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Articles

Discrete-time model for malaria transmission with constant releases of sterile mosquitoes

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Pages 225-246 | Received 11 Jul 2018, Accepted 18 Nov 2018, Published online: 29 Nov 2018

ABSTRACT

In this study, we first formulate a baseline discrete-time mathematical model for malaria transmission where the survival function of mosquitoes is of Beverton–Holt type. We then introduce sterile mosquitoes to the baseline model to explore the transmission dynamics with sterile mosquitoes. We derive formulas for the reproductive number R0 of infection and determine the existence and uniqueness of endemic fixed points as well, for the models with or without sterile mosquitoes. We then study the impact of the releases of sterile mosquitoes on the disease transmissions by investigating the effects of varying the release rates of the sterile mosquitoes. We use a numerical example to illustrate our results for all cases and finally give brief discussions of our findings.

AMS SUBJECT CLASSIFICATIONS:

1. Introduction

Mosquito-borne diseases, such as malaria, are a big concern for the public health worldwide. Malaria is a leading cause of death in many developing countries. There are 3.2 billion people, half of the world's population, living in areas at risk of malaria transmission in 106 countries and territories. In 2016, estimated 216 million cases of malaria occurred worldwide and 445,000 people died, mostly children in the African Region. About 1700 cases of malaria are diagnosed in the United States each year [Citation13, Citation23, Citation30].

Malaria is transmitted indirectly not from a human to a human but through vectors as blood-feeding mosquitoes. The effective way to prevent the transmission of such diseases is to control mosquitoes and among the biological methods in control of mosquitoes is the sterile insect technique (SIT) [Citation7], in which the natural reproductive process of the target population is disrupted. By chemical or physical methods, male mosquitoes are genetically modified to be sterile despite being sexually active. These sterile male mosquitoes are then released into the environment to mate with the wild female mosquitoes. A wild female mosquito that mates with a sterile male mosquito will either not reproduce or produce eggs that do not hatch. Repeated releases of genetically modified mosquitoes or the releases of a significantly large number of sterile mosquitoes may eventually wipe out a wild mosquito population, although it is, in practice, often more useful to consider controlling the population rather than eradicating it [Citation5, Citation8, Citation29].

Mathematical models have played an important role in providing insights into the transmission dynamics of diseases and in influencing the decision making for preventing the diseases. There are many models in the literature for the study of vector-borne diseases and models incorporating sterile mosquitoes, are also formulated for the disease transmission dynamics [Citation1, Citation9, Citation17, Citation18, Citation20, Citation28]. However, most of the models are of continuous time, based on differential equations, to take advantage of the rich theoretical foundation of the well-developed dynamical systems theory and bifurcation theory. As time steps are sufficiently small or population sizes are sufficiently large, these models are valid. The results from those continuous-time models have shown their significance in helping understand the transmission dynamics and make useful strategies for control and prevention of the disease.

On the other hand, the timescales for the dynamics of the human and mosquito populations are significantly different. Then, discrete-time models, based on difference equations, become equally important and even more appropriate [Citation3, Citation10, Citation11, Citation14]. Moreover, discrete-time models also have many strengths in the theory of population dynamics and mathematical epidemiology [Citation2, Citation10, Citation11, Citation27, Citation31].

In this paper, we first formulate a discrete-time susceptible-exposed-infective-recovered (SEIR) compartmental model for humans and a susceptible-exposed-infective (SEI) compartmental model for mosquitoes, based on the model in [Citation21], as our baseline model in Section 2. We derive a formula for the reproductive number R0 of infection and study the existence of endemic fixed points. We then incorporate the sterile mosquitoes into the baseline model such that the susceptible mosquitoes consist of wild and sterile mosquitoes in Section 3. We consider the case where the release rate of sterile mosquitoes is constant and derive a formula for the reproductive number. We investigate the impact of the releases of sterile mosquitoes on the malaria transmission based on the reproductive number and the endemic fixed points, and give a numerical example to verify our theoretical results in Section 4. Brief discussions of our findings are provided in Section 5.

2. Baseline model for malaria transmission

To study the impact of interactive wild and sterile mosquitoes on the malaria transmission, based on the discrete-time malaria model in [Citation21], we formulate a new discrete-time model as our baseline model in this section, where the survivability of mosquitoes assumes the Beverton–Holt type of nonlinearity instead of the Ricker type in [Citation21].

Since the mosquitoes lifespan is shorter than their infective period, we assume that mosquitoes are unable to recover after infection. Then we divide the mosquito population into groups of susceptible, exposed or incubating, and infective individuals, and denote their numbers, at generation n, by Sv(n), Ev(n) and Iv(n), respectively [Citation6, Citation21].

For the human population, we divide it into groups of susceptible, exposed or incubating, infective and recovered individuals. We let Sh(n) be the number of susceptible humans, Eh(n) the number of exposed or incubating humans who are infected but not infectious yet, Ih(n) the number of infective humans who are infected and also infectious, Rh(n) the number of humans who are recovered from infection but partly lose their immunity [Citation19, Citation24–26], and Nh(n)=Sh(n)+Eh(n)+Ih(n)+Rh(n), the total number of humans at time n.

