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ABSTRACT
We develop a delay differential equation model for the interactive wild and sterile mosquitoes. Different from the existing modelling studies, we assume that only those sexually active sterile mosquitoes play a role for the interactive dynamics. We consider the cases where the release amount is either constant or described by a given function of time. For the constant releases, we establish a threshold of releases to determine whether the wild mosquito suppression succeeds or fails. We study the existence and stability of the model equilibria. When the releases are described by given functions, the trivial equilibrium is no longer globally but locally uniformly asymptotically stable if the amount of releases is below the threshold whereas it is still globally uniformly asymptotically stable if the release amount is above the threshold. Numerical examples demonstrating the model dynamical features and brief discussions of our findings are also provided.
1. Introduction
In the face of outbreaks of mosquito-borne diseases, such as dengue fever, malaria and Zika, plaguing the world, traditional prevention and control programs are to eradicate or suppress wild mosquitoes, involving the use of insecticides. The spraying insecticides bas been an effective method to control these disease outbreaks in the short term in some areas. However, the method of spraying insecticide does not typically last long enough to keep the mosquito population density low enough to avoid the risk of epidemic. Regular spraying of insecticides can also lead to serious environmental problems and insecticide resistance [Citation22]. These problems lead to the urgent need of finding environmentally-safe ways to control the outbreaks of the mosquito-borne diseases. Through the continuous efforts of scientists, effective biological control methods have been developed and used. These include the genetic approaches [Citation9,Citation15,Citation21,Citation24], the Wolbachia driven mosquito control technique [Citation4,Citation25,Citation27], and the sterile insect technique (SIT) [Citation26].
Mathematical models have been developed and analysed to investigate the effects of these biological control measures. Modelling of transgenesis and paratransgenesis has helped us to gain insight into better strategies in genetically engineering mosquitoes [Citation16,Citation17,Citation19,Citation20]. Models based on difference or differential equations to study the suppression effects on the control of mosquitoes by releasing Wolbachia-infected male mosquitoes have been formulated with the earliest difference equation model, incorporating laboratory data, by Caspari and Watson [Citation7] in 1959. The work then has been followed one after another in [Citation11–14,Citation29–33] and the references therein. To investigate the impacts of utilizing SIT on mosquito control, mathematical models have also been formulated and analysed in [Citation1–3,Citation6,Citation16–18]. Several strategies of releases of sterile mosquitoes are proposed, qualitatively analysed, and compared in these studies. Different from the methods of transgenesis and paratransgenesis, releasing sterile or Wolbachia-infected male mosquitoes has a common basis in mathematical modelling process.
For these biological control measures, the major difficulty is to estimate how many among the released engineered mosquitoes are sexually active. Existing models involving sterile mosquitoes assume the entire population of sterile mosquitoes is sexually active and its population dynamics are governed by a different differential equation coupled with an equation for the dynamics of the wild mosquitoes. However, many sterile mosquitoes have a short sexual lifespan and lose their ability to mate. As a result, their presence has no influence on the dynamics of wild mosquitoes after they lose their mating ability.
We, in this paper, focus on further developing models for the interactive wild and sterile mosquitoes with the idea that only sexually active mosquitoes are considered and their death is ignored. We describe our model formulation in detail in Section 2. We consider two different cases of releases of sterile mosquitoes. Model dynamics for the sterile mosquitoes staying at a constant level and varying as a given function are investigated in Sections 3 and 4, respectively. Numerical examples are also provided to confirm our analytic results. Brief discussions are given in Section 5.
2. Model formulation
To explore the interactive dynamics for mosquitoes after sterile mosquitoes are released into the field of wild mosquitoes, the following ordinary differential equations model was introduced in [Citation6]:
(1)
(1) Here w and g are the numbers of wild and sterile mosquitoes at time t, respectively, a is the total number of offspring per wild mosquito, per unit of time,
is the fraction of mates with wild mosquitoes,
and
, i = 1, 2, are the density-independent and density-dependent death rates of wild and sterile mosquitoes, respectively, and B is the rate of releases of sterile mosquitoes.
