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Articles

A Wolbachia infection model with free boundary

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Pages 515-542 | Received 17 Jul 2018, Accepted 06 Jun 2020, Published online: 29 Jun 2020

Abstract

Scientists have been seeking ways to use Wolbachia to eliminate the mosquitoes that spread human diseases. Could Wolbachia be the determining factor in controlling the mosquito-borne infectious diseases? To answer this question mathematically, we develop a reaction-diffusion model with free boundary in a one-dimensional environment. We divide the female mosquito population into two groups: one is the uninfected mosquito population that grows in the whole region while the other is the mosquito population infected with Wolbachia that occupies a finite small region. The mosquito population infected with Wolbachia invades the environment with a spreading front governed by a free boundary satisfying the well-known one-phase Stefan condition. For the resulting free boundary problem, we establish criteria under which spreading and vanishing occur. Our results provide useful insights on designing a feasible mosquito releasing strategy that infects the whole mosquito population with Wolbachia and eradicates the mosquito-borne diseases eventually.

1. Introduction

Recently, several public health projects were launched in China [Citation27], USA [Citation1] and France [Citation22], with an aim to fight mosquito populations that transmit Zika virus, Dengue fever and Chikungunya. All of these projects involve the release of male Aedes aegypti mosquitoes infected with the Wolbachia bacteria to the wild. For instance, 20000 male Aedes aegypti mosquitoes carrying Wolbachia bacteria were released on Stock Island of the Florida Keys in the week of April 20, 2017. Google's Verily is about to release 20 million machine-reared Wolbachia-infected mosquitoes in Fresno (see [Citation1]). A factory in Southern China is manufacturing millions of ‘mosquito warriors’ (male Aedes aegypti mosquitoes carrying Wolbachia bacteria) to combat epidemics transmitted by mosquitoes [Citation27].

The science behind these projects is based on the following two facts: (i) Wolbachia often induces cytoplasmic incompatibility (CI) which leads to early embryonic death when Wolbachia-infected males mate with uninfected females and (ii) Wolbachia-infected females produce viable embryos after mating with either infected or uninfected males, resulting in a reproductive advantage over uninfected females. In practice, Wolbachia has been successfully transferred into Aedes aegypti or Aedes albopictus by embryonic microinjections, and the injected infection has been stably maintained with complete CI and nearly perfect maternal transmission [Citation2,Citation16,Citation17,Citation23,Citation31,Citation32,Citation34]. Thus, the bacterium is expected to invade host population easily driving the host population to decline. Successful Wolbachia invasion in Aedes aegypti has been observed by Xi et al. in the laboratory caged population within seven generations [Citation33].

By releasing Aedes albopictus mosquitoes infected with Wolbachia bacteria into the wild, it is expected that over a long time period, the wild Aedes aegypti mosquito population would decline drastically and hopefully be completely replaced by infected mosquitoes so that the mosquito-borne infectious diseases such as Zika, Dengue fever and Chikungunya would be eradicated. To qualitatively examine if Wolbachia can effectively invade the wild uninfected mosquito population, Zheng, Tang and Yu [Citation38] considered the following model: (1) dudt=u[b1δ1(u+v)]for t>0,dvdt=vb2vu+vδ2(u+v)for t>0,(1) where u denotes the number of reproductive infected insects and v denotes uninfected ones, b1 and b2 denote half of the constant birth rates for the infected and uninfected insects respectively. The parameter δ1 (resp. δ2) denotes the density-dependent death rate for the infected (resp. uninfected) population. The birth rate of uninfected mosquitoes is diminished by the factor v/(u+v) due to the sterility caused by cytoplasmic incompatibility (CI) for mating between infected males and uninfected females.

Let us now recall the origin of system (Equation1) with some details. Let rf and rm denote the number of released female mosquitoes and the number of released males, respectively, and suppose the released mosquitoes were infected with Wolbachia. Also, assume that rf and rm satisfy (2) drfdt=δ1rfT(t),t>0,drmdt=δ1rmT(t),t>0,(2) where T(t)=rf+rm+If+Im+Uf+Um denotes the total population size, with Uf, Um, If and Im standing for the numbers of uninfected reproductive females, uninfected reproductive males, and infected reproductive females and males other than those from releasing, respectively. Let bI (resp. bU) be the natural birth rate of the infected (resp. uninfected) mosquitos and 0δ1 be the proportion of mosquitos born female. Then the proportion of mosquitos born male is 1δ. With complete CI (see Table ) and perfect maternal transmission, we have (3) dIfdt=δbI[If+rf]δ1IfT(t),t>0,dImdt=(1δ)bI[If+rm]δ1ImT(t),t>0,dUfdt=δbUUfUmrm+Im+Umδ2UfT(t),t>0,dUmdt=(1δ)bUUfUmrm+Im+Umδ2UmT(t),t>0.(3)

Table 1. Strong CI, × means ‘no offspring’.

One can easily verify that both rf and rm approach 0 as t+. We denote by (4) u(t)=If+Imandv(t)=Uf+Um.(4) Assuming equal determination case, which means that δ=1/2, If=Im and Uf=Um, then system (Equation1) can be obtained by setting b1=bI/2 and b2=bU/2. In order to obtain the spatiotemporal dynamics of (Equation1), Huang et al. [Citation14,Citation15] studied the following reaction-diffusion system: (5) ut=d1Δu+u(b1δ1(u+v)),t>0,xΩ,vt=d2Δv+vb2vu+vδ2(u+v),t>0,xΩ,uν=vν=0,t>0,xΩ,u(0,x)=u0(x),v(0,x)=v0(x),xΩ.(5) In (Equation5), d1 and d2 are the diffusion rates, Δ denotes the Laplace operator in the spatial variable x, and ν denotes the unit outward normal vector to the boundary of Ω. We mention that (Equation5) is obtained from a delay differential equation model in [Citation38] after ignoring the delay factor and incorporating the spatial inhomogeneity. Similarly, there has been several mathematical models formulated to describe the Wolbachia spreading dynamics [Citation13,Citation35,Citation36,Citation39]. These models focused on studying the subtle relation between the threshold releasing level for Wolbachia-infected mosquitoes and several important parameters including the CI intensity and the fecundity cost of Wolbachia infection.

We also note that female Aedes aegypti mosquitoes infected with the Wolbachia bacteria were initially released at a specific site. Hence, the infected female mosquitoes initially occupy only a small region, while the wild uninfected females are distributed over the whole area.