Then, the model equations for humans and mosquitoes described in Figure  are given by (2.1) Sh(n+1)=Λ+(1λh(n))Sh(n)α1h+θhRh(n)α8h,Eh(n+1)=λh(n)Sh(n)α2h+(1γh)Eh(n)α3h,Ih(n+1)=γhEh(n)α4h+(1ηh)Ih(n)α5h,Rh(n+1)=ηhIh(n)α6h+(1θh)Rh(n)α7h,Sv(n+1)=bvNv(n)αbv(Nv(n))+(1λv(n))Sv(n)α1v,Ev(n+1)=λv(n)Sv(n)α2v+(1γv)Ev(n)α3v,Iv(n+1)=γvEv(n)α4v+Iv(n)α5v,(2.1) where the parameters are given in Table . We assume that the infection has no effects on the birth and that the intraspecific competition only affects the survivability of the susceptible newborns. We further assume that the survivability of the susceptible newborns, αbv(Nv(n)), has the Beverton–Holt-II type of nonlinearity as (2.2) αbv(Nv(n)):=kbv1+ηbvNv(n).(2.2)

Figure 1. Schematic diagram for the malaria transmission. The mosquito population is divided into three groups, Sv, Ev and Iv and the human population is divided into four groups, Sh, Eh, Ih and Rh. The mosquito infection rate λv(n) is given in (Equation2.3), and the human infection rate λh(n) is given in (Equation2.5).

Figure 1. Schematic diagram for the malaria transmission. The mosquito population is divided into three groups, Sv, Ev and Iv and the human population is divided into four groups, Sh, Eh, Ih and Rh. The mosquito infection rate λv(n) is given in (Equation2.3(2.3) λv(n)=βhrIh(n)Nh(n),(2.3) ), and the human infection rate λh(n) is given in (Equation2.5(2.5) λh(n)=G(L(n)),(2.5) ).

Table 1. The parameters for the malaria transmission model.

To determine the infection rates λv(n) and λh(n), we let r be the average number of bites that a single mosquito makes on all human hosts. Then the total number of bites made by susceptible mosquitoes is rSv, among which, a proportion of Ih/Nh goes to the infected hosts, causing rSv(Ih/Nh)βh=βhr(Ih/Nh)Sv new infections for mosquitoes, where βh is the transmission probability per bite to a susceptible mosquito. Thus the infection rate for mosquitoes is given by (2.3) λv(n)=βhrIh(n)Nh(n),(2.3) where we assume βhr<1.

Similarly, let r1 be the average number of bites a human host receives from all mosquitoes. Then the total number of bites received by susceptible human hosts is r1Sh, among which, a proportion of Iv/Nv is from infected mosquitoes, causing r1Sh(Iv/Nv)βv=βvr1(Iv/Nv)Sh new infections for hosts, where βv is the transmission probability per bite to a susceptible human. Here, however, we need to note that the number of infected mosquitoes can be sufficiently larger than the total number of humans, that is, Iv(n)/Nh(n)>1. Then, it may lead to the human infection rate greater than 1 even when βvr<1, and thus Sh(n+1)<0 from (Equation2.1), if we directly define λh(n) in the same way as for λv(n). Thus we consider the following factor for the infection rate for humans: (2.4) L(n):=βvr1Iv(n)Nv(n).(2.4) Moreover, the total number of bites that all mosquitoes make is the same as the total number of bites all human hosts receive. We then need the following balance constraint: rNv(n)=r1Nh(n), and this balance constraint and (Equation2.4) immediately lead to L(n)=βvrNv(n)Nh(n)Iv(n)Nv(n)=βvrIv(n)Nh(n). Since the number of infected mosquitoes can be sufficiently larger than the total number of humans, which may lead to the factor L(n) much greater than 1, we assume that the infection rate for humans is given by (2.5) λh(n)=G(L(n)),(2.5) where G is a positive function of L, satisfying the following conditions [Citation10]: (2.6) G:[0,)[0,1],G(0)=0,G(L)>0,G(L)<0,L0,limLG(L)=1.(2.6)

2.1. The reproductive number

We derive a formula for the reproductive number by investigating the local stability of the infection-free fixed point. The Jacobian matrix of system (Equation2.1) at the infection-free fixed point (Ev,Iv,Eh,Ih,Sv,Sh,Rh)=(0,0,0,0,S0v,S0h,0) with (2.7) S0v=kbvbv(1α1v)ηbv(1α1v),S0h=Λ1α1h,(2.7) has the form (2.8) J1:=F+T0C,(2.8) where F is the fertility matrix given by F=000rβhα2vs0vs0h00000G(0)rβvα2h000000, T is the transition matrix given by T=(1γv)α3v000γvα4vα5v0000(1γh)α3h000γhα4h(1ηh)α5h, and matrix C has the form C=c11000α1hθhα8h00(1θh)α7h, where c11=kbvbv1+ηbvS0vkbvbvηbvS0v(1+ηbvS0v)2+α1v<1, since c111=(1α1v)1α1vkbvbv1<0. Then, the next generation matrix of system (Equation2.1) [Citation4, Citation12, Citation15, Citation16] is given by Q=F(IT)1=00q13q140000q31q32000000, where q13:=rβhα4hα2vS0vS0h(1(1γh)α3h)(1(1ηh)α5h),q14:=rβhα2vS0vS0h(1(1ηh)α5h),q31:=G(0)rα2hβvγvα4v(1α5v)(1(1γv)α3v),q32:=G(0)rα2hβv1α5v.

The characteristic polynomial of Q is det(ρIQ)=ρ2(ρ2q13q31)=ρ2(ρ2q13q32), which leads to the eigenvalues as ρ1=0 and ρ2=q13q32. According to [Citation16], the infection-free fixed point is locally asymptotically stable if and only if the eigenvalues ρ1,2<1. Thus we define the net reproductive number of infection as R0:=ρ2=q13q32.

