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Research Article

Dynamic analysis of stochastic delay mutualistic system of leaf-cutter ants with stage structure and their fungus garden

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Pages 565-584 | Received 22 Jun 2021, Accepted 17 Mar 2022, Published online: 18 Jul 2022

Abstract

In this paper, we propose a stochastic delay mutualistic model of leaf-cutter ants with stage structure and their fungus garden, in which we explore how the discrete delay and white noise affect the dynamic of the population system. The existence and uniqueness of global positive solution are proved, and the asymptotic behaviours of the stochastic model around the positive equilibrium point of the deterministic model are also investigated. Furthermore, the sufficient conditions for the persistence of the population are established. Finally, some numerical simulations are performed to show the effect of random environmental fluctuation on the model.

1. Introduction

Mutualism is a universal phenomenon in the ecosystem, which refers to the cooperative survival relationship among species. For example, euryhaline tilapia co-exist and evolve with chlorella, the similiar relationship also happens to pollinating fine moths and Euphorbiacecae plants, legumes and rhizobium, and so on. Sometimes, predation behaviours also lead to reciprocal effect. Leaf mites can increase cucumber yield by consuming some cucumber seedlings. These interesting phenomena have attracted the attention of scholars. In 1925, Lotka and Volterra [Citation16, Citation20] have successively established a mathematical model of the interaction between biological populations, which was known as Lotka–Volterra model. After that, many scholars have studied the dynamic behaviour of Lotka–Voterra cooperative model. For example, Yao et al. [Citation24] added impulse and harvest term on the basis of previous studies, and generalized the research results of Lotka–Volterra cooperation model by coincidence degree theory and extension theorem of inequality analysis in 2006. Then in 2017, Feng et al. [Citation8] supplemented delay and impulse to Lotka–Volterra cooperation model, by constructing appropriate Lyapunov functions, sufficient conditions for the existence and uniform asymptotic stability of periodic solutions of the system are obtained. It is worth mentioning that there are also plenty of models based on reality that are different from Lotka–Volterra model, for instance, Kang et al. [Citation12] established a model of leaf-cutter ants and their fungus garden, which introduces in detail about how they cooperate with each other. The model is as follows: (1) {dAdt=(raFdaA)A,dFdt=(rfaA2b+aA2dfFrcA)F,(1) where A(t) is the total biomass of ants including workers, larvae, pupae and eggs at time t, F(t) is the total biomass of the fungus at time t, rc=cra and a=p2q(1p) can be considered as a parameter measuring the division of labour in the colony of ants. The meaning of other parameters is shown in Table , all parameters are positive constants.

Table 1. The meaning of some parameters.

The mutualism relationship between leaf-cutter ants and their fungus garden that we intend to study in this paper is very fascinating. The ant is very good at cutting leaves with its tail, hence it was named the ‘leaf-cutter ant’. In the Amazon rainforest, they carry leaves back to their nest not for food, but to plant fungus. To supply a place with good environment for fungus' growth, the leaves will be cut into smaller pieces and stack together after they are moved into the nest, then, under the meticulous care of the leaf-cutter ants, the mycelia quickly grows into a fungus garden. In fact, the relationship between the leaf-cutter ants and their fungus garden just like human plant mushroom and eat them, except that they don't sell the fungus to other leaf-cutter ant tribes. After the leaf-cutter ants have scattered their mycelia, if they do not look after the mycelia carefully, such as weeding, fertilizing and spraying, they will not produce a good harvest, exactly as humans grow crops.