To model the spatial spreading of Wolbachia in the wild mosquito population and explore the possibility that the infection can indeed occupy the whole region, it is natural to consider system (Equation5) under the setting of a free boundary problem.

In this work, we consider the following free boundary problem in one-dimensional space: (6) ut=d1uxx+u(b1(x)δ1(u+v)),t>0,0<x<h(t),vt=d2vxx+vb2(x)vu+vδ2(u+v),t>0,x>0,ux(t,0)=vx(t,0)=0,u(t,h(t))=0,t>0,h(t)=μux(t,h(t)),t>0,h(0)=h0,u(0,x)=u0(x),x[0,h0],v(0,x)=v0(x),x[0,+).(6) The equation governing the movement of the spreading front x=h(t) is deduced in a manner similar to that in Section 1.3 of  [Citation3]. It is known as the one-phase Stefan condition in the literature. This type of free boundary condition has been widely used in previous work such as [Citation5–7,Citation10,Citation18–20,Citation24,Citation28–30].

We will first analyse system (Equation6) with constant birth rates b1 and b2 in Section 3. Environmental variables such as available water surfaces and humidity have huge impacts on birth rates [Citation8]. This is why we also extend our study to the case with space-dependent birth rates b1(x) and b2(x) in Section 4, while we assume that the natural death rate is spatially independent for simplicity.

Throughout this paper, we assume that b1(x) and b2(x) satisfy the following conditions, unless otherwise stated: θ(0,1) such that biC0,θ[0,+)L[0,+),bi0, i=1,2.(B1)C0,θ([0,+)) is the Hölder space with Hölder exponent θ. The initial conditions u0 and v0 are assumed to be bounded and satisfy (7) u0C2([0,h0]),u0(0)=u0(h0)=0,u0(x)>0for all x(0,h0),v0C2[0,)L[0,)andv0>0.(7) For the free boundary problem (Equation6)–(Equation7), the main question we are concerned about is whether the infected population can eventually occupy the whole space or not.

Definition 1.1

The notion of vanishing and spreading

If the infected population eventually occupies the whole space, i.e. limth(t)=+, we say spreading occurs; otherwise, we say vanishing occurs.

The main goal of this work is to derive conditions under which the spreading occurs. If spreading occurs, then the whole mosquito population will become infected with Wolbachia bacteria and this leads to the extinction of the mosquito population and eventually the eradication of mosquito-borne diseases.

Organization of the paper. The paper is organized as follows. We first establish the global existence and uniqueness of solutions to the free boundary problem (Equation6) in Section 2. In Section 3, we present a detailed analysis of a specific case of model (Equation6). In Section 4, we study the population dynamics of infected mosquitoes in a heterogeneous environment with a free boundary condition. In order to better understand the effects of dispersal and spatial variations on the outcome of the competition, we study system (Equation6) over a bounded domain with Neumann boundary conditions. We summarize our results in the last section.

2. Global existence of smooth solutions

Using arguments that are similar to those in [Citation11], we can establish the following result concerning the existence and uniqueness of solutions to system (Equation6)–(Equation7).

Theorem 2.1

Local existence

Consider system (Equation6) with initial conditions (Equation7). Assume that b1 and b2 satisfy (B1). Then, there exists T>0 such that (Equation6) admits a unique solution (u,v,h(t)) satisfying

  1. (u,v,h)C(1+θ)/2,1+θ(Q)×C(1+θ)/2,1+θ(Q)×C1+θ/2([0,T]),

  2. uC(1+θ)/2,1+θ(Q)+vC(1+θ)/2,1+θ(Q)+hC1+θ/2([0,T])K,

where 0<θ<1 is the Hölder exponent in (B1), Q={(t,x)R2, such that t[0,T] and x[0,h(t)]},Q={(t,x)R2, such that t[0,T] and x[0,+)},K and T are constants that depend only on h0, θ, u0C2([0,h0]) and v0C2([0,+)).

The next result provides some bounds on the solutions to system (Equation6) with initial conditions (Equation7).

Lemma 2.1

Let (u,v,h) be a solution of (Equation6) for t[0,T] for some T>0. Then,

  1. 0<u(t,x)M1 for all t(0,T] and x[0,h(t)), where M1:=maxb1L([0,))δ1, u0L([0,h0]).

  2. 0<v(t,x)M2 for all t(0,T] and x[0,+), where M2:=maxb2L([0,))δ2, v0L([0,+)).

  3. 0<h(t)Λ for all t(0,T], where Λ>0 depends on μ, d1, u0L([0,h0]) and u0C[0,h0].

Proof.

The strong maximum principle yields that u(t,x)>0 for all t(0,T] and x[0,h(t)), and v(t,x)>0 for all t(0,T] and x[0,+). Note that u(t,h(t))=0 yields that ux(t,h(t))<0for all t(0,T]. Thus, h(t)>0 for t(0,T]. Next, we consider the initial value problem (8) u(t)=u(t)(b1L([0,))δ1u(t)), for t>0,u(0)=u0L([0,h0]).(8) From the comparison principle, we know that u(t,x)maxb1L([0,))δ1, u0L([0,h0]). Similarly, we can show that v(t,x)maxb2L([0,))δ2, v0L([0,+)). To prove (iii), we first consider the auxiliary function (9) ω1(t,x):=M12M(h(t)x)M2(h(t)x)2(9) for t[0,T] and x[h(t)M1,h(t)], where M=max1h0,b1L([0,))2d1, u0C[0,h0]M1. We have (10) ω1td1ω1xx2d1M1M2b1M1u[b1δ1(u+v)]=utd1uxx,ω1(t,h(t))=0=u(t,h(t)),ω1(t,h(t)M1)=M1u(t,h(t)M1).(10) We also note that u0(x)=xh0u0(s)ds(h0x)u0C[0,h0] and ω1(0,x)=M1M(h0x)[2M(h0x)]M1M(h0x),for x[h0M1,h0]. Thus, ω1(0,x)u(0,x). Applying the comparison principle, we get ω1(t,x)u(t,x),for t[0,T]andx[h(t)M1,h(t)]. Since ω1(t,h(t))=0=u(t,h(t)), we then have ux(t,h(t))ω1x(t,h(t))=2MM1. Consequently, h(t)=μux(t,h(t))Λ with Λ:=2μMM1.

Bearing the above result in mind, we can show that the local solution obtained in Theorem 2.1 can indeed be extended to all t>0.

Theorem 2.2

Global existence and uniqueness

System (Equation6)–(Equation7) admits a unique solution for t[0,).

Proof.