Hence, (2.9) R0:=G(0)rβhγhα2hα4h(1(1γh)α3h)(1(1ηh)α5h)rβvγvα2vα4v(1α5v)(1(1γv)α3v)S0vS0h,(2.9) with S0v and S0h given in (Equation2.7).

We have the following results.

Theorem 2.1

Define the reproductive number of infection, R0, in (Equation2.9) for system (Equation2.1). Then, the infection-free fixed point is locally asymptotically stable if R0<1, and is unstable if R0>1.

2.2. The endemic fixed point

We next explore the existence of endemic fixed points of system (Equation2.1).

The components of an endemic fixed point need to satisfy the following equations: (2.10a) Sh=Λ+(1λh)Shα1h+θhRhα8h,(2.10a) (2.10b) Eh=λhShα2h+(1γh)Ehα3h,(2.10b) (2.10c) Ih=γhEhα4h+(1ηh)Ihα5h,(2.10c) (2.10d) Rh=ηhIhα6h+(1θh)Rhα7h,(2.10d) (2.10e) Sv=kbvbvNv1+ηbvNv+(1λv)Svα1v,(2.10e) (2.10f) Ev=λvSvα2v+(1γv)Evα3v,(2.10f) (2.10g) Iv=γvEvα4v+Ivα5v.(2.10g)

Solving (Equation2.10a)–(Equation2.10d), we have (2.11) Eh=ω1hλhSh,Ih=ω1hω2hλhSh,Rh=ω1hω2hω3hλhSh,Sh=Λ1α1h+A1hλh,(2.11) where ω1h:=α2h1(1γh)α3h,ω2h:=γhα4h1(1ηh)α5h,ω3h:=ηhα6h1(1θh)α7h,A1h:=α1hθhα8hω1hω2hω3h.

Then we have (2.12) Nh=Λ(1+A2hλh)1α1h+A1hλh,(2.12) where A2h:=ω1h+ω1hω2h+ω1hω2hω3h.

Solving (Equation2.10e)–(Equation2.10g), we have (2.13) Sv=kbvbvNv(1(1λv)α1v)(1+ηbvNv),Ev=ω1vλvSv,Iv=ω1vω2vλvSv,(2.13) where (2.14) ω1v:=α2v1(1γv)α3v,ω2v:=γvα2vα4v(1α5v)(1(1γv)α3v).(2.14) Substituting (Equation2.11)–(Equation2.13) into (Equation2.3) and (Equation2.5), respectively, we obtain (2.15) λv=βhrIhNh=βhrω1hω2hλh1+A2hλh(2.15) and L(λh)=βvrIvNh=βvrω1vω2vα2vSv(1α1h+A1hλh)Λ(1+A2hλh)λv=βvrω1vω2vα2vSv(1α1h+A1hλh)Λ(1+A2hλh)βhrω1hω2hλh1+A2hλh=R021α1h+A1hλhG(0)(1α1h)(1+A2hλh)2Sv(λh)S0vλh. Hence, there exists an endemic fixed point if and only if there is a positive solution to equation λh=G(L(λh)), or equivalently, H(λh):=G(L(λh))λh for 0λh1.

Notice that L(0)=0 and hence G(L(0))=0 by (Equation2.6). Then H(0)=0 and H(1)<0 since G(L)>0 and limLG(L)=1. Moreover, it follows from Sv(0)=S0v and L(0)=R02/G(0) that H(0)=G(0)L(0)1=R021. Thus if R0>1, there exists a positive solution to H(λh)=0, that is, to λh=G(L(λh)). In other words, there exists an endemic fixed point if R0>1.

Theorem 2.2

For system (Equation2.1), if the reproductive number R0>1, there exits an endemic fixed point.

3. The interactive transmission model with sterile mosquitoes

Now suppose that sterile mosquitoes are released into a wild mosquito population and we let B(n) be the number of the sterile mosquitoes released at time n. Since sterile mosquitoes do not reproduce, there is no maturation process from larvae to adults for sterile mosquitoes. Hence, the number of sterile mosquitoes at n is just the number of released sterile mosquitoes, and the size of total mosquitoes is Nv(n)+B(n). We assume sterile mosquitoes are constantly released such that B(n):=b>0 is a positive constant. After the sterile mosquitoes are released, the mating interaction between the wild and sterile mosquitoes takes place. The number of susceptible offspring produced per mating is bvNv(n)Nv(n)+b. Then the model for the interactive mosquitoes and the human population can be described as (3.1) Sv(n+1)=kbv1+ηbvNv(n)Nv(n)bvNv(n)Nv(n)+b+(1λv(n))Sv(n)α1v,Ev(n+1)=λv(n)Sv(n)α2v+(1γv)Ev(n)α3v,Iv(n+1)=γvEv(n)α4v+Iv(n)α5v,Sh(n+1)=Λ+(1λh(n))Sh(n)α1h+θhRh(n)α8h,Eh(n+1)=λh(n)Sh(n)α2h+(1γh)Eh(n)α3h,Ih(n+1)=γhEh(n)α4h+(1ηh)Ih(n)α5h,Rh(n+1)=ηhIh(n)α6h+(1θh)Rh(n)α7h,(3.1) where the human part remains the same as described in system (Equation2.1).

Assume that all of the survival probabilities of mosquitoes αiv, i=1,2,3,4,5, in system (Equation3.1) are the same, denoted by αv. Then the interactive dynamics between the total wild mosquitoes and the constant sterile mosquitoes are governed by the following equation: (3.2) Nv(n+1)=kbvbv(Nv(n))2(1+ηbvNv(n))(Nv(n)+b)+αvNv(n).(3.2) It follows from [Citation22] that the results for (Equation3.2) can be stated as follows.