The environment inside the nest is very favourable for the growth of the fungus, which also creates hotbeds for the growth of miscellaneous bacteria. Obviously, leaf-cutter ants face two difficulties during the cultivation process. One is how to ensure that the fungus is not contaminated by other bacteria. The other is how do they protect themselves from microbes in an environment surrounded by so many bacteria. In fact, leaf-cutter ant's mouth and forelimb are infested with parasites that can produce streptomycin, they can adjust the active ingredient of streptomycin automatically according to the species of harmful bacteria that have been present. The infected fungus will be picked out and threw away by the ants, if streptomycin does not kill the bacteria. The ants are surrounded by a wide range of pathogenic bacteria all the time, why the ants are not infected by microbes is that they have chosen to live symbiosis with a bacterium of the genus Pseudonocardia. The bacteria attach to the ant's body like armour, they secrete antibiotics once the ant is invaded by the pathogen, which help the ant fight off the intruding pathogen and mould. So far, scientists have isolated a variety of strains from the leaf-cutter ant and extracted the corresponding gene sequence to test whether the corresponding antibodies they produced are active in humans. Scientists hope to use these special bacteria to produce new antibiotics to help humans treat diseases more effectively.

In this paper, the functional response function is defined as the culture rate of fungus by leaf-cutter ants. In 1975, J. R. Beddington and D. L. DeAngelis proposed Beddington–DeAngelis type functional response (φ(x,y)=xa+bx+cy) [Citation3, Citation6], referred to as B–D functional response, which is the combination of Holling II functional response and ratio-dependent functional response, has the advantages of prey-dependent functional response and ratio-dependent functional response. Obviously, both of the biomass of leaf-cutter ants and fungus can affect the fungus culture rate. Leaf-cutter ants would reduce the culture rate when there are enough fungus, and on the contrary, they would increase the culture rate when there are less fungus, however, the upper limit of culture rate is related to the number of worker ants among leaf-cutter ants. To have a better explanation of the dynamic behaviour between leaf-cutter ants and their fungus garden, B-D functional response is considered in this paper.

For the sake of making the model more realistic, many scholars introduce stage structure into population model on account of the different abilities and behaviours of species at different ages [Citation1, Citation13, Citation15, Citation18, Citation21, Citation23]. Leaf-cutter ants are classified as the maturity and the immaturity, the adult leaf-cutter ants will participate in the cultivation of fungus, while the immaturity, including juvenile ants, eggs and pupae, need to be fed by adult leaf-cutter ants. Then, we get the following model with stage structure and B-D function response (2) {dx1(t)dt=r1x2(t)y(t)d1x1(t)cx1(t),dx2(t)dt=cx1(t)d2x2(t),dy(t)dt=r2ax2(t)y(t)m+m1y(t)+m2x2(t)d3y(t)r3x2(t)y(t),(2) where x1(t),x2(t) represent the biomass of immature and mature ants, respectively, y(t) represents the biomass of fungus, di(i=1,2,3) is the rate of death, r1, r2 are parameters that measure the maximum growth rate of ants and fungus, respectively, r3 indicates the collection rate of fungus collected by adult ants, a can be considered as a parameter measuring the division of labour in the colony of ants, all parameters are strictly positive.

Time delay is an important factor in the biological mathematical model [Citation2, Citation11, Citation19, Citation22]. In this paper, we introduce gestation time into the model, it is a period that the fungus begins to grow from spores. To cultivate fungus, leaf-cutter ants need to cut leaves first, after the leaves are transported back to the nest, the gardener ants will be responsible for the subsequent cultivation of the fungus garden, the leaves are cleaned and arranged in tight rows and in suitable positions by the gardener ants, and then they are decomposed by the secretions of the gardener, which are rich in enzymes and can form a mucus layer on the leaves' surface. With abundant nutrients, the mucus layer is a natural culture medium in which the gardener ants ‘seed’ a small amount of their own fungus, and then the careful care of ants makes the fungi grow. The period of the fungus from spores to starting growing is a relatively complex gestation time, which can be called ‘gestation time’, then the model (Equation2) with time delay can be described as (3) {dx1(t)dt=r1x2(t)y(t)d1x1(t)cx1(t),dx2(t)dt=cx1(t)d2x2(t),dy(t)dt=r2ax2(tτ)y(tτ)m+m1y(tτ)+m2x2(tτ)d3y(t)r3x2(t)y(t),(3) where τ represents the gestation delay.