Let [0,Tmax) be the maximal time interval in which the unique solution exists. We will show that Tmax=. Suppose to the contrary that Tmax<. In view of Lemma 2.1, there exists positive constants M1, M2 and Λ, independent of Tmax, such that for t[0,Tmax], 0<u(t,x)M1,0<v(t,x)M2and0<h(t)Λ. Fix δ(0,Tmax) and K>Tmax. Using the standard Lp estimates together with the Sobolev embedding theorem and the Hölder estimates for parabolic equations (see Lunardi [Citation21] for eg.), we can find M3 depending only on δ, K, M1 and M2 such that u(t,)C1+θ[0,h(t))M3andv(t,)C1+θ[0,+)M3for all t[δ,Tmax), where we used the convention that u(t,x)=0 for xh(t). By virtue of the proof of Theorem 2.1 in [Citation11], there exists a τ>0 depending only on M1, M2 and M3 such that the solution of (Equation6) with the initial time Tmaxτ/2 can be extended uniquely to the time Tmax+τ/2, which contradicts the definition of Tmax. Thus, Tmax=+ and the proof is complete.

3. The special case of constant birth rates

System (Equation5) was investigated in [Citation14,Citation15] for two disjoint cases. Namely, the fitness benefit case and the fitness cost case. Define κ1 and κ2 as κ1=b1/δ1 and κ2=b2/δ2. Wolbachia is said to have the fitness benefit if κ1>κ2, which means that the local area is more (or at least equally) favourable for infected mosquitoes. The fitness cost case is represented by κ1<κ2, see [Citation38].

In this section, we assume that bi(x)=bi for i = 1, 2, where bi are positive constants. In other words, we have the constant-coefficient free boundary problem given by (11) ut=d1uxx+u(b1δ1(u+v)),t>0,0<x<h(t),vt=d2vxx+vb2vu+vδ2(u+v),t>0,x>0,ux(t,0)=vx(t,0)=0,u(t,h(t))=0,t>0,h(t)=μux(t,h(t)),t>0,h(0)=h0,u(0,x)=u0(x),x[0,h0],v(0,x)=v0(x),x[0,+).(11) System (Equation11) is essentially a competition model. For the fitness benefit case, κ1>κ2, u is the so-called superior competitor and v the inferior competitor (see [Citation11]). For the fitness cost case, κ1<κ2, (Equation11) represents a strong competition [Citation25]. Throughout this section, we always assume u is a superior competitor. That is, the Wolbachia infection has a fitness benefit. The strong competition case is usually more complicated to be studied mathematically. To the best of our knowledge, results for competition models with a free boundary are very limited in strong competition case. Further details can be seen in [Citation41,Citation42].

We organize this section as follows. In Subsection 3.1 we present some preliminary results, which play a role in proving our main results. Subsection 3.2 is devoted to the vanishing case. The invasion dynamics is studied in detail in Subsection 3.3. A rough estimation of asymptotic spreading speed of Wolbachia invasion is given in Subsection 3.4.

3.1. Preliminary results

Consider the system (12) ut=d1uxx(t,x)+u(t,x)(b1δ1u(t,x)),t>0,0<x<L,ux(t,0)=u(t,L)=0,t(0,).(12) The following result holds.

Lemma 3.1

Let L=π/2d1/b1 and d=4b1L2/π2. Then,

  1. if LL, all positive solutions of (Equation12) tend to zero in C([0,L]) as t+.

  2. If L>L, there exists a unique positive stationary solution φ of (Equation12) such that all positive solutions of (Equation12) approach φ in C([0,L]) as t+.

Proof.

(i) and (ii) follow from Propositions 3.1, 3.2 and 3.3 of  [Citation4].

We recall the following comparison principle.

Lemma 3.2

Comparison principle [Citation11]

Assume that 0T0<T<+ and h¯,h_C1([T0,T]). Denote by GT={(t,x)R2:t(T0,T], x(0,h_)} and GT1={(t,x)R2:t(T0,T] and x(0,h¯)}. Let u_C(GT¯)C1,2(GT),u¯C(GT1¯)C1,2(GT1) and v¯,v_LC([T0,T]×[0,+))C1,2((T0,T]×[0,+)). Suppose that (13a) u¯td1u¯xxδ1u¯(κ1u¯v_),T0<tT,0<x<h¯(t),u_td12u_x2δ1u_(κ1u_v¯),T0<tT,0<x<h_(t),v¯td2v¯xxδ2v¯κ2v¯u_+v¯v¯u_,T0<tT,x>0,v_td22v_x2δ2v_κ2v_u¯+v_v_u¯,T0<tT, x>0,(13a) (13b) h_(t)μu_x(t,h_(t)),T0<tT,h¯(t)μu¯x(t,h¯(t)),T0<tT,(13b) and (13c) u¯x(t,0)0,u¯(t,h¯(t))=0,T0<tT,xu_(t,0)0,u_(t,h_(t))=0,T0<tT,v¯x(t,0)0,v_x(t,0)0,T0<tT,h_(T0)h(T0)h¯(T0),u_(T0,x)u(T0,x)u¯(T0,x),0xh(T0),v_(T0,x)v(T0,x)v¯(T0,x),x0.(13c) Let (u,v,h) be the unique solution of (Equation11). Then,

  1. h(t)h¯(t), u(t,x)u¯(t,x) and v(t,x)v_(t,x) for all (t,x) in (T0,T]×[0,+).

  2. h(t)h_(t), u(t,x)u_(t,x) and v(t,x)v¯(t,x) for all (t,x) in (T0,T]×[0,+).

The following follows from Lemmas A.2 and A.3 in [Citation37].

Lemma 3.3

  1. Let a, b and q be fixed positive constants. For any given ε>0 and L>0, there exists l>maxL,π2dasuch that, if the continuous and non-negative function U(t,x) satisfies (14) UtdUxxU(abU),t>0,0<x<l,Ux(t,0)=0,U(t,l)q,t>0,(q0),(14) with U(0,x)>0 for all x[0,l), then lim inft+U(t,x)>abε uniformly on [0,L].

  2. Let a, b and q be fixed positive constants. For any given ε>0 and L>0, there exists l>max{L,π/2d/a} such that lim supt+V(t,x)<ab+ε uniformly on [0,L],where V(t,x) is a continuous and non-negative function satisfying (15) VtdVxxV(abV),t>0,0<x<l,Vx(t,0)=0,V(t,l)q,t>0,(q0),(15) and V(0,x)>0 for all x[0,l).

We are now in the position to present part of our main results.

3.2. The vanishing case

We consider the vanishing case in this subsection.