Lemma 3.1

Theorem 4.1 [Citation22]

For given r0=kbvbv, system (Equation3.2) has no, one or two positive fixed points if b>bc, b=bc or b<bc, respectively, where the release threshold bc is given by (3.3) bc:=(kbvbv1αv)2ηbv(1αv).(3.3) If b>bc, there exists no positive fixed point and N0vb=0 is the only fixed point, which is globally asymptotically stable. Thus the wild mosquito population goes extinct regardless of the initial population size. If b=bc, there exists a unique positive fixed point which is unstable and thus the trivial fixed point N0vb=0 is also globally asymptotically stable. If b<bc, there exist two positive fixed points (3.4) Nvb(±)=kbvbv(1+bηbv)(1αv)±(kbvbv(1+bηbv)(1αv))24bηbv(1αv)22ηbv(1αv),(3.4) where Nvb() is unstable and Nvb(+) is locally asymptotically stable with a basin of attraction (Nvb(),), while the trivial fixed point N0vb=0 is locally asymptotically stable with a basin of attraction (0,Nvb()).

Notice that if b>bc, for system (Equation3.2), there exists no positive fixed point and the trivial fixed point N0vb=0 is the only fixed point, which is globally asymptotically stable. Correspondingly, for system (Equation3.1), all wild mosquitoes eventually go extinct so that there will be no malaria transmission. Thus we only consider the case of b<bc, and we assume the initial size of the wild mosquitoes is greater than Nvb(), as shown in Figure .

Figure 2. Solutions approach Nv=0 or Nvb(+) depending on their initial values.

Figure 2. Solutions approach Nv=0 or Nvb(+) depending on their initial values.

3.1. The reproductive number and disease spread

We derive a formula for the reproductive number of infection after the sterile mosquitoes are released into the wild mosquito population with b<bc and the initial size in (Nvb(),) as follows.

Consider the following infection-free fixed point of system (Equation3.1): (Ev,Iv,Eh,Ih,Sv,Sh,Rh)=(0,0,0,0,S0vb,S0hb,0). It follows from (Equation2.7) that S0hb=S0h and from (Equation3.1) that (3.5) S0vb=kbvS0vb1+ηbvS0vbbvS0vbS0vb+b+S0vbα1v.(3.5) We then define function P(b):=Φ(S0vb(b),b) by (3.6) Φ(S0vb(b),b):=(1α1v)ηbvS0vb2+(1α1v)(1+bηbv)kbvbvS0vb+b(1α1v).(3.6) Clearly, there exists no, one or two positive roots of (Equation3.6) if b>bc, b=bc or b<bc, respectively, where bc is given in (Equation3.3). Therefore, in the case of b<bc, there exist two infection-free fixed points.

The Jacobian matrix Jb evaluated at an infection-free fixed point has the following form: Jb:=Fb+Tb0Cb, where Fb is the fertility matrix Fb=F, Tb is the transition matrix Tb=T, with F and T given in (Equation2.8), and Cb=c11b000α1hθhα8h00(1θh)α7h with c11b=kbvbvS0vb[2(1+ηbvS0vb)(b+S0vb)S0vb(1+2ηbvS0vb+bηbv)](1+ηbvS0vb)2(b+S0vb)2=2S0vb(1+2ηbvS0vb+bηbv)(1+ηbvS0vb)(b+S0vb)=2(1α1v)(1+2ηbvS0vb+bηbv)kbvbv. It follows from (Equation3.6) that S0vbΦ(S0vb(b),b)=2(1α1v)ηbvS0vb+(1α1v)(1+bηbv)kbvbv=(1α1v)(1+2ηbvS0vb+bηbv)kbvbv, which implies that, for the two positive roots S0vb()<S0vb(+), S0vb()Φ<0,S0vb(+)Φ>0; that is, (1α1v)(1+2ηbvS0vb()+bηbv)kbvbv<1,(1α1v)(1+2ηbvS0vb(+)+bηbv)kbvbv>1. Thus we have c11b>1 for S0vb(), and hence S0vb() is unstable.

On the other hand, it follows from c11b<1 for S0vb(+) that all eigenvalues of matrix C are inside the unit circle, and thus the local stability of this infection-free fixed point is determined by matrix Fb+Tb. It follows then from Section 2.1 that we can define the reproductive number of infection for system (Equation3.1) as (3.7) R0b(b):=G(0)rβhγhα2hα4h(1(1γh)α3h)(1(1ηh)α5h)rβvγvα2vα4v(1α5v)(1(1γv)α3v)S0vb(+)(b)S0h,(3.7) where S0vb(+)(b) equals Nvb(+) given in (Equation3.4).

By using b as a variable, S0vb(+)(b) is a function of b and so is R0b. When b=0, it is clear that S0vb(+)(b)=S0v and R0b=R0. Then R0b(b)=S0vb(+)(b)S0vR0, and, for 0<b<bc, 0<S0vb(+)(b)<S0v. Thus R0b<R0.

It follows from (Equation3.6) that P(b)=S0vbΦ(S0vb(b),b)S0vb(b)+(1α1v)(1+ηbvS0vb)=0, and we have (3.8) S0vb(b)=(1α1v)(1+ηbvS0vb)S0vbΦ(S0vb(b),b)<0.(3.8) Since S0vb(+)Φ>0, S0vb(+) is monotone decreasing with respect to b. Notice that function R0b is monotone increasing with respect to S0v(+). That is to say, the composed function R0b is monotone decreasing with respect to b. Hence, there is a unique threshold b¯ such that R0b(b¯)=1 and R0b(b)>1,if b<b¯,<1,if b>b¯. In fact, the threshold b¯ can be explicitly solved as follows.

