
,
$\forall (x,t) \in \overline {\Omega } \times \mathbb {R}$
. This further implies that
Integrating both sides of the above equation over
$\Omega \times [0,T]$
and using the T-periodicity in t together with the Neumann boundary condition, we obtain
$0=\int _0^T\int _\Omega (-\hat d(x,t)+\lambda ^*)w_F^*(x,t)\,\mathrm {d}x\mathrm {d}t$
. Since
$w_F^*\geq 0$
,
$\lambda ^*\leq 0$
, and
$\hat d(x,t)>0$
, it follows that
$w_F^*\equiv 0$
on
$\overline \Omega \times \mathbb R$
, which implies
$\boldsymbol \phi ^*\equiv 0$
. This contradicts
$\boldsymbol \phi ^*\in P\setminus \{0\}$
. Therefore,
$\phi _M^*(x)>0$
for all
$x\in \overline \Omega $
.
Similarly, we can prove that
$\phi _F^*(x)>0$
,
$\forall x \in \overline {\Omega }$
. It suffices to show that
$\phi _A^*(x)>0$
,
$\forall x\in \overline {\Omega }$
. Suppose that
$\phi _A^*(x_A)=0$
for some
$x_A \in \overline {\Omega }$
. If
$\phi _M^*(x_A) \phi _F^*(x_A)>0$
, then
$w_A^*(x_A,t)>0$
,
$\forall t>0$
, which contradicts
$\phi _A^*(x_A)=w_A^*(x_A,T)=0$
. If
$\phi _M^*(x_A) \phi _F^*(x_A)=0$
, then
$\phi _M^*(x_A)=0$
or
$\phi _F^*(x_A)=0$
, which is impossible. Therefore,
$\phi _A^*(x)>0$
,
$\forall x \in \overline {\Omega }$
, and hence,
$\boldsymbol {\phi }^*=(\phi _A^*,\phi _M^*,\phi _F^*)^T \in \mathrm {Int}(P)$
.
(ii) Define
For any
$\lambda < \eta $
, define
$p_{\lambda }(x,t)= p(x,t)+\lambda $
,
$\forall (x,t) \in \overline {\Omega } \times \mathbb {R}$
,
$\tilde {p}_{\lambda }=\tilde {p} +\lambda $
and let
$R_{\lambda }[\boldsymbol {w}]$
be the unique nonnegative T-periodic solution of
where
$\boldsymbol {w}=(w_M,w_F)^T \in C_T(\overline {\Omega } \times \mathbb {R},\mathbb {R}_+^2)$
. It is easy to verify that for any
$(x,t) \in \overline {\Omega } \times \mathbb {R}$
and
$\lambda < \eta $
, there holds
where
Let
$G=C_{0}^{1}(\overline \Omega _0)$
and
$G_+=\{w \in G: w(x) \geq 0,~ \forall x \in \overline {\Omega }_0\}$
. Then
$(G, G_+)$
is an ordered Banach space with
We denote by
$C_T(\mathbb {R},G)$
the Banach space of continuous T-periodic functions from
$\mathbb {R}$
to G, which is equipped with the maximum norm and the positive cone
$C_T(\mathbb {R},G_+)$
. It is easy to see that
$\mathrm {Int}(C_T(\mathbb {R},G_+))$
is nonempty.
Define two linear operators
$\mathcal {L}_M$
and
$\mathcal {L}_F$
by
with their domains
$\mathcal {D}(\mathcal {L}_M)= \mathcal {D}(\mathcal {L}_F)\subset C_T(\overline {\Omega }_0 \times \mathbb {R},\mathbb {R})$
being
For convenience, let
$q_M=\theta \gamma $
and
$q_F=(1-\theta )\gamma $
. Next, we find a suitable
$\overline {\lambda }< \eta $
such that
where
$\boldsymbol {\tilde {w}}=(\tilde {w}_M,\tilde {w}_F)^T \in C_T(\overline {\Omega }_0 \times \mathbb {R},\mathbb {R}^2_+)$
.
For
$N=M,F$
, let
$\lambda _N$
be the principal eigenvalue of
with the positive eigenvector
$\tilde {w}_N \in \mathrm {Int}(C_T(\mathbb {R},G_+)),$
where
$\max _{(x,t) \in \overline {\Omega }_0 \times \mathbb {R}}\vert \tilde {w}_N(x,t) \vert =~1 $
. It is easy to verify that there exist
$a>0$
and
$b>0$
such that
$a \tilde {w}_M \leq \tilde {w}_F \leq b \tilde {w}_M$
. Hence, there exist
$c_M>0$
,
$c_F>0$
,
$\hat {c}_M>0,$
and
$\hat {c}_F>0$
such that
and
Choose
$\hat {\eta }<\eta $
such that
$\lambda _M> \hat {\eta }$
and
$\lambda _F> \hat {\eta }$
. For any
$\lambda \in [\hat {\eta },\eta )$
, there exists
$\hat {c}_{1}>0$
independent of
$\lambda $
such that
$e^{\int _{s}^{t} p_{\lambda }(x,\tau )\mathrm {d} \tau }> \hat {c}_{1}$
,
$\forall s,t \in \mathbb {R}$
with
$0 \leq s \leq t \leq T$
,
$x \in \overline {\Omega }_0$
. Moreover, there exist
$\hat {c}_2>0$
,
$\tilde {c}_M>0,$
and
$\tilde {c}_F>0$
such that
$\rho (x,t)> \hat {c}_2$
,
$\theta \gamma (x,t) \geq \tilde {c}_M$
,
$(1-\theta ) \gamma (x,t) \geq \tilde {c}_F$
,
$\forall (x,t) \in \overline {\Omega }_0 \times \mathbb {R}$
. An easy computation yields that
and
Therefore,
where
$b_N=\hat {c}_1^2 \hat {c}_2 \tilde {c}_N c_N \hat {c}_N$
for
$N=M,F$
. We set
$\tilde {\lambda }_N=\frac {b_N}{\hat {\eta }-\lambda _N} +\eta $
for
$N=M,F$
,
$\overline {\lambda }= \max (\tilde {\lambda }_M,\tilde {\lambda }_F,\hat {\eta })$
. It is easy to see that
$\overline {\lambda }<\eta $
. A straightforward computation gives rise to
By similar arguments, we also get
Therefore, (3.3) holds true.
Letting
$\tilde {w}_A= R_{\overline {\lambda }}\boldsymbol {\tilde {w}}$
, we then have
For
$N=A,M,F$
, define
We then obtain that
$e^{\overline {\lambda } T} U(T,0) \boldsymbol {\tilde {\phi }} \geq \boldsymbol {\tilde {\phi }}$
. Choose
$\sigma \in (0,\eta -\overline {\lambda })$
small enough. It follows from [Reference Wang, Zhang and Zhao20, Lemma 2.1] that
$r_{P}(e^{\overline {\lambda }T} U(T,0)) \geq 1$
, and hence,
$r_{P}(e^{(\eta -\sigma ) T}U(T,0))>r_{P}(e^{\overline {\lambda }T} U(T,0)) \geq 1$
. With Lemma 3.1, we have
$\alpha (e^{(\eta -\sigma )T}U(T,0)B) \leq e^{-\sigma T}\alpha (B)$
for any bounded set
$B \subseteq P$
. It then follows from Lemma 2.2 that
$e^{(\eta -\sigma ) T}U(T,0)$
, and hence,
$U(T,0)$
admits the principal eigenvalue with an eigenvector
$\phi ^*\in P \setminus \{0\}$
.
For
$N=M,F$
, let
$\hat {\lambda }_N$
be the principal eigenvalue of
with the positive eigenvector
$\hat {w}_N \in \mathrm {Int}(C_T(\overline {\Omega }\times \mathbb {R},\mathbb {R}_+))$
satisfying
$\max _{(x,t) \in \overline {\Omega } \times \mathbb {R}}\vert \hat {w}_N(x,t) \vert =1 $
. Note that
$\lambda ^*=- \frac {\ln r_{P}( U(T,0))}{T}\leq \min (\eta ,\hat {\lambda }_M,\hat {\lambda }_F)$
. Since
$r_{P}(e^{(\eta -\sigma ) T}U(T,0))>1$
, we have
$\lambda ^*<\eta -\sigma $
. In view of
$\lambda _0:=\hat {\lambda }_M=\hat {\lambda }_F$
, choosing
$\hat {w}_A=0$
, we then have
Define
By the comparison arguments, it is not hard to verify that
$e^{\lambda _0 T} U(T,0) \boldsymbol {\hat {\psi }} \gg \boldsymbol {\hat {\psi }}$
. Using [Reference Wang, Zhang and Zhao20, Lemma 2.1] again, we have
$r_{P}(e^{\lambda _0 T} U(T,0))>1$
, which implies that
$\lambda ^*<\lambda _0$
. Therefore,
$\lambda ^*<\min (\eta ,\hat {\lambda }_M,\hat {\lambda }_F)$
. Now one can easily obtain
$\boldsymbol {\phi }^* \in \mathrm {Int}(P)$
by using the arguments similar to those in case (i).