An ecosystem is inevitably disturbed by random fluctuations such as temperature, humidity, rainfall, and sometimes even artificial factors in the wild [Citation5, Citation7, Citation10, Citation11, Citation14, Citation17, Citation25]. The sudden torrential rain will bring devastating disaster to the leaf-cutter ants, and the drought will increase the death rate of the fungus garden, hence, it is reasonable to consider the random fluctuations in the mutualistic system and it will enrich the dynamic behaviour of the population. Therefore, we consider that the mortality of leaf-cutter ants and fungus are affected by the natural environment in our model.

Let didi+σiB˙i(t),i=1,2,3,where Bi(t)(i=1,2,3) denote the independent standard Brownian motion, and σi2(i=1,2,3) denote the intensity of white noise [Citation17, Citation25]. Then, the stochastic system takes the form (4) {dx1(t)=(r1x2(t)y(t)d1x1(t)cx1(t))dt+σ1x1(t)dB1(t),dx2(t)=(cx1(t)d2x2(t))dt+σ2x2(t)dB2(t),dy(t)=[r2ax2(tτ)y(tτ)m+m1y(tτ)+m2x2(tτ)d3y(t)r3x2(t)y(t)]dt+σ3y(t)dB3(t),(4) and (5) xi(θ)=ϕi(θ),y(θ)=ψ(θ),θ[τ,0],i=1,2,(5) where ϕi(θ),(i=1,2) and ψ(θ) are nonnegative continuous function on [τ,0], ϕi(θ)>0,ψ(θ)>0,θ[τ,0],(i=1,2).

We have the following problems to be solved about this system: first, investigate whether the model has a globally unique positive solution, and how about the asymptotic behaviour of the model, then we want to know whether the sufficient conditions for the persistence of the model can be established. Naturally, we hope to analyse the influence of the environment disturbance on the model.

2. Existence and uniqueness of global positive solutions

In this section, we study the existence and uniqueness of positive solution of system, which is the basis of studying the long-time behaviour of the model. First of all, we present some known results and lemma which will be used later.

In general, consider the d-dimensional stochastic differential equation (6) dX(t)=f(X(t),t)dt+g(X(t),t)dB(t),fortt0,(6) with the initial value X(0)=x0Rd. B(t) denotes a d-dimensional standard Brownian motion defined on the complete probability space (Ω,F,{Ft}t0,P). The family of all nonnegative functions V(X,t) defined on Rd×[t0,), such that they are continuously twice differentiable in X and once in t. The differential operator L of Equation (Equation6) is defined by Mao [Citation17]. L=t+i=1dfi(X,t)Xi+12i,j=1d[gT(X,t)g(X,t)]ij2XiXj.If L acts on a function VC2,1(Rd×[t0,);R+), then LV(X,t)=Vt(X,t)+VX(X,t)f(X,t)+12trace[gT(X,t)VXX(X,t)g(X,t)],where Vt=Vt, VX=(VX1,,VXd), VXX=2VXiXj(i,j=1,2,,d). In view of Itô formula, if X(t)Rd, then dV(X,t)=LV(X,t)dt+VX(X,t)g(X,t)dB(t).

Theorem 2.1

For any given initial condition (Equation5), model (Equation4) has a unique global positive solution (x1(t),x2(t),y(t)) on tτ, and the solution will remain in R+3 with probability 1.

Proof.

Since the coefficients of system satisfy the local Lipschitz condition, then for any initial value (Equation5), there exists a unique local solution (x1(t),x2(t))y(t))R+3 on [τ,τe), where τe is the explosion time. To verify that this solution is global, we only need to prove that τe= a.s. For this purpose, let n0>0 be large enough such that every component of (ϕ1(θ),ϕ2(θ),φ(θ)),θ[τ,0] lay within the interval [1n0,n0]. For each integer n>n0, we define stopping time: τn=inf{t[τ,τe):min{x1(t),x2(t),y(t)}1normax{x1(t),x2(t),y(t)}n}.We can see that, τn is increasing as n, define ( denotes the empty set). Let τ=limnτn and ττe a.s. Hence to complete the proof, we need to show that τ=.