Since 1=S0vb(+)(b¯)/S0vR0, we have S0vb(+)(b¯)=S0vR02. It follows from (Equation3.5) that (3.9) b¯=kbvbvS0vb(1α1v)(1+ηbvS0vb)S0vb=kbvbvS0v(1α1v)(R02+ηbvS0v)S0vR02.(3.9) Thus, the infection-free fixed point of system (Equation3.1) is locally asymptotically stable if b>b¯ and unstable if b<b¯. We also notice that if b=bc, the infection-free fixed point with the unique positive component S0vb(+) is unstable and if b>bc, there exists no such positive component S0vb(+). Thus all wild mosquitoes will be wiped out if bbc and hence b¯<bc. We summarize the results as follows.

Theorem 3.1

Assume sterile mosquitoes are released into the wild mosquito population constantly with the number of releases b>0. Define the two threshold values of releases bc and b¯ in (Equation3.3) and (Equation3.9), respectively. Then we have the following results.

  • If b>bc, there exists no positive fixed point for the interactive mosquitoes system (Equation3.2) and the only trivial fixed point N0vb=0 of system (Equation3.2) is globally asymptotically stable. Correspondingly, for system (Equation3.1), all wild mosquitoes are wiped out and there will be no infection.

  • If b=bc, the unique positive fixed point of system (Equation3.2) is unstable and the trivial fixed point N0vb=0 of system (Equation3.2) is globally asymptotically stable. Correspondingly, for system (Equation3.1), all wild mosquitoes are wiped out as well and hence there is no infection.

  • If b¯<b<bc, the sterile and wild mosquitoes coexist, but the reproductive number R0b<1 and the infection-free fixed point of system (Equation3.1) associated with the locally asymptotically stable positive fixed point Nvb(+) given in (Equation3.4) of system (Equation3.2) is locally asymptotically stable. Thus the infection will eventually go extinct.

  • If b<b¯ such that R0b>1, then the infection-free fixed point of system (Equation3.1) is unstable. The disease spreads when the initial size of wild mosquito is greater than Nvb() given in (Equation3.4).

3.2. Endemic fixed point

Similarly as in Section 2.2, we determine the existence of endemic fixed points for system (Equation3.1) as follows.

The components of an endemic fixed point for the human part are the same as those in Section 2.2 for the baseline model. The components for the wild mosquitoes at an endemic fixed point satisfy the following system: (3.10a) Svb=kbv1+ηbvNvbNvbbvNvbNvb+b+(1λv)Svbα1v,(3.10a) (3.10b) Evb=λvSvbα2v+(1γv)Evbα3v,(3.10b) (3.10c) Ivb=γvEvbα4v+Ivbα5v,(3.10c) which leads to Nvb=kbvbvNvb2(1+ηbvNvb)(b+Nvb)+αNvb with αiv=α for i=1,,5. Then Nvb=S0vb=Nvb(+) given in (Equation3.4).

Solving (Equation3.10a)–(Equation3.10c), we have (3.11) Svb=kbvbvNvb2(1+ηbvNvb)(b+Nvb)(1(1λv)α1v),Evb=λvα2v1(1γv)α3vSvb=ω1vλvSvb,Ivb=γvα4v1α5vλvα2v1(1γv)α3vSvb=ω2vλvSvb,(3.11) where ω1v and ω2v are defined in (Equation2.14). Then we obtain (3.12) Svb=Nvb1+λv(ω1v+ω2v).(3.12) Substituting (Equation3.11) and (Equation2.15) into L(t) in (Equation2.5) yields L(λh)=βvrIvbNh=βvrω1vα2vSvb(1α1h+A1hλh)Λ(1+A2hλh)λv=βvrω1vα2vSvb(1α1h+A1hλh)Λ(1+A2hλh)βhrω1hω2hλh1+A2hλh=R0b21α1h+A1hλhG(0)(1α1h)(1+A2hλh)2Svb(λh)S0vbλh. It follows from (Equation3.12) that Svb(λh)S0vb=11+λv(ω1v+ω2v). Then (3.13) L(λh)=R0b21α1h+A1hλhG(0)(1α1h)(1+A2hλh)211+λv(ω1v+ω2v)λh=R0b21α1h+A1hλhG(0)(1α1h)(1+A2hλh)(1+Dλh)λh,(3.13) where D=A2h+βhrω1hω2h(ω1v+ω2v)>0 and A2h>0. Hence, there exists an endemic fixed point if and only if there is a positive solution to equation λh=G(L(λh)), or equivalently, (3.14) Hb(λh):=G(L(λh))λh,(3.14) for 0λh1.

Notice that L(0)=0 and hence G(L(0))=0 by (Equation2.6). Then Hb(0)=0 and Hb(1)<0 since G(L)>0 and limLG(L)=1. Moreover, it follows from L(0)=R0b2/G(0) that Hb(0)=G(0)L(0)1=R0b21. Thus if R0b>1, there exists a positive solution λ to Hb(λh)=0, that is, to λ=G(L(λ)). Then there exists an endemic fixed point if R0b>1.

Next, we prove the uniqueness of this endemic fixed point.