Theorem 3.1

Let (u,v,h) be the solution of system (Equation11) with initial data (Equation7). If h<+, then limt+u(t,)C[0,h(t)]=0andlimt+v(t,x)=κ2 uniformly in any bounded subset of [0,+).

Proof.

Theorem 2.1 yields that for θ(0,1), there is a constant Cˆ depending on θ, (u0,v0), h0 and h such that (16) uC(1+θ)/2,1+θ(G)+vC(1+θ)/2,1+θ(G)+h(t)C1+θ/2([0,))Cˆ,(16) where G:={(t,x)[0,)×[0,h(t)]}. Suppose that lim supt+u(t,)C([0,h(t)])=ε>0. Then, there exists a sequence (tk,xk) in (0,)×[0,h(t)], where tk as k, such that u(tk,xk)ε2 for all kN. Note that 0xk<h(tk)<h<. By passing to a subsequence if necessary, it follows that xkx0(0,h) as k. Define uk(t,x):=u(t+tk,x)andvk(t,x)=v(t+tk,x) for t(tk,) and x[0,h(t+tk)]. It follows from (Equation16) and standard parabolic regularity that {(uk,vk)} has a subsequence {(uki,vki)} satisfying (uki,vki)(u~,v~) as ki, where (u~,v~) is the solution to the following system (17) u~t=d1u~xx+u~b1δ1(u~+v~),(t,x)(,+)×(0,h),v~t=d2v~xx+v~b2v~u~+v~δ2(u~+v~),(t,x)(,+)×(0,h),(17) with u~(t,h)=0 for all tR. Since u~(0,x0)=limkiuki(0,xki)=limkiu(tki,xtki)ε2, the maximum principle implies that u~>0 in (,+)×(0,h). Hence, we can apply Hopf Lemma at the point (0,h) to obtain u~x(0,h)<0. Therefore, we have ux(tki,h(tki))=xuki(0,h(tki))<0 for large i. This, together with the Stefan condition, implies that h(tki)>0 .

On the other hand, h<+ implies h(t)0 as t (see Lemma 3.3 in [Citation12]). This is a contradiction. Thus, limt+u(t,)C[0,h(t)]=0. Next, we prove that limt+v(t,x)=κ2. Having limt+u(t,)C[0,h(t)]=0 implies that, for any ε(0,1), there exists T>0 such that 0u(t,x)ε for all t>T and x(0+). Thus, (18) vtd2vxx+vb2vε+vδ2(ε+v),t>T,x>0,vx(t,0)=0,v(t,+)0,t>T,v(T,x)>0.(18) By Lemma 3.3 and the arbitrariness of ϵ, we have lim inft+v(t,x)b2/δ2=κ2 uniformly in any bounded subset of [0,+). This, together with the fact lim supt+v(t,x)κ2, shows that limt+v(t,x)=κ2.

3.3. The invasion dynamics

Theorem 3.2

Suppose (u,v,h) is the solution of system (Equation11) under conditions (Equation7). If h=+, then limt+u(t,x)=κ1 and limt+v(t,x)=0 uniformly in any compact subset of [0,+).

Proof.

Consider the system (19) u~(t)=δ1u~(κ1u~),t>0,u~(0)=u0L([0,h0]).(19) Then, limt+u~(t)=κ1 and u(t,x)u~(t). Consequently, we have lim supt+u(t,x)κ1 uniformly for x[0,+). In a similar manner, we can obtain that lim supt+v(t,x)κ2 uniformly for x[0,+). Since κ1>κ2 then, for δ=(κ1κ2)/2, there exists T1>0 such that v(t,x)κ2+δ for all t>T1 and x0. If h=+, then for any given L, there exists l>{L,π/2d1/δ1δ} such that u satisfies (20) utd1uxx+δ1u(δu),t>T1,0<x<l,ux(t,0)=0,u(t,l)0,t>T1,u(T1,x)>0,0<x<l.(20) By Lemma 3.3, we know that for sufficiently small ε>0, lim inft+u(t,x)>δε uniformly in any compact subset of [0,L]. Since h=+, there exists T2>T1 such that h(T2)>L and u(t,x)δ/2 for all t>T2 and 0x<L. Then, (u,v) satisfies (21) ut=d1uxx+u[b1δ1(u+v)],t>T2,0<x<L,vt=d2vxx+vb2vu+vδ2(u+v),t>T2,0<x<L,ux(t,0)=vx(t,0)=0,t>T2,u(T2,x)δ2,v(T2,x)κ2+δ,0<x<L.(21) Let (u_,v¯) be the solution to the following problem: (22) u_t=d1u_xx+u_[b1δ1(u_+v¯)],t>T2,0<x<L,v¯t=d2v¯xx+v¯b2v¯u_+v¯δ2(u_+v¯),t>T2,0<x<L,xu_(t,0)=v¯x(t,0)=0,t>T2,u_(t,L)=δ2,v¯(t,L)=κ2+δ,t>T2,u_(T2,x)=δ2,v¯(T2,x)=κ2+δ,0xL.(22) It follows from the comparison principle that u(t,x)u_(t,x)andv(t,x)v¯(t,x)for t>T2and0xL. By Corollary 3.6 of [Citation26], we have limt+u_(t,x)=u_L(x)andlimt+v¯(t,x)=v¯L(x)uniformly in [0,L]. Here, (u_L,v¯L) satisfies (23) d1xxu_L+u_L[b1δ1(u_L+v¯L)]=0,0<x<L,d2xxv¯L+v¯Lb2v¯Lu_L+v¯Lδ2(u_L+v¯L)=0,0<x<L,xu_L(0)=xv¯L(0)=0,u_L(L)=δ2,v¯L(L)=κ2+δ.(23) Letting L+, it follows from standard elliptic regularity and a diagonal procedure that (u_L(x),v¯L(x)) converges to (u_(x),v¯(x)) uniformly on any compact subset of [0,+), where (u_,v¯) satisfies (24) d1xxu_+u_[b1δ1(u_+v¯)]=0,x>0d2xxv¯+v¯b2v¯u_+v¯δ2(u_+v¯)=0,x>0xu_(0)=xv¯(0)=0,u_(x)δ2,v¯(x)κ2+δ,0<x<+.(24) We consider now the following system: (25) du1dt=u1(b1δ1(u1+v1)),t>0,dv1dt=v1b2v1u1+v1δ2(u1+v1),t>0,u1(0)=δ2,v1(0)=κ2+δ.(25) Since κ1>κ2, then (u1,v1)(κ1,0) as t+ (see Lemma 2.2 of [Citation38], for e.g.). Then, the solution (U,V) of the problem (26) Ut=d1Uxx+U(b1δ1(U+V)),t>0,x0,Vt=d2Vxx+Vb2VU+Vδ2(U+V),t>0,x0,Ux(t,0)=Vx(t,0)=0,t>0,U(0,x)=δ2,V(0,x)=κ2+δ,x0.(26) satisfies (U(t,x),V(t,x))(κ1,0), as t+, uniformly in x[0,+). By the comparison principle, we have u_U and v¯V for t0, which immediately yields that limt+u(t,x)=κ1andlimt+v(t,x)=0. The criteria for spreading and vanishing are given in the following theorem.