3.2 Basic reproduction ratio
In this section, we introduce and characterize the basic reproduction ratio
$\mathcal {R}_0$
for system (1.1), and then quantitatively analyze the effects of parameters on
$\mathcal {R}_0$
.
Following the notations and procedure in [Reference Wang, Zhang and Zhao20, Section 3], we choose
where
$C_T(\mathbb {R},X)$
is the Banach space of all T-periodic and continuous functions from
$\mathbb {R}$
to X. Let
$\{\Phi (t,s),\ t\geq s\}$
be the evolution family on X associated with the following linear system:
It is easy to verify that
$\omega (\Phi )<0$
. For each
$t\in \mathbb {R}$
, we define
$F(t):P\rightarrow P$
by
where
$(w_A,w_M,w_F)^T\in P$
. Thus, the operator
$\mathcal {L}$
on
$\mathbb {P}$
is defined by
and the basic reproduction ratio of mosquitoes is given by
For any
$\mu>0$
, let
$\{U(t,s;\mu )\}$
be the evolution family on X of the following system:
Proposition 3.1
$\mathcal {R}_0 -1$
and
$r_{P}(U(T,0))-1$
have the same sign. Moreover,
$\mathcal {R}_0^{-1}$
is the unique solution of
$r_{P}(U(T,0;\mu ))=1$
.
Proof It is easy to see that (H1)–(H4) in [Reference Wang, Zhang and Zhao20, Section 3] hold true. Thanks to Lemma 3.2, if there exists a domain
$\Omega _0 \subseteq \Omega $
such that
$\tilde {p}(x)=-\eta $
for all
$x \in \Omega _0$
, then (H5) in [Reference Wang, Zhang and Zhao20, Section 3] is true. Thus, the desired conclusion follows from Lemma 2.1 and [Reference Wang, Zhang and Zhao20, Theorem 3.4].
To prove the conclusion in the general case, we first take a series of functions
$d_{\epsilon }$
and domains
$\Omega _{\epsilon } \subseteq \Omega $
such that
$d_{\epsilon } \leq d \leq d_{\epsilon }+\epsilon $
and
$\tilde {p}^{\epsilon }(x)=\max _{x \in \Omega } \tilde {p}^{\epsilon } (x)$
,
$\forall x \in \overline {\Omega }_{\epsilon }$
, where
$\tilde {p}^{\epsilon }(x)=T^{-1}\int _{0}^{T} [-d_{\epsilon }(x,t)-\gamma (x,t)] \mathrm {d} t$
,
$\forall x \in \overline {\Omega }$
.
We can define
$\mathcal {R}_0^{\epsilon }$
and
$\{U^{\epsilon }(t,s): t \geq s \}$
(resp.
$\mathcal {R}_{0,\epsilon }$
and
$\{U_{\epsilon }(t,s): t \geq s \}$
) in the same way as
$\mathcal {R}_0$
and
$\{U(t,s): t \geq s \}$
with d replaced by
$d_{\epsilon }$
(resp.
$d_{\epsilon }+\epsilon $
). It is easy to see that
$\mathcal {R}_{0,\epsilon } \leq \mathcal {R}_{0} \leq \mathcal {R}_0^{\epsilon }$
and
$r_{P}(U_{\epsilon }(T,0)) \leq r_{P}(U(T,0)) \leq r_{P}(U^{\epsilon }(T,0))\leq r_{P}(U_{\epsilon }(T,0)) e^{\epsilon T}$
. Moreover,
$\mathcal {R}_{0,\epsilon }-1$
and
$r_{P}(U_{\epsilon }(T,0)) -1$
have the same sign, and
$\mathcal {R}_{0}^{\epsilon }-1$
and
$r_{P}(U^{\epsilon }(T,0)) -1$
share the same sign.
If
$r_{P}(U(T,0))=1$
, then
$\mathcal {R}_0=1$
by [Reference Wang, Zhang and Zhao20, Lemma 3.2(i)]. If
$r_{P}(U(T,0))>1$
, one can choose
$\epsilon>0$
small enough such that
$r_{P}(U(T,0)) e^{-\epsilon T}>1$
. We then have
$r_{P}(U_{\epsilon }(T,0))\geq r_{P}(U(T,0)) e^{-\epsilon T}>1$
, and hence,
$\mathcal {R}_0\geq \mathcal {R}_{0,\epsilon }>1$
. By the similar arguments, we can also prove that
$\mathcal {R}_0<1$
whenever
$r_{P}(U(T,0))<1$
.
Repeating the above arguments, we see that
$\mu \mathcal {R}_0-1$
and
$r_{P}(U(T,0;\mu ))-1$
have the same sign for all
$\mu>0$
. By the comparison arguments, it is not hard to prove that
$\mathcal {R}_0\geq \theta (1-\theta )\underline {\rho }\underline {\gamma } [{\overline {d}}(\overline {d}+\overline {\gamma })]^{-1}>0$
, where
$\underline {\rho }=\min _{(x,t)\in \overline {\Omega } \times \mathbb {R}} \rho (x,t)$
,
$\underline {\gamma }=\min _{(x,t)\in \overline {\Omega } \times \mathbb {R}} \gamma (x,t)$
,
$\overline {\gamma }=\max _{(x,t)\in \overline {\Omega } \times \mathbb {R}} \gamma (x,t)$
, and
$\overline {d}=\max _{(x,t)\in \overline {\Omega } \times \mathbb {R}} \max (d(x,t),\hat {d}(x,t))$
. Therefore,
$\mathcal {R}_0^{-1}$
is the unique solution of
$r_{P}(U(T,0;\mu ))=1$
.
Theorem 3.1 The following statements are valid:
-
(i) $\frac {\mathcal {R}_0}{\theta (1-\theta )}$
is independent of
$\theta $
. In particular,
$\mathcal {R}_0$
is increasing with respect to
$\theta \in (0,\frac {1}{2}]$
and decreasing with respect to
$\theta \in [\frac {1}{2},1)$
, and goes to
$0$
as
$\theta \rightarrow 0^{+}$
or
$\theta \rightarrow 1^{-}$
. -
(ii) If, in addition, all parameters are independent of t, then $\mathcal {R}_0$
is decreasing with respect to D. Moreover,
$\mathcal {R}_0 \rightarrow \theta (1-\theta ) \max _{x \in \overline {\Omega }} \frac {\rho (x)\gamma (x)}{(d(x)+\gamma (x))\hat {d}(x)}$
as
$D \rightarrow 0^{+}$
and
$\mathcal {R}_0 \rightarrow \theta (1-\theta )\frac {\int _{\Omega } \frac {\rho (x)\gamma (x)}{d(x)+\gamma (x)} \mathrm {d} x}{\int _{\Omega } \hat {d} (x) \mathrm {d} x}$
as
$D \rightarrow +\infty $
.
Proof (i) It suffices to prove that
$\mathcal {R}_0^{-1}$
is an eigenvalue of the weighted eigenvalue problem
with a strongly positive eigenvector. With Proposition 3.1, we have
$r_{P}(U(T,0;\mathcal {R}_0^{-1}))=1$
. Thanks to Lemma 3.2, there exists a strongly positive eigenvector
$(w_A^*,w_M^*,w_F^*)^T$
such that
Multiplying the second and third equations by
$1-\theta $
and
$\theta $
, respectively, and then making difference, we obtain
where
$z=(1-\theta ) w_M^* - \theta w_F^*$
. It is easy to verify that
$z \equiv 0$
due to the Krein–Rutman theorem. Let
$w_S^*=\theta ^{-1}w_M^*=(1-\theta )^{-1}w_F^*$
. Then (3.4) holds with
$\mu =\mathcal {R}_0^{-1}$
,
$w_A=w_A^*$
, and
$w_S=w_S^*$
.
(ii) If all parameters are independent of t, then so are
$w_A^*$
and
$w_S^*$
. By the first equation of (3.4), it then follows that
which yields that
$\mathcal {R}_0^{-1}$
is an eigenvalue of the weighted eigenvalue problem
Now the desired conclusion follows from the same arguments as in [Reference Allen, Bolker, Lou and Nevai1, Lemma 2.3].