We rewrite the equation Hb(λh)=G(L(λh))λh as Hb(x)=G(L(x))x for convenience and take the first derivative with respect to variable x. Then we have Hb(x)=G(L(x))L(x)1, where G(L(x))>0. It follows from (Equation3.13) that L(x)=R0b2G(0)(1α1h)l1(x)(1+A2hx)2(1+Dx)2, where l1(x):=kx2+2A1hx+(1α1h) and we write (3.15) k:=A1h(A2h+D)A2hD(1α1h).(3.15) If k<0, there exists a unique positive solution x satisfying l1(x)=0. Then L(x)>0 if x(0,x) and L(x)<0 which leads to Hb(x)<0 if x>x. Thus Hb(x) is monotone decreasing if x>x.

We then prove that L(x) is monotone decreasing for x<x, which is equivalent to L(x)<0 for x<x.

It follows from (Equation3.13) that L(x)=2R0b2G(0)(1α1h)l2(x)(1+A2hx)3(1+Dx)3, where (3.16) l2(x)=[kA2hD]x33A1hA2hDx23A2hD(1α1h)x+A1h(A2h+D)(1α1h).(3.16) Since A1h<A2hD(1α1h)/(A2h+D) from (Equation3.15) with k<0, the constant part of function l2(x) becomes A1h(A2h+D)(1α1h)<A2hD(1α1h)A2h+D(A2h+D)(1α1h)<1α1hA2h+D(A2hD2)23D24<0. Hence there exists at most one positive solution x0 satisfying l2(x0)=0 from Descartes' rule of sign. Thus L(x)<0 if x<x0 and L(x)>0 if x>x0.

In order to compare x with x0, we assume x>x0. Then, for any x(x0,x), L(x) is increasing and L(x)>0 but L(x)=0, which is a contradiction. Thus, x<x0, and then L(x)<0 for x<x. It follows from (3.17) Hb(x)=G(L(x))(L(x))2+G(L(x))L(x),(3.17) where G(L(x))<0, that Hb(x)<0 for x<x. Therefore, function Hb(x) is concave down for x<x and then decreasing for x>x with Hb(0)>0, which implies that the endemic fixed point is unique.

If k>0, then A1h>0 and thus L(x)>0 for all x. If the constant part of l2(x) from (Equation3.16) is negative, l2(x)<0 for all x(0,1] and thus L(x)<0, which implies that Hb(x)<0 from (Equation3.17). Since function Hb(x) is concave down for all x(0,1], the endemic fixed point is unique.

If the constant part of l2(x) from (Equation3.16) is positive, there exists a unique solution x1 such that L(x1)=0. Then L(x)>0 if x<x1 and L(x)<0 if x>x1. Hence Hb(x) is monotone increasing if x<x1 and monotone decreasing if x>x1 with Hb(0)=Rb21>0, where there exists a unique solution x¯ such that Hb(x¯)=0. Therefore, function Hb(x) is monotone increasing for x<x¯ and monotone decreasing for x>x¯. Thus the endemic fixed point is unique.

In summary, we have shown that when R0b>1, the infection-free fixed point is unstable and there exists a unique endemic fixed point.

In the following, we prove that if R0b<1, there exists no endemic fixed point.

If k<0, it follows from the analysis above that Hb(x) is concave down if x<x and monotone decreasing if x>x. Thus there exists no intersection with x-axis since Hb(0)=Rb21<0.

If k>0, it follows from (Equation3.16) that Hb(x) is concave down with Hb(0)<0 if the constant part of l2(x) is negative, and hence there exists no endemic fixed point. On the other hand, if the constant part of l2(x) is positive, Hb(x) is monotone increasing if x<x1 and monotone decreasing if x>x1 with Hb(0)=Rb21<0. Thus Hb(x1)=0 and L(x1)=0 since L(x1)=0 from (Equation3.17). Then Hb(x1)=G(L(x1))L(x1)1=1<0, which implies that Hb(x)<0 for all x(0,1]. Hence Hb(x) is monotone decreasing for all x(0,1] and there exists no endemic fixed point.

4. Impact of releases of sterile mosquitoes

To study the impact of the releases of sterile mosquitoes on the malaria transmission dynamics, first, it is clear that if bb¯, there is no infection. We then consider the interval (0,b¯), where b¯ is given in (Equation3.9). For each b(0,b¯), the corresponding reproductive number R0b>1 and there exists a unique endemic fixed point associated with λ to equation Hb(λ)=0.

With this λ(b), we have Hbλλ=λ(b)=G(λ)L(λ)1<0. Then it follows from (Equation3.13) that L(b)=L(R0b(b),λ(b)) since both R0b and λ can be regarded as functions of variable b. Thus (4.1) L(b)=L(b)R0b(b)R0b(b)+L(b)λ(b)λ(b)=2R0bλ(b)(1α1h+A1hλ(b))G(0)(1α1h)(1+A2hλ(b))(1+Dλ(b))R0b(b)+L(λ(b))λ(b),(4.1) and by taking the derivative of equation Hb(λ(b))=0 with respect to b on both sides, Hb(b)=G(L(b))L(b)λ(b)=2R0bG(λ)λ(b)(1α1h+A1hλ(b))G(0)(1α1h)(1+A2hλ(b))(1+Dλ(b))R0b(b)+(G(λ)L(λ)1)λ(b)=0. Equivalently, (4.2) λ(b)(G(λ)L(λ)1)=2R0bG(λ)λ(b)(1α1h+A1hλ(b))G(0)(1α1h)(1+A2hλ(b))(1+Dλ(b))R0b(b),(4.2) where G(λ(b))L(λ(b))1<0 and R0b<0. Hence λ(b)<0, which implies that λ is monotone decreasing with respect to b.