Theorem 3.3

If h0π/2d1/δ1(κ1κ2):=h0, then h=+.

Proof.

Note that h(t) is nondecreasing. We only need to show that h<+ implies hh0. It follows from Theorem 3.1 that h<+ implies limt+u(t,)C[0,h(t)]=0andlimt+v(t,x)=κ2 uniformly in any bounded subset of [0,+). Assume that h>h0. Then for sufficiently small ε>0, there exists T>0 such that h(t)>π2d1δ1(κ1κ2)εandv(t,x)κ2+εδ1for tTandx[0,+). Let u_ be the solution of the following problem (27) u_td1u_xx=δ1u_κ1κ2εδ1u_,t>T,0<x<h(T),u_x(t,0)=0=u_(t,h(T)),t>T,u_(T,x)=u(T,x),0<x<h(T).(27) By the comparison principle, we have u_(t,x)u(t,x) for all tT and x[0,h(T)]. Since h(t)>π/2d1/δ1(κ1κ2)ε for t>T then, by Lemma 3.1, we know that limt+u_=U_>0 uniformly in any compact subset of (0,h(T)), where U_ is the unique positive solution of (28) d1U_xx=δ1U_κ1κ2εδ1U_,0<x<h(T),U_x(t,0)=0=U_(t,h(T)).(28) Thus, lim inft+u(t,x)limt+u_(t,x)=U_(x)>0, which is a contradiction. Therefore, hh0 and this completes the proof.

Theorem 3.4

If h0<h0, then there exists μ¯>0 such that h=+ as μμ¯.

Proof.

Since lim supt+v(t,x)κ2+ε uniformly for x[0,+), then there exists T1>0 such that v(t,x)κ2 when t>T1. So, (u,h) satisfies (29) utd1uxx+δ1u[κ1κ2u],t>T1,0<x<h(t),h(t)=μux(t,h(t)),t>T1,ux(t,0)=0,u(t,h(t))=0,t>T1,u(T1,x)>0,0<x<h(T1).(29) Note that, u(T1,x) depends on μ. So, we consider the following problem. (30) u~(t,x)t=d1u~xx+u~(b1δ1(u~+v~)),t>0,0<x<h0,v~(t,x)t=d2v~xx+v~b2v~u~+v~δ2(u~+v~),t>0,0<x<h0,u~x(t,0)=v~x(t,0)=0,t>0,u~(t,h0)=0,t>0,u~(0,x)=u0(x),0<x<h0,v~(0,x)=maxκ2,v0L([0,+)),0<x<h0,v~x(t,h0)=maxκ2,v0L([0,+)),t>0.(30) It follows from the comparison principle that u(T1,x)u~(T1,x)for all (t,x)[0,+)×[0,h0]. Clearly, u~(T1,x) is independent of μ. Now, we consider the following system. (31) u_td1u_xx=δ1u_[κ1κ2u_],t>T1,0<x<h_(t),u_x(t,0)=0=u_(t,h_(t)),t>T1,h_(t)=μu_x(t,h_(t)),t>T1,u_(T1,x)=u~(T1,x),x[0,h0],h_(T1)=h0.(31) By Lemma 3.2, we know that h_(t)h(t) for t>T1. It follows from [Citation10, Lemma 3.7] that h_=+ if μμ¯, where μ¯:=max1,u~(T1,x)κ1κ2d1(h0h0)0h0u~(T1,x)dx. This implies that h=+.

By Theorems 3.3 and 3.4, we can also derive spreading criteria in terms of the diffusion coefficient d1, for any fixed h0.

Theorem 3.5

Spreading criteria

Let d1=4δ1(κ1κ2)h02/π2, where h0 is any prefixed positive constant. Then, spreading occurs provided that either

  1. 0<d1d1 or

  2. d1>d1 and μμ¯.

Our next result is a criterion on ‘vanishing’.

Theorem 3.6

Assume that h0<π2d1b1=π2d1δ1κ1<h0. Then, there exists μ_>0 such that h<+, whenever μμ_.

Proof.

Consider the following problem (32) u¯td1u¯xx=u¯(b1δ1u¯),t>0,0<x<h¯(t),u¯x(t,0)=0,u¯(t,h¯(t))=0,t>0,h¯(t)=μux(t,h¯(t)),t>0,u(0,x)=u0(x),h¯(0)=h0,x[0,h0].(32) Lemma 3.2 applies and yields that h(t)h¯(t)andu(t,x)u¯(t,x)for t>0and0xh(t). Furthermore, by Lemma 3.8 of [Citation10], there exists μ_>0 such that h¯<+ in the case μμ_, where μ_=δ~γ~h024M~,γ~=12π22d1h02b1, and δ~, M~ are such that π22d1(1+δ~)2h02b1=γ~ and u0(x)M~cosπ2xh0(1+δ~/2),for x[0,h0]. Therefore, h<+.

3.4. The spreading speed

If spreading occurs, it is important to estimate the spreading speed of h(t). Following an idea in [Citation12], one can obtain a rough estimate of the spreading speed as stated in the following theorem.

Theorem 3.7

[Citation12]

Suppose that κ1>κ2 and let (u,v,h) be the solution of (Equation11). If h=+, u0(x)κ1 in [0,h0), v0(x)>0 in [0,+) and lim infx+v0(x)κ2, then lim supt+h(t)ts, where s is the minimal speed of the travelling waves to the problem related with (Equation11) in the entire space. This estimation of the spreading speed is independent of μ.

However, in the fitness benefit case, we can derive an estimate better than the one in Theorem 3.4. We first recall Proposition 5.1 of  [Citation11].