3.3 Threshold dynamics
In this section, we establish the threshold dynamics for system (1.1) in terms of the basic reproduction ratio
$\mathcal {R}_0$
. For this purpose, we choose the feasible domain of system (1.1) as
To guarantee the positive invariance of
$\Sigma (t)$
for system (1.1), we further assume that
-
(A) $\frac {\partial K(x,t)}{\partial t}\geq -(d(x,t)+\gamma (x,t)) K(x,t)$
for all
$x\in \overline {\Omega },\, t>0$
.
Note that assumption (A) holds true automatically in the autonomous case. For convenience, we also let
Then we have the following threshold-type result on the global dynamics of system (1.1).
Theorem 3.2 Let
$\boldsymbol {u}(x,t;\boldsymbol {\phi })$
be the unique solution of system (1.1) with
$\boldsymbol {u}(\cdot ,0;\boldsymbol {\phi })=\boldsymbol {\phi }\in \Sigma (0)$
. Then the following statements are valid:
-
(i) If $\mathcal {R}_0\leq 1$
, then
$\lim \limits _{t\rightarrow +\infty }\Vert \boldsymbol {u}(\cdot ,t;\boldsymbol {\phi }) \Vert _{C(\overline {\Omega },\mathbb {R}^3)}=0$
for all
$\boldsymbol {\phi } \in \Sigma (0)$
. -
(ii) If $\mathcal {R}_0>1$
, then system (1.1) admits a unique positive T-periodic solution
$\boldsymbol {u}^*(x,t)$
, and
$\lim \limits _{t\rightarrow +\infty }\Vert \boldsymbol {u}(\cdot ,t;\boldsymbol {\phi })- \boldsymbol {u}^*(\cdot ,t) \Vert _{C(\overline {\Omega },\mathbb {R}^3)}=0$
for all
$\boldsymbol {\phi } \in \tilde {\Sigma }$
.
Proof By [Reference Martin and Smith16, Corollary 4] and assumption (A), it follows that for every initial value function
$\boldsymbol {\phi }:=(\phi _A,\phi _M,\phi _F)^T\in \Sigma (0)$
, system (1.1) admits a unique non-continuable solution
$\boldsymbol {u}(\cdot ,t;\boldsymbol {\phi }):=(u_A,u_M,u_F)^T(\cdot ,t;\boldsymbol {\phi })$
on its maximal interval of existence
$[0,t_{\boldsymbol {\phi }})$
with
$\boldsymbol {u}(\cdot ,0;\boldsymbol {\phi })=\boldsymbol {\phi }$
, where
$t_{\boldsymbol {\phi }}\leq \infty $
, and
$\boldsymbol {u}(\cdot ,t;\boldsymbol {\phi })\in \Sigma (t)$
for all
$t\in [0,t_{\boldsymbol {\phi }})$
. Thus,
$0\leq u_A(x,t;\boldsymbol {\phi })\leq K(x,t),\ \forall \ x \in \bar {\Omega },\ t\geq 0$
. Then we can use standard comparison arguments to further show that solutions of system (1.1) are ultimately bounded and uniformly bounded. Hence, we can define the solution maps
$S(t):\Sigma (0)\rightarrow \Sigma (t)$
associated with system (1.1) by
where
$\boldsymbol {u}(x,t;\boldsymbol {\phi })$
is the unique solution of system (1.1) with
$\boldsymbol {u}(x,0;\boldsymbol {\phi })=\boldsymbol {\phi }(x),\ \forall \ x \in \bar {\Omega }$
. Let
$S:=S(T):\Sigma (0)\rightarrow \Sigma (0)$
be the Poincaré map associated with system (1.1) on
$\Sigma (0)$
. By the theory of periodic semiflows (see [Reference Zhao22]), it suffices to prove a threshold-type result on the global dynamics for S on
$\Sigma (0)$
in terms of
$\mathcal {R}_0$
. By using the arguments similar to those in [Reference Hsu, Wang and Zhao9], we see that S is
$\alpha $
-contracting in the sense that
$\lim _{n\to \infty }\alpha (S^n(B))=0\ \text { for any bounded set}\ B \subset \Sigma (0)$
. It follows from [Reference Magal and Zhao14, Theorem 2.6] that S admits a strong global attractor that attracts each bounded set in
$\Sigma (0)$
. By the essentially same arguments as those for [Reference Bai and Zhao3, Theorem 3.1], we can easily show that the desired statements are valid in the case where
$\mathcal {R}_0<1$
or
$\mathcal {R}_0>1$
.
It remains to consider the critical case where
$\mathcal {R}_0=1$
. In view of Proposition 3.1, we have
$r_{P}(U(T,0))=1$
. It follows from Lemma 3.2 that 0 is the principal eigenvalue of system (3.2), which corresponds to an eigenvector
$\boldsymbol {\psi }=(\psi _A,\psi _M,\psi _F)^T \in \mathrm {Int}(P)$
. Motivated by [Reference Cui, Lam and Lou5, Reference Li and Zhao10], we define
Then we have the following observation.
Claim.
$c(nT;\boldsymbol {\phi })$
is non-increasing in n, and
$c(nT;\boldsymbol {\phi })$
is not a constant sequence whenever
$\boldsymbol {\phi } \in \tilde {\Sigma }$
.
Indeed, since
$\phi _A(\cdot )\leq K(\cdot ,0)$
, we have
$u_A(\cdot ,t,\boldsymbol {\phi })\leq K(\cdot ,t)$
for all
$t\geq 0$
. Fix
$n_0>0$
, we let
Then
$(\bar {u}_A(x,t;\boldsymbol {\phi }),\bar {u}_M(x,t;\boldsymbol {\phi }),\bar {u}_F(x,t;\boldsymbol {\phi }))$
satisfies (3.2) with
$\lambda =0$
. Since
$(u_A,u_M,u_F)(x,t;\boldsymbol {\phi })$
satisfies system (1.1), it follows that
and
By the comparison principle, we have
This implies that
Moreover,
$c(nT;\boldsymbol {\phi })$
is non-increasing in n. In addition, if
$\boldsymbol {\phi }:=(\phi _A,\phi _M,\phi _F)\in \tilde \Sigma $
, then one can show that
and hence,
which implies that the first inequality in (3.6) must be strict. Thus, the strong comparison principle ensures
It then follows that
$c(n T;\boldsymbol {\phi })< c(n_0 T;\boldsymbol {\phi })$
whenever
$n> n_0$
, and hence,
$c(nT;\boldsymbol {\phi })$
is not a constant sequence whenever
$\boldsymbol {\phi } \in \tilde \Sigma $
.
Now we show that
$\lim _{n\rightarrow \infty }S^n\boldsymbol {\phi }=0$
for all
$\boldsymbol {\phi }\in \tilde \Sigma $
. Clearly,
$\lim _{n\rightarrow \infty }c(nT;\boldsymbol {\phi })=c_0\geq 0$
. Let
$\omega (\boldsymbol {\phi })$
be the omega limit set of the forward orbit through
$\boldsymbol {\phi }$
for S. For any given
$\boldsymbol {\varphi } \in \omega (\boldsymbol {\phi })$
, there is a sequence
$n_k\to \infty $
such that
$\lim _{k\rightarrow \infty }S^{n_k}\boldsymbol {\phi }=\boldsymbol {\varphi }$
. It then follows that
By the definition of
$c(t;\boldsymbol {\varphi })$
, we have
It then follows from the above claim that
$\boldsymbol {\varphi }=(0,0,0)$
, and hence,
$\omega (\boldsymbol {\phi })=\{(0,0,0)\}$
. The desired conclusion can be derived easily for all
$\boldsymbol {\phi }\in \Sigma (0) \setminus \tilde \Sigma $
.
Remark 3.2 In view of [Reference Wu and Zhao21, Lemma 2.1], it turns out that the zero solution is globally asymptotically stable for system (1.1) in
$\Sigma (0)$
if
$\mathcal {R}_0\leq 1$
, and the positive periodic solution
$u^*(x,t)$
is globally asymptotically stable for system (1.1) in
$\tilde {\Sigma }$
if
$\mathcal {R}_0> 1$
.
4 Propagation in a periodic habitat
In this section, we investigate the spreading properties and time–space periodic traveling waves for system (1.2) in a spatially periodic habitat. Throughout this section, we always assume that the function
$K(x,t)$
satisfies condition (A) as stated in Section 3.3.
4.1 The periodic initial value problem
We first consider the following reaction–diffusion system with periodic initial value:
where
$\mathcal {X}=PC(\mathbb {R},\mathbb {R}^3)$
represents the set of all continuous and L-periodic functions from
$\mathbb {R}$
to
$\mathbb {R}^3$
with the maximum norm
$\|\cdot \|_{\mathcal {X}}$
, and
$\mathcal {X}_+=\{\psi \in \mathcal {X}: \ \psi (x)\ge 0,\forall x\in \mathbb {R}\}$
is a positive cone of
$\mathcal {X}$
. Clearly,
$(\mathcal {X},\mathcal {X}_+)$
is a strongly ordered Banach space.