Since mosquitoes undergo complete metamorphosis, going through four distinct stages of development during a lifetime [Citation8], inclusion of the larvae maturation to adults makes more realistic models. Instead of explicitly formulating stage-structured models, by using time delay τ for the larvae maturation of wild mosquitoes, the following delay differential equations model was studied in [Citation5]:
(2)
(2) where
is the survival rate of lava mosquitoes, and the initial conditions for the model system are
with
, and
and
both positive and continuous in
.
We note that in both of models (Equation1(1)
(1) ) and (Equation2
(2)
(2) ), the sterile mosquitoes are assumed to have their own dynamic equation. However, among the sterile mosquitoes determined by the second equation of both (Equation1
(1)
(1) ) and (Equation2
(2)
(2) ), some are sexually active, which can play a role in the interactive dynamics, but some have lost their abilities to mate. The interactive dynamics of wild and sterile mosquitoes after sterile mosquitoes are released are mostly affected only by the mating between the two types of mosquitoes. Thus, the modelling focus should be only on those sexually active sterile mosquitoes. Recently the author in [Citation28] introduced a non-autonomous mosquito population suppression model, in which it is assumed that the number of Wolbachia-infected mosquitoes released is a given positive function. It is also assumed that the released mosquitoes have no offspring and their losses can only be supplemented by new releases.
Employing the modelling ideas in [Citation28] and assuming that the death and the dynamics of the sterile mosquitoes with respect to the time variable are negligible, we use the following model equation for the wild mosquitoes
(3)
(3) where
is the number of sterile mosquitoes released. It is a non-negative valued function of
that satisfies
for t<0.
Comparing (Equation3(3)
(3) ) with both (Equation1
(1)
(1) ) and (Equation2
(2)
(2) ), we see that
is no longer a variable whose dynamics are governed by an independent equation, and Equation (Equation3
(3)
(3) ) is no longer an autonomous, but a non-autonomous or a time-varying equation. Many classical methods and techniques, such as qualitative analysis and the characteristic root method, are then not applicable.
For given , with an initial function
, and a given non-negative function
specified separately on
,
is said to be a solution of (Equation3
(3)
(3) ) through
if
satisfies (Equation3
(3)
(3) ) on
and
for
. It is easy to see that, since
for
, we have
for all
. We assume that the release starts at time t = 0. Then
for
and
denotes the number of the released sterile mosquitoes for the first time.
Before we proceed with further analysis of model dynamics, a few definitions are given below.
Definition 2.1
[Citation28] A solution of (Equation3
(3)
(3) ) is said to be
Stable if for any
and
, there exists
such that
and
for
implies
If δ is independent of
, then
is uniformly stable.
Stable from the right-side (or left-side) if for any
and
, there exists
such that
and
(or
) for
implies
Semi-stable if it is stable from the right-side (or left-side) but unstable from the left-side (or right-side).
Globally asymptotically stable if it is uniformly stable, and for any
3. Sterile mosquitoes staying at a constant level
In reality, sterile mosquitoes cannot be released continuously, but only at discrete times. We now assume that the sterile mosquitoes are released impulsively with a constant amount c at time kT, , where T>0.
The lifespan of male mosquitoes is relatively short, usually less than 10 days. While mating behaviour, one of the critical behaviours that characterize the mosquito life strategy, is the least understood and most understudied [Citation10,Citation23], it is well known that males can mate several times and their sexual lifespan is even much shorter than their ages. During the period when the sterile mosquitoes are sexually active, it is reasonable to neglect their death, and more importantly the efficacy of the releases of sterile mosquitoes is closely related to how often they are released and how the waiting time for the next release is correlated with the sexually active period of the sterile mosquitoes.