Proposition 3.1

[Citation11]

For any given constants d1>0, b1>0, δ1>0 and β[0,2b1d1), the problem (33) d1U+βU=b1Uδ1U2for x(0,),U(0)=0,(33) admits a unique positive solution U=Uβ, which depends on d1,b1,δ1,β, and satisfies Uβ(x)κ1 as x+. Moreover, U(x)>0 for x0, and for each μ>0, there exists a unique β0=β0(μ,d1,b1,δ1)(0,2b1d1) such that μUβ0(0)=β0.

Our result reads:

Theorem 3.8

Assume κ1>κ2. If h=+, then β0(μ,κ1κ2,d1)lim inft+h(t)tlim supt+h(t)tβ0(μ,b1,δ1,d1), where β0 is determined by Proposition 3.1.

Proof.

Note that (34) utd1uxx=u[b1δ1(u+v)]u(b1δ1u),t>0,0<x<h(t),ux(t,0)=0,u(t,h(t))=0,t>0,h(t)=μux(t,h(t)),t>0,u(0,x)=u0(x),x[0,h0].(34) Thus, the pair (u,h) is a subsolution to the problem (35) u¯td1u¯xx=u¯(b1δ1u¯),t>0,0<x<h¯(t),u¯x(t,0)=0,u¯(t,h¯(t))=0,t>0,h¯(t)=μu¯x(t,h¯(t)),t>0,u¯(0,x)=u0(x),h¯(0)=h0,x[0,h0].(35) By the comparison principle, h(t)h¯(t) for t>0. Theorem 4.2 of [Citation10] yields that limt+h¯(t)t=β0(μ,b1,δ1,d1). Hence lim supt+h(t)tβ0(μ,b1,δ1,d1). Note that lim supt+v(t,x)κ2 uniformly for x[0,+) and h=+. Then, there exists Tε>0 such that v(t,x)κ2+ε and h(Tε)>π2d1κ1κ2εwhen t>Tε. Next, we consider the following problem (36) u_td12u_x2=u_(κ1κ2εu_),t>Tε,0<x<h_(t),u_x(t,0)=0,u_(t,h_(t))=0,t>Tε,h_(t)=μu_x(t,h_(t)),t>Tε,u_(Tε,x)=u(Tε,x),x[0,h(Tε)].(36) By the comparison principle, we obtain h(t)h_(t) for t>Tε. From Theorem 3.3, we know that h_()=+. Using a similar argument as above, we have limt+(h_(t)/t)=β0(μ,κ1κ2,d1). Therefore, lim inft+h(t)tβ0(μ,κ1κ2,d1).

4. The free boundary problem with a heterogeneous birth rate

In this section, we consider the free boundary problem (Equation6)–(Equation7) with the heterogeneous birth rates b1(x) and b2(x).

4.1. Some useful lemmas

In this subsection, we first study a related eigenvalue problem: (37) dφxx+b(x)φ+λφ=0,x(0,h0),φx(0)=φ(h0)=0.(37) Problem (Equation37) admits a positive principal eigenvalue λ1 determined by (38) λ1=infφW1,2((0,h0))0h0[dφx2b(x)φ2]dx, φx(0)=φ(h0)=0,0h0φ2dx=1.(38) We state two hypotheses that we refer to when needed. We use a generic symbol B(x) in the statement of the hypotheses. The function B(x) will be replaced accordingly (by b, b1 or b2) in the rest of this Section. B(x)C1[0,+)L[0,+)and B(x) is positive somewhere in (0,h0).(B2) B(x)C1[0,+) and b_<B(x)<b¯ for all x[0,+),where b_ and b¯ are two positive constants.(B3)

Remark 4.0.1

In order to compare the principal eigenvalues λ1 associated with different parameters, we denote the principal eigenvalue λ1 by λ1(d,h0). When we fix h0 and study the property of λ1 as d varies, we write λ1=λ1(d). Similarly, we write λ1=λ1(h0) when d is fixed while h0 varies.

We gather the following known results about the dependance of λ1 on d and h.

Lemma 4.1

[Citation40]

Suppose that b(x) satisfies (B2), where B(x) is replaced by b(x). Then, λ1=λ1(d) has the following properties:

  1. λ1(d) is increasing with respect to d.

  2. λ1(d)+ as d+ and λ1(d)maxx[0,l]b(x)<0 as d0.

  3. For any fixed h0>0, there exists d=d>0 such that

    • λ1(d)<0 for 0<d<d,

    • λ1(d)>0 for d>d, and

    • λ1(d)=0 for d=d.

Lemma 4.2

[Citation40]

Assume that (B3) holds, where B(x) is replaced by b(x). Then, λ1=λ1(h0) has the following properties:

  1. λ1(h0) is monotone decreasing with respect to h0.

  2. λ1(h0)+ as h00 and limh0+λ1(h0)<0.

  3. For any fixed d>0, there exists h0=h0>0 such that

    • λ1(h0)>0 for 0<h0<h0,

    • λ1(h0)<0 for h0>h0,

    • λ1(h0)=0 for h0=h0.

For the reader's convenience, we also recall some facts related to the following problem (39) vt=d2vxx+vb2(x)δ2v,t>0,x>0,vx(t,0)=0,t>0,v(0,x)=v0(x),x[0,+).(39) The proof of the next lemma follows from Lemma 5.2 and Lemma 6.2 of [Citation40].

Lemma 4.3

Assume that b2(x) satisfies (B3), where B(x) is replaced by b2(x). Let v(t,x) be the unique solution of (Equation39) with an initial condition v0C2[0,)L[0,)andv0>0. Then, limt+v(t,)=φv uniformly in any compact subset of [0,), where φv is the unique positive solution of the following elliptic problem (40) d2vxx+vb2(x)δ2v)=0,x>0,vx(0)=0.(40)

4.2. Sharp criteria for spreading and vanishing

Let us first consider the vanishing case.

Theorem 4.1

Let (u,v,h) be the solution of system (Equation6) subject to initial conditions (Equation7). If h<+ and b2 satisfies (B3), where we replace B(x) by b2(x), then limt+u(t,)C[0,h(t)]=0andlimt+v(t,x)=φv uniformly in any bounded subset of [0,+).

The proof is similar to that of Theorem 3.1, above.

In order to obtain sharp criteria for spreading, we require stronger conditions on b1(x) and δ1. Namely, we assume that (41) b1(x)δ1φv is positive somewhere in [0,h0].(41) Our assumption (Equation41) is not excessive in the sense that, when bi and δi(i=1,2) are constant, we have φv=b2/δ2. Consequently, b1δ1φv is a positive constant over the interval [0,+).

Theorem 4.2

Assume that b1(x)δ1φv(x) satisfies (B2) and b2(x) satisfies (B3) (where B is replaced accordingly). If 0<d1<d1, then spreading occurs.