Moreover, it follows from (Equation2.11) that Ih(b)=Λω1hω2hλ(b)1α1h+A1hλ(b), and then Ih(b)=ω1hω2hΛ(1α1h)(1α1h+A1hλ)2λ<0. Thus component Ih(b) of the unique endemic fixed point is monotone decreasing with respect to b. In other words, we can reduce the population of the infected humans by increasing the amount of sterile mosquitoes.

Furthermore, it follows from (Equation3.11) and (Equation3.12) that Iv(b)=ω2vλv(b)Nvb+(b)1+λv(b)(ω1v+ω2v). Taking the derivative with respect to b, we obtain Iv(b)=ω2v(1+λv(b)(ω1v+ω2v))2Nvb+λv+(1+(ω1v+ω2v)λv)Nvb+(b). In addition, it follows from (Equation2.15) that λv(b)=βhrω1hω2hλ(1+A2hλ)2<0, and Nvb+(b)<0 from (Equation3.8). Consequently, Iv(b)<0, which implies that component Iv(b) of the unique endemic fixed point is also monotone decreasing with respect to b. As a result of increasing the releases of the sterile mosquitoes, we can possibly wipe out all of the wild mosquitoes eventually.

Therefore, for b(0,b¯), even though R0b(b)>1 such that the disease spreads and goes to a positive steady state for n, we can still reduce the components of the infected humans and mosquitoes by increasing the releases of the sterile mosquitoes to make the transmission under control.

We provide an example below to demonstrate our findings.

Example 4.1

We use the following parameters for the transmission model: (4.3) Λ=6;kbv=0.5;ηbv=1.5;bv=80;γv=0.8;βv=0.1;βh=0.15;α1h=0.3;α2h=0.7;α3h=0.5;α4h=0.8;α5h=0.8;α6h=0.8;α7h=0.8;α8h=0.8;γh=0.7;ηh=0.25;θh=0.5;r=6;(4.3) and α1v=α2v=α3v=α4v=α5v=0.5.

Before the sterile mosquitoes are released, the reproductive number of infection for system (Equation2.1) is R0=1.3039>1 and hence the disease spreads. After the releases of sterile mosquitoes, we have the existence threshold value bc=42.0743 such that if b=45>42.0743, there exists no positive fixed point and thus the infection eventually dies out, as shown in Figure . If b<42.0743, there exist two positive fixed points with components Nvb(±)(b)=26.33330.5000b±0.6667(39.50.75b)21.500b, where Nvb() is unstable and Nvb(+) is locally asymptotically stable with initial value larger than Nvb(). Nvb(+) is a decreasing function with respect to release value b.

Figure 3. With the parameters given in (Equation4.3), the threshold value is bc=42.0743. If b=45>bc, the trivial fixed point is globally asymptotically stable, which leads to the infection eventually extinct as shown above.

Figure 3. With the parameters given in (Equation4.3(4.3) Λ=6;kbv=0.5;ηbv=1.5;bv=80;γv=0.8;βv=0.1;βh=0.15;α1h=0.3;α2h=0.7;α3h=0.5;α4h=0.8;α5h=0.8;α6h=0.8;α7h=0.8;α8h=0.8;γh=0.7;ηh=0.25;θh=0.5;r=6;(4.3) ), the threshold value is bc=42.0743. If b=45>bc, the trivial fixed point is globally asymptotically stable, which leads to the infection eventually extinct as shown above.

However, the threshold value for the disease spread is b¯=21.2330 such that the reproductive number R0b(b)<1 if b=30>21.2330 and thus the disease dies out eventually, which is shown in Figure  (right). If b<21.2330, the reproductive number R0b(b)>1, which makes the infection-free fixed point unstable and thus the disease spreads. Moreover, R0b(b) is monotone decreasing with respect to b, as shown in Figure  (left).

Figure 4. With the same parameters in (Equation4.3), the threshold values are b¯=21.2330 and bc=42.0743, respectively. By using b as a variable, the horizontal axis is for b and the vertical axis is for R0b(b) (left). The reproductive number R0b(0)=R0=1.3039 at b=0. For b=30, the infection-free fixed point is stable since R0b(b)<1. Then the infected humans and infected mosquitoes eventually go extinct (right).

Figure 4. With the same parameters in (Equation4.3(4.3) Λ=6;kbv=0.5;ηbv=1.5;bv=80;γv=0.8;βv=0.1;βh=0.15;α1h=0.3;α2h=0.7;α3h=0.5;α4h=0.8;α5h=0.8;α6h=0.8;α7h=0.8;α8h=0.8;γh=0.7;ηh=0.25;θh=0.5;r=6;(4.3) ), the threshold values are b¯=21.2330 and bc=42.0743, respectively. By using b as a variable, the horizontal axis is for b and the vertical axis is for R0b(b) (left). The reproductive number R0b(0)=R0=1.3039 at b=0. For b=30, the infection-free fixed point is stable since R0b(b)<1. Then the infected humans and infected mosquitoes eventually go extinct (right).

For b<21.2330, R0b(b)>1, the corresponding λh(b) determined in (Equation3.14) is a positive and decreasing function as shown in Figure  (left). Corresponding to λh(b), there exists a unique endemic fixed point whose components Iv(b) and Ih(b) are monotone decreasing with respect to b as shown in Figure  (right), which indicates that increasing the releases of sterile mosquitoes reduces the disease spread.