Let the sexual lifespan of sterile male mosquitoes be . If the new sterile mosquitoes are released exactly at the end of the sexual lifespan of sterile mosquitoes such that
, then the sterile mosquitoes in the field are almost kept at a constant level. In this case, we let
for all
. Then model equation (Equation3
(3)
(3) ) becomes
(4)
(4) An equilibrium of (Equation4
(4)
(4) ) satisfies the following quadratic equation in w:
(5)
(5) Define the threshold value of releases for (Equation4
(4)
(4) ) by
(6)
(6) We have the following existence results for the equilibria of Equation (Equation4
(4)
(4) ).
Lemma 3.1
Equation (Equation4(4)
(4) ) has a trivial equilibrium
and, in addition,
Two positive equilibria
(7)
(7) if and only if
.
A unique positive equilibrium
if
.
No positive equilibrium if
.
For the case when , Equation (Equation4
(4)
(4) ) exhibits more complex dynamics such as the bistable dynamics. We give relatively complete stability results below.
Theorem 3.2
Let . Then the following conclusions for (Equation4
(4)
(4) ) hold.
The origin
is uniformly asymptotically stable, and
for any
.
Positive equilibrium
is unstable.
Positive equilibrium
is uniformly asymptotically stable, and
for any
.
Proof.
(1) For any and
, it suffices to show that the solution of (Equation4
(4)
(4) ) with
for given c satisfies
(8)
(8) In fact, if (Equation8
(8)
(8) ) is not true, then there exists
such that
and
for
. Hence
. However, it follows from (Equation4
(4)
(4) ) that
which leads to
from Equation (Equation4
(4)
(4) ), a contradiction to
for
in (Equation5
(5)
(5) ). Thus (Equation8
(8)
(8) ) holds.
We next prove
(9)
(9) From (Equation8
(8)
(8) ), we have
. We now claim
. Otherwise,
, and we then can choose an infinite time sequence
such that
as
and
.
On the other hand, from (Equation4(4)
(4) ), we have
(10)
(10) Because
is bounded, it has a convergent subsequence. For convenience, we assume
. Then
. Taking the limit on both sides of (Equation10
(10)
(10) ) yields
and then
which indicates
. Hence,
. This contradicts (Equation8
(8)
(8) ), and as a result, we get
and
. Hence (Equation9
(9)
(9) ) is proved.
(2) The instability of follows directly from (1).
(3) For each , similarly as showing (Equation8
(8)
(8) ), we can show that
for
implies
(11)
(11) In fact, if (Equation11
(11)
(11) ) is not true, then there is
as the least time such that
or
. If
, then
. It follows from (Equation4
(4)
(4) ) that
and consequently
. It contradicts the fact that
for
If , then by a similar argument,
. This is a contradiction to the fact that
for
. This proves the uniform stability of
.
Finally, we show
By using an argument similar to that in the proof of (Equation11
(11)
(11) ), we can show
for
and hence the lower limit
. Let
be a monotonic time sequence such that
, as
, and
. Then we have, from (Equation4
(4)
(4) ), that
(12)
(12) Since
is bounded, we let
. Then
. Taking the limit on both sides of (Equation12
(12)
(12) ) yields
which leads to
Thus
, and further
. The proof will then be completed if we can prove
.
To this end, we let be an infinite divergent sequence along which
, as
, and
. Then from (Equation4
(4)
(4) ), we obtain (Equation10
(10)
(10) ). By using an argument similar to that in the proof above in (1), we have
This implies
, and hence the proof is complete.
If the number of sterile mosquitoes released is the same as the threshold value such that , for all
, then Equation (Equation3
(3)
(3) ) is reduced to (Equation4
(4)
(4) ) with c replaced by
where
is given in (Equation6
(6)
(6) ); that is, Equation (Equation3
(3)
(3) ) becomes
(13)
(13) with a unique positive equilibrium
The dynamics of (Equation13
(13)
(13) ) can be summarized as follows.
Theorem 3.3
For model equation (Equation13(13)
(13) ), we have the following results.
Equilibrium
is uniformly asymptotically stable and
, if
for
;
Equilibrium
is globally uniformly asymptotically stable from the right-side that
if
for
.