Define a C2-function V(x1,x2,y)=x1(t)+1lnx1(t)+x2(t)+1lnx2(t)+y(t)+1lny(t), applying the Ito^ formula [Citation17], one can compute that LV(x1,x2,y)=(11x1(t))(r1x2(t)y(t)d1x1(t)cx1(t))+(11y(t))(r2ax2(tτ)y(tτ)m+m1y(tτ)+m2x2(tτ)d3y(t)r3x2(t)y(t))+(11x2(t))(cx1(t)d2x2(t))+σ122+σ222+σ322=r1x2(t)y(t)d1x1(t)cx1(t)r1x2(t)y(t)x1(t)+d1+c+cx1(t)d2x2(t)cx1(t)x2(t)+d2+[r2ax2(tτ)y(tτ)m+m1y(tτ)+m2x2(tτ)d3y(t)r3x2(t)y(t)r2ax2(tτ)y(tτ)y(t)[m+m1y(tτ)+m2x2(tτ)]+d3+r3x2(t)]+σ122+σ222+σ322d1+c+d2+d3+(r1r3)x2(t)y(t)+(r3d2)x2(t)+r2ax2(tτ)y(tτ)m+m1y(tτ)+m2x2(tτ)+σ122+σ222+σ322.Then we have K1,K2 and K3, where K1=max{d1+c+d2+d3+σ122+σ222+σ322,2r3d2}, K2=∣r1r3, K3=r2am. (7) LVK1(1+x2(t)2)+K2x2(t)y(t)+K3x2(tτ)y(tτ).(7) Note that x2(t)2V(x1(t),x2(t),y(t)). We can get dV[K1(1+V(x1(t),x2(t),y(t)))+K2x2(t)y(t)+K3x2(tτ)y(tτ)]dt+σ1(x1(t)1)dB1(t)+σ2(x2(t)1)dB2(t)+σ3(x3(t)1)dB3(t).For any nn0 and t1[0,τ], we obtain (8) EV(x1(t1τn),x2(t1τn),y(t1τn))K4+K1E0t1τnV(x1(t),x2(t),y(t))dt,(8) where K4=V(ϕ1(θ),ϕ2(θ),ψ(θ))+K1τ+K20τx2(t)y(t)dt+K30τx2(tτ)y(tτ)dt<.Equation (Equation8) and the Gronwall inequality [Citation4, Citation17] imply that EV(x1(t1τn),x2(t1τn),y(t1τn))K4eτK1,for0t1τ,nn0.It then follows that (9) EV(x1(τnτ),x2(τnτ),y(τnτ))K4eτK1fornn0.(9) We can hence show that ττa.s. Additionally, letting n in Equation (Equation9) gives (10) EV(x1(t1),x2(t1),y(t1))K4eτK1for0t1τ.(10) For t2(τ,2τ], (11) EV(x1(t2τn),x2(t2τn),y(t2τn))K5+K1E0t2τnV(x1(t),x2(t),y(t))dt,(11) where K5=V(ϕ1(θ),ϕ2(θ),ψ(θ))+2K1τ+K2τ2τx2(t)y(t)dt+K3τ2τx2(tτ)y(tτ)dt<. Similarly, we obtain that τ2τa.s. and EV(x1(t2τn),x2(t2τn),y(t2τn))K5e2τK1.Repeating this procedure, one can show τmτ with probability one for any inter m1. Therefore, τ= a.s.

3. Asymptotic property

After introducing white noise into the deterministic model, the solution of the system will be disturbed under the influence of random fluctuations. Therefore, we will discuss the asymptotic property of stochastic system (Equation4) around positive equilibrium point of the corresponding deterministic model (Equation3).