Let
$\{\mathcal {U}(t,s): t \geq s\}$
be the evolution family associated with homogeneous system on
$\mathcal {X}$
:
that is,
$\mathcal {U} (t,0)\boldsymbol {\phi }=\boldsymbol {u}(\cdot ,t;\boldsymbol {\phi })$
, where
$\boldsymbol {u}(x,t;\boldsymbol {\phi }):=(u_A(x,t;\boldsymbol {\phi }),u_M(x,t;\boldsymbol {\phi }),u_F(x,t;\boldsymbol {\phi }))$
is the unique solution of system (4.2) with
$\boldsymbol {u}(\cdot ,0;\boldsymbol {\phi })=\boldsymbol {\phi }\in \mathcal {X}$
.
According to [Reference Wang, Zhang and Zhao20, Section 3], we choose
where
$C_T(\mathbb {R},\mathcal {X})$
is the Banach space of all T-periodic and continuous functions from
$\mathbb {R}$
to
$\mathcal {X}$
. Then we can use the arguments similar to those in Section 3.2 to define the basic reproduction ratio
$\mathcal {R}_0$
and obtain the following result.
Lemma 4.1
$\mathcal {R}_0 -1$
and
$r_{\mathcal {X}_+}(\mathcal {U}(T,0))-1$
have the same sign.
Note that the feasible domain of system (1.2) takes the form
For convenience, we also let
By the arguments similar to those for Theorem 3.2, we have the following threshold-type result on the global dynamics of system (4.1).
Lemma 4.2 Let
$\boldsymbol {u}(x,t;\boldsymbol {\varphi })$
be the unique solution of system (4.1) with
$\boldsymbol {u}(\cdot ,0;\boldsymbol {\varphi })=\boldsymbol {\varphi }\in \Upsilon (0)$
. Then the following statements are valid:
-
(i) If $\mathcal {R}_0\leq 1$
, then
$\lim \limits _{t\rightarrow +\infty }\Vert \boldsymbol {u}(\cdot ,t;\boldsymbol {\varphi }) \Vert _{\mathcal {X}}=0$
for all
$\boldsymbol {\varphi }\in \Upsilon (0)$
. -
(ii) If $\mathcal {R}_0>1$
, then system (4.1) admits a unique positive time–space periodic solution
$\boldsymbol {u}^*(x,t)$
with
$\boldsymbol {u}_A^*(x,t)\leq K(x,t)$
for all
$(x,t)\in \mathbb {R}^2$
, and
$\lim \limits _{t\rightarrow +\infty }\Vert \boldsymbol {u}(\cdot ,t;\boldsymbol {\varphi })- \boldsymbol {u}^*(\cdot ,t) \Vert _{\mathcal {X}}=0$
for all
$\boldsymbol {\varphi }\in \tilde {\Upsilon }$
.
4.2 Spreading speeds and traveling waves
In this section, we first establish the well-posedness of system (1.2), and then investigate the spreading speeds and traveling waves of system (1.2) in the case where
$\mathcal {R}_0>1$
.
Let
$\mathcal {C}$
be the space of all bounded and continuous functions from
$\mathbb {R}$
to
$\mathbb {R}^3$
. It is easy to see that
$\mathcal {C}^{+}:=\{\boldsymbol {u}\in \mathcal {C}:\boldsymbol {u}(x)\geq \boldsymbol {0},\ \forall x\in \mathbb {R} \} $
is a positive cone of
$\mathcal {C}$
. For
$\boldsymbol {u}:=(u_1,u_2,u_3)$
and
$\boldsymbol {v}:=(v_1,v_2,v_3)\in \mathcal {C}$
, we write
$\boldsymbol {u}\geq \boldsymbol {v}$
provided that
$u_{i}(x)\geq v_{i}(x)$
,
$\forall \ i=1,2,3$
,
$x\in \mathbb {R}$
, and
$\boldsymbol {u}> \boldsymbol {v}$
if
$\boldsymbol {u}\geq \boldsymbol {v}$
and
$\boldsymbol {u}\neq \boldsymbol {v}$
. We equip
$\mathcal {C}$
with the compact open topology, that is,
$\boldsymbol {u}^{m}\rightarrow \boldsymbol {u}$
in
$\mathcal {C}$
means that the sequence of
$\boldsymbol {u}^{m}(x)$
converges to
$\boldsymbol {u}(x)$
as
$m\rightarrow \infty $
uniformly for x in any compact set of
$\mathbb {R}$
. Define
where
$\mid \cdot \mid $
denotes the usual norm in
$\mathbb {R}^3$
. Then
$(\mathcal {C},\|\cdot \|)$
is a normed space. For
$\boldsymbol {r}\in \mathcal {C}^{+}$
, we define
$\mathcal {C}_{\boldsymbol {r}}:=\{\boldsymbol {u}\in \mathcal {C}:\boldsymbol {0}\leq \boldsymbol {u} \leq \boldsymbol {r}\}$
. Besides, we let
where
$\mid \cdot \mid $
denotes the usual norm in
$\mathbb {R}^3$
.
Assume that
$\mathcal {R}_0>1$
. In view of Lemma 4.2, there exist two periodic solutions
$\boldsymbol {0}:=(0, 0,0)$
and
$\boldsymbol {u}^*(x,t):=(u_1^*(x,t),u_2^*(x,t),u_3^*(x,t))$
for system (1.2). Then we have the following result.
Lemma 4.3 For any initial value
$\phi \in \mathcal {C}_{\boldsymbol {u}^*(\cdot ,0)}$
, system (1.2) admits a unique solution
$\boldsymbol {u}(x,t;\boldsymbol {\phi })$
with
$\boldsymbol {u}(\cdot ,0;\boldsymbol {\phi })=\boldsymbol {\phi }$
and
$\boldsymbol {u}(\cdot ,t;\boldsymbol {\phi })\in \mathcal {C}_{\boldsymbol {u}^*(\cdot ,t)}$
for
$t\in [0,\infty )$
. Moreover,
$\boldsymbol {u}(x,t;\boldsymbol {\phi })\ge \boldsymbol {u}(x,t;\boldsymbol {\psi })$
for all
$x\in \mathbb {R}$
and
$t\ge 0$
provided that
$\boldsymbol {\phi }\ge \boldsymbol {\psi }$
in
$\mathcal {C}_{\boldsymbol {u}^*(\cdot ,0)}$
.
Proof Following the arguments similar to those for[Reference Zhao23, Lemma 4.1], we first rewrite system (1.2) as an abstract integral equation in the Banach space
$(\mathcal {C}_{\boldsymbol {u}^*(\cdot ,0)},\|\cdot \|_{\infty })$
, and then apply [Reference Martin and Smith16, Corollary 5] with
$\tau =0$
to obtain the desired conclusions.
In Lemma 4.3, we use the Banach space
$(\mathcal {C}_{\boldsymbol {u}^*(\cdot ,0)},\|\cdot \|_{\infty })$
to establish the existence and uniqueness of solutions and the comparison principle for system (1.2). Further, one can use the same arguments as in [Reference Zhao23, Theorem 4.4 and Remark 1] to show that the solution maps of system (1.2) induce a monotone semiflow on
$(\mathcal {C}_{\boldsymbol {u}^*(\cdot ,0)},\|\cdot \|)$
with respect to the compact open topology. Thus, we can define a family of operators
$\{Q_t\}_{t\ge 0}$
from
$ \mathcal {C}_{\boldsymbol {u}^*(\cdot ,0)}$
to
$ \mathcal {C}_{\boldsymbol {u}^*(\cdot ,t)}$
by
where
$\boldsymbol {u}(\cdot ,t;\boldsymbol {\phi })$
is the solution of system (1.2) with
$\boldsymbol {u}(\cdot ,0;\boldsymbol {\phi })=\boldsymbol {\phi }\in \mathcal {C}_{\boldsymbol {u}^*(\cdot ,0)}$
.
By the arguments similar to those in [Reference Fang, Lai and Wang6, Section 4], we have the following two results on the continuity and
$\alpha $
-contraction property for
$Q_t$
.
Lemma 4.4
$Q_t[\boldsymbol {\phi }]$
is continuous in
$(t,\boldsymbol {\phi })$
in the sense that if
$\boldsymbol {\phi }_n \rightarrow \boldsymbol {\phi }$
in
$\mathcal {C}_{\boldsymbol {u}^*(\cdot ,0)}$
and
$t_n \rightarrow t$
as
$n \rightarrow \infty $
, then
$Q_{t_n}[\boldsymbol {\phi }_n] \rightarrow Q_{t}[\boldsymbol {\phi }]$
in
$\mathcal {C}$
.