Proof.
(1) The proof is the same as that in Theorem 3.2 (1) and is omitted.
(2) To show the uniform stability of from the right-side, it suffices to verify that
on
for any
implies
(14)
(14) By an argument similar to that in the proof of Theorem 3.2, we can easily prove that
for
. Hence, if (Equation14
(14)
(14) ) is not true, then there is
such that
, and
for
and
. From (Equation13
(13)
(13) ), we get
which yields
, a contradiction to the non-negativity of
. This shows that (Equation14
(14)
(14) ) is true and hence
is uniformly stable from the right-side.
We then prove that , if
for
. To this end, we just need to prove
. Otherwise, if
, then there must be an increasing sequence
such that
,
, as
, and
.
From (Equation13(13)
(13) ), we have
(15)
(15) Since
is bounded, it has a convergent subsequence. For simplicity, we let
, and then
. Taking the limit on both sides of (Equation15
(15)
(15) ), we have
which leads to
, a contradiction to the fact that
for all non-negative
. This forces
and hence
. The proof is complete.
Remark 3.1
Theorem 3.3 tells us that is semi-stable, that is, stable from the right-side and unstable from the left-side. Such a phenomenon of semi-stability is due to the coincidence of unstable equilibrium
and stable equilibrium
when
.
We give an example below to demonstrate the results from Theorems 3.2 and 3.3.
Example 3.1
Given and parameters
(16)
(16) we have threshold
. With the release
, there exist two positive equilibria
and
. The origin
is uniformly asymptotically stable,
is unstable, and
is uniformly asymptotically stable. Solutions with initial values
, for
, approach
and solutions with initial values
, for
, approach the positive equilibrium
, as shown in the left figure in Figure . If the amount of releases is the same as the threshold
, there exists a unique positive equilibrium
which is globally uniformly asymptotically stable from the right-side such that
, for
,
, and
, for
,
, as shown in the right figure in Figure .
Figure 1. With parameters given in (Equation16(16)
(16) ) and
, the threshold of releases is
. When the amount of releases is c = 6.6874 less than the threshold, there exist two positive equilibria
. The origin
is uniformly asymptotically stable,
is unstable, and
is uniformly asymptotically stable. Solutions approach either
or
, depending on their initial values, as shown in the left figure. When the amount of releases is equal to the threshold
, there exists a unique positive equilibrium
which is globally uniformly asymptotically stable from the right-side such that
, for
,
, and
, for
,
, as shown in the right figure.
![Figure 1. With parameters given in (Equation16(16) a0=50,μ0=0.3,μ1=0.2,ξ1=0.1,τ=9,(16) ) and t0=0, the threshold of releases is g∗=7.4304. When the amount of releases is c = 6.6874 less than the threshold, there exist two positive equilibria E∓. The origin E0 is uniformly asymptotically stable, E− is unstable, and E+ is uniformly asymptotically stable. Solutions approach either E0 or E+, depending on their initial values, as shown in the left figure. When the amount of releases is equal to the threshold g∗, there exists a unique positive equilibrium E∗=8.3709 which is globally uniformly asymptotically stable from the right-side such that limt→∞w(t,g∗,0,ϕ)=0, for ϕ(t)<E∗, t∈[−9,0], and limt→∞w(t,g∗,0,ϕ)=E∗, for ϕ(t)>E∗, t∈[−9,0], as shown in the right figure.](/cms/asset/a8210afc-4343-4c0b-8f91-e8d31e53184a/tjbd_a_1682201_f0001_oc.jpg)
In the case of , equilibrium
is the unique equilibrium. Its dynamics are described as follows.
Theorem 3.4
Assume that . Then the origin
of (Equation13
(13)
(13) ) is globally uniformly asymptotically stable.
The proof follows directly from that of Theorem 4.2 and is omitted.
From Theorems 3.2, 3.3, and 3.4, it is easy to obtain the following useful conclusion.