Proof.

First, we consider the following equation: (42) v¯t(t,x)=d2v¯xx+v¯(b2(x)δ2v¯),t>0,x>0,v¯x(t,0)=0,t>0,v¯(0,x)=v0(x).(42) Since b2(x) satisfies the hypotheses of Lemma 4.3, all solutions of (Equation42) with non-trivial non-negative initial values converge to φv as t.

It follows, from the comparison principle, that vv¯ for all t>0 and x>0. Since limt+v¯(t,x)=φv uniformly in any compact subset of [0,), then, for any ε>0, there exists T>0 such that v(t,x)φv+ε, for tT.

Consider the following eigenvalue problem: (43) d1ϕxx+ϕ(b1(x)δ1(φv+ε))+λϕ=0,x(0,h0),ϕx(0)=ϕ(h0)=0.(43) It is well known that the principal eigenvalue λ1 can be characterized by λ1=infϕH1(0,h0)0h0d1ϕx2(b1(x)δ1(φv+ε))ϕ2,0h0ϕ2=1. Using (iii) of Lemma 4.1, for any fixed h0, there exists d1 such that λ1(d1)<0for all 0<d1<d1,λ1(d1)=0for d1=d1,andλ1(d1)>0for d1>d1. In this theorem, we have 0<d1<d1. Let us set u_=δϕ1(x), for tT and x[0,h0] (here ϕ1(x) is the corresponding eigenfunction of λ1). Choose δ>0, small enough, so that δϕ1(x)minλ1δ1, u(T,x)for x[0,h0]. A straightforward calculation leads to (44) u_td1u_xxu_(b1(x)δ1(φv+ε)δ1u_)=δϕ1(x)(λ1+δ1δϕ1(x))0for t>T,0<x<h0,u_x(t,0)=0,t>T,u_(t,h0)=0,t>T,u_(0,x)=δϕ1u(T,x),0xh0.(44) By the comparison principle, we have uu_, for tT and x[0,h0]. Thus, lim inftu(t,)C[0,h0]δϕ1(0)>0. By Theorem 4.1, we have h=+. Therefore, spreading occurs.

Theorem 4.3

Suppose that b1(x)δ1φv(x) satisfies (B3) and b2(x) satisfies the hypotheses of Lemma 4.3. If h0>h, then h=+ (i.e. the species u spreads eventually).

Proof.

Similarly, we consider the following equation (45) v¯t=d2v¯xx+v¯(b2(x)δ2v¯),t>0,x>0,v¯x(t,0)=0,t>0,v¯(0,x)=v0(x).(45) Since b2(x) satisfies the hypotheses of Lemma 4.3, all solutions of (Equation45) with nontrivial and nonnegative initial conditions converge to φv as t.

It follows from the comparison principle that vv¯ for t>0, x>0. Since lim supt+v¯(t,x)=φv uniformly in any compact subset of [0,). So for any ε>0, there exists T>0 such that v(t,x)φv+ε for tT.

Consider the following eigenvalue problem: (46) d1ϕxx+ϕ(b1(x)δ1(φv+ε))+λϕ=0,x(0,h0),ϕx(0)=ϕ(h0)=0.(46) The principal eigenvalue λ1 is characterized by λ1=infϕH1(0,h0)0h0d1ϕx2(b1(x)δ1(φv+ε))ϕ2,0h0ϕ2=1. Since b1(x)δ1φv satisfies the hypotheses of (B3). Then by Lemma 4.2, for any fixed d1, there exists h such that λ1(h0)<0 for all h0>h, λ1(h0)=0 for h0=h, and λ1(h0)>0 for h0<h.

If h0>h, then we set u_=δϕ1(x), for tT, x[0,h0] (here ϕ1(x) is the corresponding eigenfunction of λ1). Choose δ>0 small enough so that δϕ1(x)min{λ1/δ1,u(T,x)} for x[0,h0]. After a straightforward calculation, we obtain (47) u_td12u_x2u_(b1(x)δ1(φv+ε)δ1u_)=δϕ1(x)(λ1+δ1δϕ1(x))0,t>T,0<x<h0,xu_(t,0)=0,t>T,u_(t,h0)=0,t>T,u_(0,x)=δϕ1u(T,x),0xh0.(47) By the comparison principle, we have uu_ for tT, x[0,h0]. Hence, lim inftu(t,)C[0,h0]δϕ1(0)>0. Similarly, we have h=+; hence, spreading occurs.

Theorem 4.4

If d1>d1 and u0 is small enough, then ‘vanishing’ occurs.

Proof.

We consider the following problem as an auxiliary to the first equation of (Equation6): (48) ut=d1uxx+u(b1(x)δ1u),t>0,0<x<h0,ux(t,0)=u(t,h0)=0,t>0,u(0,x)=u0(x),x[0,h0].(48) Denote the principal eigenvalue λ1 and the corresponding positive eigenfunction φ1 satisfy (49) d1φxx+φb1(x)+λφ=0,0<x<h0,φx(0)=φ(h0)=0.(49) One can verify that there exists d1 such that λ1>0, when d1>d1. Furthermore, it follows, from Theorem 4.2 in [Citation40], that there exists a constant B such that φ1(x)2h0Bφ1(x) for all x[0,h0]. Now, we can use the following auxiliary functions, which were constructed in [Citation40]. Let h¯(t)=h01+αα2eαt,for t0andu¯(t,x)=βeαtφ1xh0h¯(t),for t0and0xh¯(t). The conditions on α and β will be determined later. If we let 0<α1, direct calculations show that h02h¯2(t)b1xh0h¯(t)b1(x)h02h¯2(t)b1xh0h¯(t)b1(x)+h02h¯2(t)1b1(x)b1xh0h¯(t)b1(x)+b1C([0,2h0])h02h¯2(t)12h0b1C1([0,2h0])+b1C([0,2h0])h0h¯(t)1. Since h¯(t)h0 as α0, we can find sufficiently small α1, such that h02h¯2(t)b1xh0h¯(t)b1(x)λ14for αα1. Moreover, there exists α2>0, small enough, such that 2h02Bα14λ1and1(1+α)234,for αα2. Let α=min{1,λ1/4,α1,α2}. Direct calculation leads to u¯td1u¯xxb1(x)u¯=αu¯βeαtφ1xh0h¯xh0h¯(t)h¯2(t)βeαtd1φ1xh0h¯(t)h02h¯2(t)b1(x)u¯=αu¯βeαtφ1xh0h¯xh0h¯(t)h¯2(t)+h02h¯2(t)b1xh0h¯(t)b1(x)u¯+h02h¯2(t)λ1u¯αu¯2h02Bα2u¯λ1u¯4+λ1u¯(1+α)2u¯λ14+λ14+λ14+3λ14=0. Furthermore, we choose 0<βh0α2/2μφ1(h0). Then, μu¯x(t,h¯(t))=βμeαtφ1(h0)h0h¯(t)βμeαtφ1(h0)h0α22eαt=h¯(t). In order to apply the comparison principle, we choose u0 small enough such that u0(x)βφ1x1+α2,for x[0,h0]. Thus, we have (50) u¯td1u¯xxu¯(b1(x)δ1u¯)0,t>0,0<x<h¯(t),u¯x(t,0)=u¯(t,h(t))=0,t>0,h¯(t)μu¯x(t,h¯(t)),t>0,u¯(0,x)=βφ1x1+α2u0(x),x[0,h0],h¯(0)=h01+α2>h0.(50) Form the comparison principle, we have h(t)h¯(t) for t>0 and u(t,x)u¯(x,t)for t>0andx[0,h(t)]. So, hlimt+h¯(t)=h0(1+α)<+. This implies that vanishing occurs.