Lemma 4.5 For any
$I=[a,b] \subset \mathbb {R}$
and
$t>0$
, there exists
$\zeta (t) \in (0,1)$
such that
$\alpha ((Q_t[B])_I) \leq \zeta (t)\alpha (B_I)$
for any bounded
$B \subseteq \mathcal {C}_{\boldsymbol {u}^*(\cdot ,0)}$
.
As a consequence of [Reference Fang, Yu and Zhao7, Theorem 2.1], we have the following result on time–space periodic traveling waves of system (1.2).
Theorem 4.1 Assume that
$\mathcal {R}_0>1$
. Then there exist two numbers
$c_+^*$
and
$c_-^*$
such that the following statements are valid:
-
(i) For any $c \geq c_+^*$
, system (1.2) admits rightward traveling waves
$\boldsymbol {U}(x,t,x-ct)$
connecting
$\boldsymbol {u}^*(x,t)$
to
$\boldsymbol {0}$
, that is,
$\boldsymbol {U}(x,t,-\infty )=\boldsymbol {u}^*(x,t)$
and
$\boldsymbol {U}(x,t,+\infty )=\boldsymbol {0}$
, with the wave profile
$\boldsymbol {U}(x,t,\xi )=\boldsymbol {U}(x+L,t+T,\xi )$
being continuous and non-increasing in
$\xi $
. For any
$c <c_+^*$
, system (1.2) admits no such traveling waves connecting
$\boldsymbol {u}^*(x,t)$
to
$\boldsymbol {0}$
. -
(ii) For any $c \geq c_-^*$
, system (1.2) admits leftward traveling waves
$\boldsymbol {V}(x,t,x+ct)$
connecting
$\boldsymbol {0}$
to
$\boldsymbol {u}^*(x,t)$
, that is,
$\boldsymbol {V}(x,t,-\infty )=\boldsymbol {0}$
and
$\boldsymbol {V}(x,t,+\infty )=\boldsymbol {u}^*(x,t)$
, with the wave profile
$\boldsymbol {V}(x,t,\xi )=\boldsymbol {V}(x+L,t+T,\xi )$
being continuous and non-decreasing in
$\xi $
. For any
$c <c_-^*$
, system (1.2) admits no such traveling waves connecting
$\boldsymbol {0}$
to
$\boldsymbol {u}^*(x,t)$
.
The following result shows that the minimum wave speed coincides with the spreading speed, which follows from [Reference Fang, Yu and Zhao7, Theorem 2.3].
Theorem 4.2 Assume that
$\mathcal {R}_0>1$
and let
$c_{\pm }^*$
be given in Theorem 4.1. Let
$\boldsymbol {u}(x,t;\boldsymbol {\phi })$
be a solution of (1.2) with
$\boldsymbol {u}(\cdot ,0;\boldsymbol {\phi })=\boldsymbol {\phi }=(\phi _A,\phi _M,\phi _F)^T\in \mathcal {C}_{\boldsymbol {u}^*(\cdot ,0)}$
. Then the following statements are valid:
-
(i) If $0\leq \boldsymbol {\phi }(\cdot )\leq \boldsymbol {\varphi } (\cdot )\ll \boldsymbol {u}^*(\cdot ,0)$
, for some
$\boldsymbol {\varphi } (\cdot )\in \mathcal {X}$
, and there exist
$x_1$
,
$\tilde {x}_1$
, and
$\delta _1>0$
with
$x_1<\tilde {x}_1$
such that
$\phi _N(x)> \delta _1$
for
$x < x_1$
and
$\phi _N(x)=0$
for
$x>\tilde {x}_1$
,
$N=A,M,F$
, then $$ \begin{align*}\lim_{t\rightarrow\infty}\sup_{\ x\geq ct}\Vert\boldsymbol{u}(x,t;\boldsymbol{\phi})\Vert=0,\ \forall c>c_{+}^*,\!\!\quad\lim_{t\rightarrow\infty }\sup_{x\leq ct}\Vert\boldsymbol{u}(x,t;\boldsymbol{\phi})-\boldsymbol{u}^*(x,t)\Vert=0, \ \forall c<c_{+}^*.\end{align*} $$
-
(ii) If $0\leq \boldsymbol {\phi }(\cdot )\leq \boldsymbol {\varphi } (\cdot )\ll \boldsymbol {u}^*(\cdot ,0)$
, for some
$\boldsymbol {\varphi } (\cdot )\in \mathcal {X}$
, and there exist
$x_2$
,
$\tilde {x}_2$
, and
$\delta _2>0$
with
$x_2>\tilde {x}_2$
such that
$\phi _N(x)> \delta _2$
for
$x> x_2$
and
$\phi _N(x)=0$
for
$x<\tilde {x}_2$
,
$N=A,M,F$
, then $$ \begin{align*} &\lim_{t\rightarrow\infty}\sup_{\ x\leq - ct}\Vert\boldsymbol{u}(x,t;\boldsymbol{\phi})\Vert=0,\ \forall c>c_{-}^* \ ,\quad \\ & \lim_{t\rightarrow\infty}\sup_{\ x\geq -ct}\Vert\boldsymbol{u}(x,t;\boldsymbol{\phi})-\boldsymbol{u}^*(x,t)\Vert=0, \ \forall c<c_{-}^*.\end{align*} $$
Since (4.1) cannot be linearized at the zero solution, we are unable to use the linear operators approach in [Reference Liang and Zhao12] to estimate
$c_+^*$
and
$c_-^*$
. Next, we show that the spreading speeds can be determined by an associated homogeneous system.
For
$\mu \in \mathbb {R}$
, substituting
$(u_{A}(x,t),u_{M}(x,t),u_{F}(x,t))=e^{-\mu x}(v_A(x,t),v_M(x,t),v_F(x,t))$
into (4.2), we obtain
Let
$\{L_{\mu }(t,s)\}_{t\geq s}$
be the evolution family on
$\mathcal {C}$
generated by system (4.3), that is,
$L_{\mu }(t,0)\boldsymbol {\varphi }=\boldsymbol {v}(\cdot ,t;\boldsymbol {\varphi })$
, where
$\boldsymbol {v}(x,t;\boldsymbol {\varphi })$
is the unique solution of system (4.3) with
$\boldsymbol {v}(x,0;\boldsymbol {\varphi })=\boldsymbol {\varphi }$
. It is not hard to see that for any
$\mu \geq 0$
,
$L_{\mu }(t,0) \mathcal {X}_+ \subset \mathcal {X}_+$
, and
Substituting
$(v_{A}(x,t),v_{M}(x,t),v_{F}(x,t))=e^{-\lambda t}(w_A(x,t),w_M(x,t),w_F(x,t))$
into (4.3), we obtain the following periodic eigenvalue problem:
Using the arguments in Lemma 3.2(i), we have the following observation.
Lemma 4.6 If
$r_{\mathcal {X}_+}(L_{\mu }(T,0)) \geq 1$
, then eigenvalue problem (4.4) admits a principal eigenvalue,
$\lambda ^*(\mu )$
, which is associated with an eigenvector
$\boldsymbol {\phi }^*=(\phi _A^*,\phi _M^*,\phi _F^*)^T \in \mathrm {Int}(\mathcal {X}_+)$
. Further,
$\lambda ^*(\mu )$
can be expressed as
$ \lambda ^*(\mu )=- \frac {1}{T} \log r_{\mathcal {X}_+}(L_{\mu }(T,0)), $
and
$e^{-\lambda ^*(\mu ) t}\boldsymbol {\phi }^*(x,t)$
is a positive solution of system (4.3).
We define
no matter whether it is the principal eigenvalue of (4.4) or not. Now we can give a computation formula for two spreading speeds.
Theorem 4.3 Assume that
$\mathcal {R}_0>1$
. Then there holds
Proof We first introduce the following evolution system:
and the associated linear eigenvalue problem:
We next prove
$c_+^*=\inf _{\mu>0} -\frac {\lambda ^*(\mu )}{\mu }$
by dividing the proof into four steps, and
$c_-^*=\inf _{\mu>0} -\frac {\lambda ^*(-\mu )}{\mu }$
can be obtained similarly.
Step 1: The principal eigenvalue of the nonlinear eigenvalue problem (4.4) coincides with that of the linear eigenvalue problem (4.6). It then suffices to show the following two claims.
Claim 1. If
$\lambda ^*$
is the principal eigenvalue of eigenvalue problem (4.4), then it is also the principal eigenvalue of (4.6).