Corollary 3.5
The origin of (Equation13
(13)
(13) ) is globally uniformly asymptotically stable if and only if
.
Remark 3.2
Corollary 3.5 provides a necessary and sufficient condition for the global asymptotic stability of . In comparison, the conditions for the global asymptotic stability of
given in [Citation5] (for the case when the dynamics of
is governed by a differential equation) are only sufficient but not necessary.
4. Sterile mosquitoes varying as a function of time
In this section we no longer assume and assume that
is not a constant but a non-negative given function of time for all t>0. Then (Equation3
(3)
(3) ) has no positive constant equilibrium, and we have the following results.
Theorem 4.1
Assume that there exist two positive constants such that
for all t>0. Then the trivial equilibrium
of (Equation3
(3)
(3) ) is uniformly asymptotically stable. Moreover,
for any
for
and
for any
for
. Here
are given in (Equation7
(7)
(7) ) with c replaced by
.
Proof.
We first show that is locally uniformly stable.
Choose such that
(17)
(17) and let
for each
. Then to show that
is locally uniformly stable, it is equivalent to showing
(18)
(18) whenever
.
Assume (Equation18(18)
(18) ) does not hold. Then there exists
such that
(19)
(19) Thus
, and a substitution of
into (Equation3
(3)
(3) ) yields
(20)
(20) We claim
. In fact, if
, then by (Equation3
(3)
(3) ), we have, for
,
and thus
which contradicts (Equation19
(19)
(19) ). Hence
.
Furthermore, it follows from (Equation20(20)
(20) ) that
a contradiction to (Equation17
(17)
(17) ). Therefore (Equation18
(18)
(18) ) holds and thus
is uniformly stable.
We next show that
(21)
(21) It suffices to prove that
Assume, otherwise,
. It is easy to prove
. Then we may choose a strictly monotone increasing sequence
such that
and
as
. It follows from Equation (Equation3
(3)
(3) ) that
(22)
(22) which yields, by substituting
, for all t>0, into (Equation22
(22)
(22) ),
(23)
(23) Since
is bounded, from (Equation18
(18)
(18) ), sequence
contains a convergent subsequence, such that
for the elements in the subsequence. Then, taking the limit on both sides of (Equation23
(23)
(23) ) as
, we have
which leads to
This implies
, a contradiction to
. Thus (Equation21
(21)
(21) ) holds and therefore
is locally uniformly asymptotically stable.
Finally, we show that
To this end, it suffices to show that if
for
, then
for all
. We prove this by contradiction again as follows.
Otherwise, we can choose such that
and
for
. Hence
and it follows from (Equation3
(3)
(3) ) that
which leads to
a contradiction. The proof thus is complete.
For the case of in Section 3, Equation (Equation3
(3)
(3) ) becomes (Equation4
(4)
(4) ) and has only the unique trivial equilibrium
and it is globally uniformly asymptotically stable. Now we show that the dynamics are similar to those for the case of variable releases as follows.
Theorem 4.2
Assume . Then the trivial equilibrium
of (Equation3
(3)
(3) ) is globally uniformly asymptotically stable.
Proof.
The uniform stability of can be proved by an argument similar to that in the proof of Theorem 3.2. To complete the proof, it suffices to show that for any
, we have
(24)
(24) In fact, if we set
, then we only need to prove
If the limit exists, it is easy to show that (Equation24
(24)
(24) ) holds. Now, we assume that the limit
does not exist. Then there is an increasing sequence
such that
as
and
From (Equation3
(3)
(3) ), we have
(25)
(25) Given that
and
are bounded, they all have convergent subsequences. For convenience and without loss of generality, we let
,
, and
. Then
and by taking limit on both sides of (Equation25
(25)
(25) ), we have
which leads to
, a contradiction to the non-negativity of
for all non-negative w. The proof is complete.
We provide an example below to confirm the results from Theorems 4.1 and 4.2. The parameters are the same as in Example 3.1 but the release functions are no longer constant functions.