Moreover, we can derive vanishing criteria in terms of the coefficient μ when d1>d1.

Theorem 4.5

Suppose that d1>d1. For any given u0, there exists μ depending on u0 and h0, such that vanishing occurs whenever μμ.

Proof.

As in the proof of the Theorem 4.4, let λ1 and φ1 satisfy Equation (Equation49). We still define u¯, h¯(t) as follows u¯(t,x)=β1eαtφ1xh0h¯(t),for t0,0xh¯(t).h¯(t)=h01+αα2eαt,for t0. Here, we also let α=min{1,14λ1,α1,α2} and choose β1>0 large enough such that u0(x)β1φ1x1+α2,forx[0,h0]. For this fixed β1, we choose 0<μh0α22β1φ1(h0)=:μ such that μu¯x(t,h¯(t))=β1μeαtφ1(h0)h0h¯(t)β1μeαtφ1(h0)h0α22eαt=h¯(t). Then, we have (51) u¯td1u¯xxu¯(b1(x)δ1u¯)0,t>0,0<x<h¯(t),u¯x(t,0)=u¯(t,h(t))=0,t>0,h¯(t)μu¯x(t,h¯(t)),t>0,u¯(0,x)=β1φ1x1+α2u0(x),x[0,h0],h¯(0)=h01+α2>h0.(51) Form the comparison principle, we have h(t)h¯(t), for t>0, and u(t,x)u¯(x,t),for t>0andx[0,h(t)]. Thus, hlimt+h¯(t)=h0(1+α)<+. This implies that vanishing occurs.

Next, we will prove the following conclusions.

Theorem 4.6

Assume that b1(x) satisfies (B3), where B(x) is replaced by b1(x). If hh, then the species u vanishes eventually.

Proof.

Choose l[h,h]. Consider the following equation: (52) u¯td1u¯xx+u¯(b1(x)δ1u¯)=0,t>0,0<x<l,u¯x(t,0)=u¯(t,l)=0,t>0,u¯(0,x)=u0(x),x[0,h0],u¯(0,x)=0,x[h0,l].(52) It follows from the comparison principle that 0uu¯ for t>0 and x(0,l). Since lπ/2d1/maxx[0,+)b1(x)=:h, Proposition 3.1 of [Citation4] yields that limt+u¯(t,)C[0,l]=0. Consequently, limt+u(t,)C[0,l]=0.

Under some assumptions, stated below, we can obtain the asymptotic spreading speed from Theorem 3.6 of [Citation9].

Theorem 4.7

Assume that b1(x) satisfies (B3), where B(x) is replaced by b1(x). If h=+, then lim supt+h(t)tβ0μ,maxx[0,+)b1(x),δ1,d1. Furthermore, if b1(x)δ1φv satisfies (B3), then lim inft+h(t)tβ0μ,minx[0,+)(b1(x)δ1φv), δ1,d1.

5. Summary and conclusions

We studied a reaction-diffusion model with a free boundary in one-dimensional environment. The model is developed to better understand the dynamics of Wolbachia infection under the assumptions supported by recent experiments such as perfect maternal transmission and complete CI.

In the special case of constant birth rates, we only considered the fitness benefit case. For the fitness benefit case, where the environment is more favourable for infected mosquitoes, our results show that the spreading of Wolbachia infection occurs if either the size of the initial habitat of infected population h0 is large enough, say h0h0 (Theorem 3.3), or the boundary moving coefficient μ is sufficiently large (μμ¯) in case of h0<h0 (Theorem 3.4). A rough estimate on the spreading speed of h(t) is also provided. Moreover, if h0<π/2d1/b1<h0 and μμ_, then the infection cannot spread and h<+.

The case of inhomogeneous (spatially dependent) birth rates is treated in Section 4. Detailed criteria for spreading and vanishing are derived in Subsection 4.2 with the aid of spectral properties of relevant eigenvalue problems.

Acknowledgments

The authors wish to thank the anonymous reviewers for their valuable comments and suggestions, which greatly helped us improve the presentation of this work. Y. Liu and Z. Guo were supported by National Science Foundation of China (No. 11371107, 11771104), Program for Chang Jiang Scholars and Innovative Research Team in University (IRT-16R16). Y. Liu was supported by the National Natural Science Foundation of China under Grant No.11271093 and the Innovation Research for the Postgraduates of Guangzhou University under Grant No.2017GDJC-D05. M. El Smaily and L. Wang acknowledge partial support received through NSERC-Discovery grants from the Natural Sciences and Engineering Research Council of Canada (NSERC).

Disclosure statement

No potential conflict of interest was reported by the author(s).

Additional information

Funding

The authors wish to thank the anonymous reviewers for their valuable comments and suggestions, which greatly helped us improve the presentation of this work. Y. Liu and Z. Guo were supported by National Science Foundation of China (No. 11371107, 11771104), Program for Chang Jiang Scholars and Innovative Research Team in University (IRT-16R16). Y. Liu was supported by the National Natural Science Foundation of China under Grant No.11271093 and the Innovation Research for the Postgraduates of Guangzhou University under Grant No.2017GDJC-D05. M. El Smaily and L. Wang acknowledge partial support received through NSERC-Discovery grants from the Natural Sciences and Engineering Research Council of Canada (NSERC).

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