Claim 2. If
$\tilde {\lambda }$
is the principal eigenvalue of (4.6), then it is also the principal eigenvalue of (4.4).
We only prove Claim 1, and Claim 2 can be derived similarly. For any given
$\mu \geq 0$
, let
$(w_A^*,w_M^*,w_F^*)^T$
be the principal eigenvector of (4.4) corresponding to
$\lambda ^*$
, and let
$\hat {\lambda }$
be the principal eigenvalue of
Thanks to the Krein–Rutman theorem, it is easy to verify that
$\lambda ^*<\hat {\lambda }$
. Multiplying
$1-\theta $
and
$\theta $
on the second and third equations of (4.4), respectively, and then making difference, we obtain
Here,
$z=(1-\theta )w_M^*-\theta w_F^*$
is T-periodic in t and L-periodic in x. In view of
$\lambda ^*<\hat {\lambda }$
, we have
$z \equiv 0$
due to the Krein–Rutman theorem. Then we have
An easy computation yields that
$\lambda ^*$
and
$(w_A^*,w_M^*+w_F^*)$
satisfy (4.6), and the desired conclusion follows from [Reference Wang, Zhang and Zhao20, Corollary 2.6] or the Krein–Rutman theorem again.
Step 2. Systems (1.2) and (4.5) admit the same minimum wave speed of rightward time–space periodic traveling waves.
Let
An easy computation yields that
Then system (1.2) becomes
Substituting
$(u_A,u_S)(x,t):=(U_A,U_S)(x,t,\xi )\ \text {with}\ \xi =x-ct$
into (4.5), we obtain
We then have the following two claims:
Claim 3. If
$(U_A,U_S,c)$
satisfies (4.9), then
$(U_A,\theta U_S,(1-\theta )U_S,c)$
satisfies (4.8).
Claim 4. If
$(U_A,U_M,U_F,c)$
satisfies (4.8), then
$(U_A,U_M+U_F,c)$
satisfies (4.9).
Claim 3 can be obtained via a straightforward computation, and so we only prove Claim 4. Multiplying
$1-\theta $
and
$\theta $
on the second and third equations of (4.8), respectively, and then making difference, we obtain
where
$Z=(1-\theta )U_M-\theta U_F$
. Since Z is a bounded entire solution of (4.10), it is not hard to verify that Z converges to
$0$
uniformly for
$(x,\xi ) \in \mathbb {R}^2$
as
$t\rightarrow +\infty $
. Notice that Z is T-periodic in t. It then follows that
$Z \equiv 0$
for
$(x,t,\xi )\in \mathbb {R}^3$
. This implies that
$(1-\theta )U_M\equiv \theta U_F$
,
$(x,t,\xi )\in \mathbb {R}^3$
. Let
$U_S=\frac {1}{\theta }U_M=\frac {1}{1-\theta }U_F=U_M+U_F$
. A straightforward computation shows that Claim 4 holds.
Step 3. Prove the conclusion when
$\lambda ^*(\mu )$
is the principal eigenvalue of (4.4).
For any
$\mu \in \mathbb {R}$
, let
$\{Q_{\mu }(t,s)\}_{t\geq s}$
be the evolution family on
$\mathcal {X}$
generated by system (4.6) with
$\lambda =0$
and define
By the arguments similar to those for [Reference Fang, Lai and Wang6, Theorem 5.1] (see also [Reference Fang, Yu and Zhao7, Reference Liang and Zhao13]), the rightward spreading speed of (4.5) can be characterized by
$\inf _{\mu>0}-\frac {\tilde {\lambda }({\mu })}{\mu }$
, which coincides with the minimum wave speed of rightward almost pulsating waves of (4.5). By Step 2, systems (1.2) and (4.5) have the same minimum wave speed of rightward almost pulsating waves. It then follows from Step 1 that
Step 4. Complete the proof.
We take a series of functions
$d_{\epsilon }$
and domains
$\Omega _{\epsilon } \subseteq \Omega :=[0,L]$
such that
$d_{\epsilon } \leq d \leq d_{\epsilon }+\epsilon $
and
$\tilde {p}^{\epsilon }(x)=\max _{x \in \Omega } \tilde {p}^{\epsilon } (x)$
,
$\forall x \in \overline {\Omega }_{\epsilon }$
, where
$\tilde {p}^{\epsilon }(x)=T^{-1}\int _{0}^{T} [-d_{\epsilon }(x,t)-\gamma (x,t)] \mathrm {d} t$
,
$\forall x \in \overline {\Omega }$
. Then (4.4) admits the principal eigenvalue
$\lambda ^\epsilon (\mu )$
and
$\lambda _{\epsilon }(\mu )$
with d replaced by
$d_{\epsilon }$
and
$d_{\epsilon }+\epsilon $
, respectively. It is not hard to verify that
Therefore, the two corresponding perturbed systems admit the rightward spreading speeds
$c_{+}^{\epsilon }$
and
$c_{+,\epsilon }$
, respectively. Repeating the above three steps, we conclude that
Notice that
$-\frac {\lambda ^{\epsilon }(\mu )}{\mu } \rightarrow +\infty $
and
$-\frac {\lambda _{\epsilon }(\mu )}{\mu } \rightarrow +\infty $
as
$\mu \rightarrow 0$
and
$\mu \rightarrow +\infty $
. Thus, the two infima are attained at some finite values of
$\mu $
. It follows that
$ c_+^{\epsilon } \rightarrow c^*_+ $
and
$ c_{+,\epsilon } \rightarrow c^*_+ $
as
$\epsilon \rightarrow 0^+$
, which implies the desired result.
By the arguments similar to those in [Reference Liang and Zhao13, Proposition 7.4], we further have the following result.
Lemma 4.7 Assume that
$\mathcal {R}_0>1$
. Then
$c_+^*+c_-^*>0$
.
As a consequence of Lemmas 4.3, 4.5, and 4.7 and [Reference Liang and Zhao13, Theorem 5.1], we have the following result, which indicates that
$c_+^*$
and
$c_-^*$
are the rightward and leftward spreading speeds for system (1.2) with initial functions having compact supports, respectively.
Proposition 4.1 Assume that
$\mathcal {R}_0>1$
. Let
$\boldsymbol {u}(x,t;\boldsymbol {\phi })$
be a solution of (1.2) with
$\boldsymbol {u}(\cdot ,0;\boldsymbol {\phi })=\boldsymbol {\phi }=(\phi _A,\phi _M,\phi _F)^T\in \mathcal {C}_{\boldsymbol {u}^*(\cdot ,0)}$
. Then the following statements are valid:
-
(i) If $0\leq \boldsymbol {\phi }(\cdot )\leq \boldsymbol {\varphi } (\cdot )\ll \boldsymbol {u}^*(\cdot ,0)$
, for some
$\boldsymbol {\varphi } (\cdot )\in \mathcal {X}$
, and
$\phi _N(x)=0$
for x outside a bounded interval and
$N=A,M,F$
, then $$ \begin{align*}\lim_{t\rightarrow\infty}\sup_{x\geq ct}\Vert\boldsymbol{u}(x,t;\boldsymbol{\phi})\Vert=0,\ \text{for any} \ c>c_{+}^*,\end{align*} $$and
$$ \begin{align*}\lim_{t\rightarrow\infty}\sup_{x\leq - ct}\Vert\boldsymbol{u}(x,t;\boldsymbol{\phi})\Vert=0,\ \text{for any} \ c>c_{-}^* \ .\end{align*} $$
-
(ii) If $\boldsymbol {\phi }\in \mathcal {C}_{ \boldsymbol {u}^*(\cdot ,0)}$
and
$\boldsymbol {\phi } \not \equiv \boldsymbol {0}$
, then for any c and
$c'$
satisfying
$-c_{-}^*<-c' <c<c_{+}^*$
, there holds $$ \begin{align*}\lim_{t\rightarrow\infty}\sup_{-c't \leq x\leq ct}\Vert\boldsymbol{u}(x,t;\boldsymbol{\phi})-\boldsymbol{u}^*(x,t)\Vert=0.\end{align*} $$
4.3 Spreading speeds in autonomous case
In this section, we discuss the spreading speed quantitatively under the assumption that all parameters are independent of time (i.e., in the autonomous case).
Proposition 4.2 Assume all parameters are independent of time and
$\mathcal {R}_0>1$
. Then for any
$\mu \in \mathbb {R}$
, eigenvalue problem (4.4) admits a principal eigenvalue
$\lambda ^*(\mu )$
such that
$\lambda ^*(\mu )=\lambda ^*(-\mu )$
,
$\mu \in \mathbb {R}$
. Moreover,
$c_+^*=c_-^*=\inf _{\mu>0} -\frac {\lambda ^*(\mu )}{\mu }>0$
.