Example 4.1
Given and parameters in (Equation16
(16)
(16) ), we have the same release threshold
. For the non-constant function
, t>0, we have
for all t>0, and the trivial equilibrium
is the unique equilibrium. It is locally, but not globally uniformly asymptotically stable. Solutions with initial values
,
, approach
. Solutions with initial values
,
, do not approach
but a different value, as shown in the left figure in Figure .
Figure 2. With parameters given in (Equation16(16)
(16) ) and
, the threshold of releases is
. When the release function is
for t>0, the origin
is the unique equilibrium and
is locally, but not globally uniformly asymptotically stable. Solutions with initial values
,
, approach
, and solutions with initial values
,
, approach a different value, as shown in the left figure. When we take
, since
, the origin
is the unique equilibrium and is globally uniformly asymptotically stable, as shown in the right figure.
![Figure 2. With parameters given in (Equation16(16) a0=50,μ0=0.3,μ1=0.2,ξ1=0.1,τ=9,(16) ) and t0=0, the threshold of releases is g∗=7.4304. When the release function is g(t)=((6t+1)/(t+1)) for t>0, the origin E0 is the unique equilibrium and E0 is locally, but not globally uniformly asymptotically stable. Solutions with initial values ϕ(t)<E−(1)=0.1017, t∈[−9,0], approach E0, and solutions with initial values ϕ(t)>E−(6)=2.8683, t∈[−9,0], approach a different value, as shown in the left figure. When we take g(t)=g∗(2+e−t), since inft∈(0,∞)g(t)=2g∗>g∗, the origin E0 is the unique equilibrium and is globally uniformly asymptotically stable, as shown in the right figure.](/cms/asset/834ffbb2-2227-4594-93ad-d85f4bb99c6e/tjbd_a_1682201_f0002_oc.jpg)
If the amount of releases is , for t>0, then
. The origin
is the unique equilibrium and is globally uniformly asymptotically stable. All solutions approach
, as shown in the right figure in Figure .
5. Concluding remarks
We formulated a new model (Equation4(4)
(4) ) for the interactive dynamics between wild and sterile mosquito populations. This model includes a time delay due to the larval stage of the mosquito life cycle. Adopting modelling methods in [Citation28], which are based on the short sexual lifespan of male mosquitoes, we ignored the dynamics of the sterile mosquito population. In place of a dynamical equation for sterile mosquitoes, we introduced a function
which is the amount of sterile mosquitoes released into the wild population. We then first assumed that the releases of sterile mosquitoes are kept as a constant so that
, and gave complete mathematical analysis for the model dynamics in Section 3. We derived a threshold value
for the releases and showed that if the release amount exceeds the threshold with
, the origin
is globally uniformly asymptotically stable; that is, all wild mosquitoes are wiped out regardless of their initial sizes. If the release amount is below the threshold with
, the origin
is locally uniformly asymptotically stable and there exist two positive equilibria
such that
is unstable and
is uniformly asymptotically stable. In this case, wild mosquito populations of high density will not be eliminated, whereas those of low density will be. Although it may rarely happen biologically, for mathematical completion, we also showed that at the critical situation where the release amount is exactly as
, for all
,
is uniformly asymptotically stable, and there exists a unique positive equilibrium which is globally uniformly asymptotically stable from the right-side.
When the releases of the sterile mosquitoes are given as non-constant functions of time, the model equation becomes a non-autonomous equation with time delay, and its mathematical analysis is much more challenging. We showed, in Section 4, that if exceeds the threshold
, the origin
is globally uniformly asymptotically stable, whereas if
and
,
is only locally but not globally uniformly asymptotically stable. Note that the release function
can be any given function of t. The model equation opens a door for modelling other ecological situations such as periodic and impulsive release functions. Further studies and new investigations are to appear in the near future.
Acknowledgments
The authors thank Professor Jim Cushing for his careful reading and instructive and useful comments and suggestions.
Disclosure statement
No potential conflict of interest was reported by the authors.
Additional information
Funding
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