Figure 5. With still the same parameters in (Equation4.3), the threshold values are b¯=21.2330 and bc=42.0743, respectively. The curve on the left figure is for λh(b) at the endemic fixed point for each b. The upper curve and lower curve on the right figure are for Ih(b) and Iv(b) at the endemic fixed point for each b, respectively. Clearly, λh(b), Ih(b) and Iv(b) are all negative if b>b¯, which implies that no endemic fixed point exists although positive Nvb(+) exists for b¯<b<bc.

Figure 5. With still the same parameters in (Equation4.3(4.3) Λ=6;kbv=0.5;ηbv=1.5;bv=80;γv=0.8;βv=0.1;βh=0.15;α1h=0.3;α2h=0.7;α3h=0.5;α4h=0.8;α5h=0.8;α6h=0.8;α7h=0.8;α8h=0.8;γh=0.7;ηh=0.25;θh=0.5;r=6;(4.3) ), the threshold values are b¯=21.2330 and bc=42.0743, respectively. The curve on the left figure is for λh(b) at the endemic fixed point for each b. The upper curve and lower curve on the right figure are for Ih(b) and Iv(b) at the endemic fixed point for each b, respectively. Clearly, λh(b), Ih(b) and Iv(b) are all negative if b>b¯, which implies that no endemic fixed point exists although positive Nvb(+) exists for b¯<b<bc.

To show the dynamical impact of the releases of the sterile mosquitoes, we also present the solutions of the disease transmission system in Figure . When no sterile mosquitoes are released, the reproductive number R0=1.3039>1 and the disease spreads as shown in Figure  (left). For b=15<b¯=21.2330, the reproductive number R0b=1.0988>1 such that the infection still exists but the number of infected humans and infected mosquitoes are reduced significantly as shown in Figure  (right).

Figure 6. (Left) With again the parameters given in (Equation4.3), the reproductive number is R0=1.3039>1 and hence the disease spreads in the absence of sterile mosquitoes. (Right) After the sterile mosquitoes are released, even the released amount is less than the threshold with b=15<b¯, the infected components Ih and Iv are reduced so that we can have the infection under control.

Figure 6. (Left) With again the parameters given in (Equation4.3(4.3) Λ=6;kbv=0.5;ηbv=1.5;bv=80;γv=0.8;βv=0.1;βh=0.15;α1h=0.3;α2h=0.7;α3h=0.5;α4h=0.8;α5h=0.8;α6h=0.8;α7h=0.8;α8h=0.8;γh=0.7;ηh=0.25;θh=0.5;r=6;(4.3) ), the reproductive number is R0=1.3039>1 and hence the disease spreads in the absence of sterile mosquitoes. (Right) After the sterile mosquitoes are released, even the released amount is less than the threshold with b=15<b¯, the infected components Ih and Iv are reduced so that we can have the infection under control.

5. Concluding remarks

To have a better understanding of the impact of releasing sterile mosquitoes on malaria transmission, we first formulated a simple compartmental SEIR model for the malaria transmission, the survivability of the susceptible newborns has the Beverton–Holt-II type of nonlinearity, in (Equation2.1) as our baseline model in Section 2. We derived a formula for the reproductive number of infection R0 in (Equation2.9) and showed that the infection-free fixed point of the baseline model is asymptotically stable if R0<1 and unstable if R0>1. We also showed that if R0>1, there exists an endemic fixed point for the baseline model.

We then introduced sterile mosquitoes into our baseline model and formulated the interactive model in Section 3. The human components are the same as those in (Equation2.1) but the mosquito components are given in (Equation3.1). We only consider the case of constant releases of sterile mosquitoes. We derived a formula for the reproductive number R0b, presented in (Equation3.7), and showed R0b<R0 for any release amount b. We proved that the infection-free fixed point is asymptotically stable if R0b<1 and is unstable if R0b>1. Using the rate of releases b as a variable, we determined threshold value bc for the existence of a unique endemic fixed point for the interactive wild and sterile mosquitoes and threshold value b¯ that determines whether R0b<1 or R0b>1, that is, whether the disease dies out or spreads.

We studied the impact of the releases of sterile mosquitoes on the transmission dynamics in Section 4 by investigating the variation of the reproductive number, R0b(b), and the infected components, Ih(b) and Iv(b), induced by λh(b), as b varies, respectively. Based on the threshold values bc and b¯, we provided Example 4.1 to confirm and demonstrate our findings. If the rate of releases b is greater than threshold bc, the trivial fixed point for the interactive wild and sterile mosquitoes is locally asymptotically stable. All wild mosquitoes are wiped out and thus there is no infection. On the other hand, if b<bc, while the wild mosquitoes cannot be wiped out and the two types of mosquitoes coexist, the disease can still go extinct when there are sufficient sterile mosquitoes with b¯<b<bc, which leads to R0b<1. In the case even if we are unable to release enough sterile mosquitoes with b<b¯, which leads to R0b>1 and thus the disease spreads when the initial wild mosquitoes is larger than Nvb() given in (Equation3.4), the infected components Ih(b) and Iv(b) are monotone decreasing with respect to b; that is, the infection can be reduced. All of the results are demonstrated by numerical simulations in Example 4.1.

In conclusion, based on the discrete-time interactive malaria model we formulated and the analysis we accomplished in this study, we showed that the releases of sterile mosquitoes have significant impacts on controlling the transmission of malaria. It will be useful and helpful in considering effective strategies for control and prevention of mosquito-borne diseases as well.

Disclosure statement

No potential conflict of interest was reported by the authors.

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