Proof Since
$\mathcal {R}_0>1$
and
$L_0(T,0)=\mathcal {U}(T,0)$
restricted on
${\mathcal {X}_+}$
, we have
$r_{\mathcal {X}_+}(L_{0}(T,0))> 1$
due to Lemma 4.1. In view of Lemma 2.1, there exists
$\tilde {\mu }$
such that
$r_{\mathcal {X}_+}(L_{\mu }(T,0))> 1$
for
$\mu \in (0,\tilde {\mu })$
. Thus, Lemma 4.6 implies that eigenvalue problem (4.4) admits a principal eigenvalue
$\lambda ^*(\mu )= -\frac {1}{T} \log r_{\mathcal {X}_+}(L_{\mu }(T,0))$
for any
$\mu \in (0,\tilde {\mu })$
. We still let
$\{Q_{\mu }(t,s)\}_{t\geq s}$
be the evolution family on
$\mathcal {X}$
generated by system (4.6) with
$\lambda =0$
and write
$\tilde {\lambda }({\mu })=-\frac {1}{T} \log r(Q_{\mu }(T,0))$
. According to Claim 1 in the proof of Theorem 4.3,
$\tilde {\lambda }(\mu )=\lambda ^*(\mu )$
is the principal eigenvalue of eigenvalue problem (4.6) for any
$\mu \in (0,\tilde {\mu })$
. We only need to prove that
$\tilde {\mu }$
can be extended to
$+\infty $
. It then suffices to prove the following three claims:
Claim 1.
$-\tilde {\lambda }(\mu )$
is convex on
$\mathbb {R}$
.
Claim 2.
$\lim \limits _{\mu \rightarrow +\infty } -\tilde {\lambda }(\mu )=+\infty $
.
Claim 3.
$\tilde {\lambda }(\mu )=\tilde {\lambda }(-\mu )$
,
$\forall \mu \in \mathbb {R}$
.
If these claims are valid, then
$\tilde {\lambda }(\mu )$
is non-increasing with respect to
$\mu \in (0,+\infty )$
, and hence,
$r(Q_{\mu }(T,0)) \geq 1$
for all
$\mu \geq 0$
. Thus, [Reference Fang, Lai and Wang6, Lemma 3.1] implies that
$\tilde {\lambda }(\mu )$
is the principal eigenvalue of (4.6). By Claim 2 in the proof of Theorem 4.3, we see that
$\lambda ^*(\mu )=\tilde {\lambda }(\mu )$
is the principal eigenvalue of eigenvalue problem (4.4) for any
$\mu \in (0,+\infty )$
. Combining with Claim 3, we can obtain the desired conclusions.
Indeed, Claim 1 follows from the proof of Theorem 4.3. For readers’ convenience, we provide more details. We take a series of functions
$d_{\epsilon }$
and define
$\lambda ^{\epsilon }(\mu )$
and
$\lambda _{\epsilon }(\mu )$
as those in the proof of Theorem 4.3. By Step 1 of Theorem 4.3, it is known that
$\lambda ^{\epsilon }(\mu )$
and
$\lambda _{\epsilon }(\mu )$
are also the principal eigenvalues of (4.6) with d replaced by
$d_{\epsilon }$
and
$d_{\epsilon }+\epsilon $
, respectively. According to [Reference Liang and Zhao12], we obtain that
$-\lambda ^{\epsilon }(\mu )$
and
$-\lambda _{\epsilon }(\mu )$
are convex on
$\mathbb {R}$
, so is
$-\tilde {\lambda }(\mu )$
by letting
$\epsilon \rightarrow 0^+$
.
In order to prove Claim 2, we choose four constants
$a_{11}$
,
$a_{12}$
,
$a_{21}$
, and
$a_{22}$
such that
and four constants
$b_{11}$
,
$b_{12}$
,
$b_{21}$
, and
$b_{22}$
such that
By the arguments in Lemma 3.2, it then follows that the following two eigenvalue problems
and
have the principal eigenvalues, which are denoted by
$\overline {\lambda }(\mu )$
and
$\underline {\lambda }(\mu )$
, respectively. By the comparison arguments, we have
$\overline {\lambda }(\mu ) \leq \tilde {\lambda }(\mu )\leq \underline {\lambda }(\mu ) $
,
$\forall \mu \in \mathbb {R}_+$
. A direct computation shows that
$\lim \limits _{\mu \rightarrow +\infty } -\frac {\overline {\lambda }(\mu )}{\mu ^2}=\lim \limits _{\mu \rightarrow +\infty } -\frac {\underline {\lambda }(\mu )}{\mu ^2}=D>0$
. This yields that
$\lim \limits _{\mu \rightarrow +\infty } -\tilde {\lambda }(\mu )=+\infty $
.
Finally, we prove Claim 3. Write
$m_{11}(x)=-d(x) -\gamma (x)$
,
$m_{12}(x)=\rho (x)\theta (1-\theta )$
,
$m_{21}(x)=\gamma (x)$
,
$m_{22}(x)=-\hat {d}(x) $
,
$\tilde {m}_{12}(x)=\tilde {m}_{21}(x)=[m_{12}(x)m_{21}(x)]^{\frac {1}{2}}$
,
$\forall x \in \mathbb {R}$
. Clearly, if
$\lambda $
,
$\phi _1$
, and
$\phi _2$
satisfy the following eigenvalue problem:
then
$\lambda $
,
$\psi _1=[m_{12}(x)]^{-\frac {1}{2}}[m_{21}(x)]^{\frac {1}{2}} \phi _1$
, and
$\psi _2=\phi _2$
satisfy the following eigenvalue problem:
Now Claim 3 follows from this observation and the adjoint eigenvalue problem.
In the rest of this section, we study the effects of diffusion coefficient on the spreading speeds. We use
$c^*(D)=c_+^*(D)=c_-^*(D)$
and
$\mathcal {R}_0(D)$
to denote the spreading speed and basic reproduction ratio of system (1.2), respectively, to emphasize that they depend on the diffusion coefficient D. From the analysis in Theorem 3.1, we have the following observation.
Lemma 4.8 Assume all parameters are independent of time. Then the following statements are valid:
-
(i) $\mathcal {R}_0(D)$
is decreasing with respect to
$D \in (0,+\infty )$
. -
(ii) $\mathcal {R}_0(D) \rightarrow \theta (1-\theta ) \max _{x \in [0,L]} \frac {\rho (x)\gamma (x)}{(d(x)+\gamma (x))\hat {d}(x)}=:\mathcal {R}_0(0)$
as
$D \rightarrow 0^{+}$
. -
(iii) $\mathcal {R}_0(D) \rightarrow \theta (1-\theta )\frac {\int _{0}^{L} \frac {\rho (x)\gamma (x)}{d(x)+\gamma (x)} \mathrm {d} x}{\int _{0}^{L} \hat {d} (x) \mathrm {d} x}:=\tilde {\mathcal {R}}_0$
as
$D \rightarrow +\infty $
.
We are ready to prove the following result on the spreading speed
$c^*(D)$
.
Theorem 4.4 Assume all parameters are independent of time. Then the following statements are valid:
-
(i) If $\mathcal {R}_0(0)>1$
, then there exists
$\underline {D}>0$
such that
$c^*(D)>0$
for all
$D\in (0,\underline {D})$
, and
$\lim \limits _{D \rightarrow 0^+} c^*(D)=0$
. -
(ii) If $\mathcal {R}_0(0)>1>\tilde {\mathcal {R}}_0$
, then there exists a unique
$D^*>0$
such that
$c^*(D)>0$
for all
$D\in (0,D^*)$
and
$\lim \limits _{D \rightarrow (D^*)^{-}} c^*(D)=0$
. -
(iii) If $\tilde {\mathcal {R}}_0>1$
, then
$c^*(D)>0$
for all
$D\in (0,+\infty )$
, and
$\lim \limits _{D \rightarrow +\infty } c^*(D)=+\infty $
.
Proof We denote
$\lambda ^*(D,\mu )$
the principal eigenvalue of (4.4) which depends on D and
$\mu $
. We remark that
$\lambda ^*(D,\mu )$
is also the principal eigenvalue of (4.6) due to Step 1 of Theorem 4.3.
(i) In view of Lemma 4.8(ii), there exists
$\underline {D}>0$
such that for all
$D\in (0,\underline {D})$
,
$\mathcal {R}_0(D)>1$
, and hence,
$c^*(D)>0$
due to Proposition 4.2. Let
$\overline {\lambda }^*(D,\mu )$
be the principal eigenvalue of eigenvalue problem (4.11). By the comparison arguments, it is easy to see that
$\overline {\lambda }^*(D,\mu ) \leq \lambda ^*(D,\mu ) $
,
$\forall (D,\mu ) \in \mathbb {R}^2_+$
. Combining with
$c^*(D)= \inf _{\mu>0} -\frac {\lambda ^*(D,\mu )}{\mu }$
, we only need to show that
It is easy to verify that
$\lim \limits _{D \rightarrow 0^+}\overline {\lambda }^*(D,\mu )$
exists for any
$\mu \geq 0$
, which is the smaller solution of the equation
$(\lambda +a_{11})(\lambda +a_{22})-a_{12}a_{21}=0$
. For convenience, we denote this limit by
$\overline {\lambda }$
. In view of
$\mathcal {R}_0(0)>1$
, it then follows that
$\frac {a_{12}a_{21}}{a_{11}a_{22}}>1$
, which implies
$\overline {\lambda }<0$
.
For any
$\epsilon>0$
, there exists
$M>0$
such that
$-\epsilon \leq \frac {-\overline {\lambda }+\epsilon }{M} \leq \epsilon $
. Moreover, there exists
$D_0>0$
such that
$\vert \overline {\lambda }^*(D,M)- \overline {\lambda } \vert \leq \epsilon $
for all
$D \in (0,D_0)$
. Therefore,
which implies that
$\lim \limits _{D \rightarrow 0^+}\inf _{\mu>0} -\frac {\overline {\lambda }^*(D,\mu )}{\mu }=0$
.
(ii) In view of Lemma 4.8, there exists
$D^*>0$
such that
$\mathcal {R}_0(D^*)=1$
, and
$\mathcal {R}_0(D)>1$
for all
$D\in (0,D^*)$
and
$\mathcal {R}_0(D)<1$
for all
$D>D^*$
. Proposition 4.2 implies that
$c^*(D)>0$
for all
$D\in (0,D^*)$
.
We next show that
$\frac {\partial }{\partial \mu } \lambda ^*(D^*,0)=0$
. For convenience, we use
$\dot {\lambda }$
,
$\dot {\psi }_1$
, and
$\dot {\psi }_2$
to represent
$\frac {\partial }{\partial \mu } \lambda $
,
$\frac {\partial }{\partial \mu } \psi _1$
, and
$\frac {\partial }{\partial \mu } \psi _2$
, respectively. Differentiating (4.13) with respect to
$\mu $
, we obtain
Multiplying the first equation of (4.14) by
$\psi _1$
and the second equation of (4.14) by
$\psi _2$
and summing them together, and multiplying the first equation of (4.13) by
$\dot {\psi }_1$
and the second equation of (4.13) by
$\dot {\psi }_2$
and summing them together, and integrating them from
$0$
to L, and then making a difference, we further have
Letting
$\mu =0$
and
$D=D^{*}$
, we see that
$\frac {\partial }{\partial \mu } \lambda ^*(D^*,0)=0$
.
Notice that
$\lambda ^*(D^*,0)=0$
due to
$\mathcal {R}_0(D^*)=1$
. For any sequence
$D_n \rightarrow D^{*}$
with
$D_n<D^{*}$
as
$n \rightarrow \infty $
, there exists
$\mu _n \rightarrow 0^+$
such that
$\frac {\lambda ^*(D_n,\mu _n)}{\mu _n} \rightarrow 0$
as
$n \rightarrow \infty $
. It follows that
$\inf _{\mu>0} -\frac {\lambda ^*(D_n,\mu )}{\mu } \leq 0$
as
$n \rightarrow \infty $
. Therefore,
$c^*(D_n) \rightarrow 0$
as
$n \rightarrow \infty $
.
(iii) In view of Lemma 4.8(i) and (iii), we have
$\mathcal {R}_0(D)>1$
, which implies that
$c^*(D)>0$
for all
$D\in (0,+\infty )$
by Proposition 4.2. We next prove the following two claims:
Claim 1. There exist
$\rho _0>0$
and
$\overline {D}_1>0$
such that
$-\lambda ^*(D,\mu ) \geq \rho _0$
holds uniformly for
$\mu \in [0,+\infty )$
when
$ D>\overline {D}_1$
.
Claim 2. For any given
$\epsilon _0>0$
,
$\lim \limits _{D \rightarrow +\infty } -\frac {\lambda ^*(D,\mu )}{D}=\mu ^2$
holds uniformly on
$[\epsilon _0,+\infty )$
, that is, for every
$\epsilon>0$
, there exists
$\overline {D}_2>0$
large enough such that
$\vert -\frac {\lambda ^*(D,\mu )}{D}-\mu ^2 \vert \leq \epsilon $
for all
$\mu \in [\epsilon _0,+\infty )$
when
$D>\overline {D}_2$
.
We postpone the proof of these two claims and complete the proof in a few lines. For any given
$M>0$
, there exists
$\delta _0>0$
such that
$\frac {\rho _0}{\delta _0} \geq M$
due to
$\rho _0>0$
. Thus,
$\frac {\rho _0}{ \mu } \geq M$
,
$\mu \in [0,\delta _0]$
. According to Claim 1, there exists
$D_1$
such that
$\frac {-\lambda ^*(D,\mu )}{\mu } \geq \frac {\rho _0}{ \mu } \geq M$
holds for all
$\mu \in (0,\delta _0]$
when
$D \geq D_1$
. By Claim 2, we can choose
$D_2$
large enough such that
$\frac {M}{D_2} \leq \frac {\delta _0}{2}$
, and
$-\frac {\lambda ^*(D,\mu )}{D} \geq \frac {\delta _0}{2} \mu \geq \frac {M}{D_2} \mu $
for all
$\mu \in [\delta _0,+\infty )$
when
$D \geq D_2$
. Therefore,
$-\frac {\lambda ^*(D,\mu )}{\mu } \geq \frac {MD}{D_2} \geq M$
for all
$\mu \in [\delta _0,+\infty )$
when
$D \geq D_2$
. Thus,
$\frac {-\lambda ^*(D,\mu )}{\mu } \geq M$
for all
$\mu \in (0,\infty )$
when
$D \geq \max (D_1,D_2)$
.
Note that
$-\lambda ^*(D,0)$
is the principal eigenvalue of (4.13) with
$\mu =0$
. It is easy to verify that
$-\lambda ^*(D,0)$
is non-increasing with respect to
$D \in (0,+\infty )$
. In view of
$\tilde {\mathcal {R}}_0>1$
, we have
$\theta (1-\theta )\int _{0}^{L} \frac {\rho (x)\gamma (x)}{d(x)+\gamma (x)} \mathrm {d} x>\int _{0}^{L} \hat {d} (x) \mathrm {d} x$
. This implies that
$\theta (1-\theta )\int _{0}^{L} \frac {\rho (x)\gamma (x)}{d(x)+\gamma (x)+\varrho } \mathrm {d} x=\int _{0}^{L} \hat {d} (x) \mathrm {d} x +\varrho L$
has a unique positive root
$\varrho _1$
. It then follows that
$-\lambda ^*(D,0)$
converges to
$\varrho _1$
as
$D \rightarrow +\infty $
. Thus, there exists
$\overline {D}_1$
such that
$-\lambda ^*(D,0)>\rho _0$
whenever
$D>\overline D_1$
for some
$\rho _0>0$
. By the proof of Proposition 4.2, we see that
$-\lambda ^*(D,\mu )$
is non-decreasing with respect to
$\mu \in (0,+\infty )$
. This proves Claim 1.
Let
$\underline {\lambda }^*(D,\mu )$
be the principal eigenvalue of the eigenvalue problem (4.12). By the comparison arguments, it is easy to see that
where
$\overline {\lambda }^*(D,\mu )$
is the principal eigenvalue of (4.11). It suffices to show that
$\lim \limits _{D \rightarrow +\infty } -\frac {\overline {\lambda }^*(D,\mu )}{D}= \lim \limits _{D \rightarrow +\infty } -\frac {\underline {\lambda }^*(D,\mu )}{D}=\mu ^2 $
holds uniformly on
$[\epsilon _0,+\infty )$
. Here, we prove that
$\lim \limits _{D \rightarrow +\infty } -\frac {\underline {\lambda }^*(D,\mu )}{D}=\mu ^2 $
holds uniformly on
$[\epsilon _0,+\infty )$
, and the other limit can be proved similarly. Since
$\underline {\lambda }^*(D,\mu )$
is the smaller solution of the equation
it follows that
A straightforward computation gives rise to
which implies Claim 2.
Finally, we remark that it remains an unsolved problem to determine the limiting behavior of the propagation speed as the diffusion coefficient approaches infinity in the critical case where
$\tilde {\mathcal {R}}_0=1$
.