System (Equation3) has the same equilibrium point as system (Equation2). Hence, we just need to consider the following equations: (12) {r1x2yd1x1cx1=0,cx1d2x2=0,r2ax2ym+m1y+m2x2d3yr3x2y=0.(12) We can deduce x1=d2x2c, from the second equation of (Equation12), we get y=d2(d1+c)r1c.

From the third equation of (Equation12), we have r2ayx2=(d3y+r3x2y)(m+m1y+m2x2)=r3m2yx22+r3y(m+m1y)x2+d3m2yx2+d3y(m+m1y),that is r3m2x22+[r3(m+m1y)+d3m2r2a]x2+d3(m+m1y)=0,where y=d2(d1+c)r1c, let b1=r3m2, b2=d3m2+r3m+r3m1d2(d1+c)r1cr2a, b3=d3(m+m1d2(d1+c)r1c).

Then (13) b1x22+b2x2+b3=0.(13) Then, if the condition H0:=b224b1b30,b2<0 is satisfied, we can get one or two positive solutions of (Equation13), simplicity, let's write them all as x2. We denote all the positive equilibria as E(x1,x2,y).

To facilitate the study, we define the following assumptions: (14) H1:{d1+c2σ12>0,d2c2σ22>0,d3(r2a)2σ32>0,r1r2acr3d2m1(d1+c)0.(14)

Theorem 3.1

For any given initial condition (Equation5), if hypothesis (H0) and (H1) are established, the solution (x1(t),x2(t),y(t)) of system (Equation4) has the property (15) lim supt1tE0t[(x1(θ)x1)2+(x2(θ)x2)2+(y(θ)y)2]dθqp,(15) where p=min{a1(d1+c2σ12),(d2c2a32m12σ2),a3(d3(r2a)2σ32)},and q=a1σ12x1+σ22x2+a3yσ32+a3x22m2+a3(x2y)2(m+m1y+m2x2)2, (x1,x2,y) is positive equilibrium point of model (Equation3), a1,a3 are positive constants to be determined later.

Proof.

Define the function (16) V=a1V1+V2+a3V3+a3V4+a5V5+a5V6+V7.(16) where V1=(x1(t)x1)22,V4=tτt(x2(r)x2)22m12dr,V2=(x2(t)x2)22,V5=y,V3=(y(t)y)22,V6=r2atτtx2(t)y(t)m+m1y(t)+m2x2(t)dr,V7=a2x1(t)+a4x2(t).And ai(i=1,2,3,4,5) are positive constants to be determined later.

The positive equilibrium satisfies the following properties: (17) {r1x2yd1x1cx1=0,cx1d2x2=0,r2ax2ym+m1y+m2x2d3yr2x2y=0.(17) By Ito^ formula, we obtain LV1=(x1(t)x1)(r1x2(t)y(t)d1x1(t)cx1(t))+12x12(t)σ12(d1+cσ12)(x1(t)x1)2+r1x2yy(t)+r1yx2(t)y(t)+σ12x1,LV2=(x2(t)x2)(cx1(t)d2x2(t))+12x22(t)σ22(d2c2σ22)(x2(t)x2)2+c2(x1(t)x1)2+σ22x2,LV3=(y(t)y)(r2ax2(tτ)y(tτ)m+m1y(tτ)+m2x2(tτ)d3y(t)r3x2(t)y(t))+12y2(t)σ32=[(m+m1y+m2x2)x2(tτ)y(tτ)(m+m1y(tτ)+m2x2(tτ))x2y(m+m1y(tτ)+m2x2(tτ))(m+m1y+m2x2)]r2a(y(t)y)d3(y(t)y)2r3(y(t)y)(x2(t)y(t)x2y)+12y2(t)σ32(d3(r2a)2σ32)(y(t)y)2+y2(tτ)(x2(tτ)x2)22(m+m1y(tτ)+m2x2(tτ))2+[(m+m1y+m2x2)x2y(tτ)(m+m1y(tτ)+m2x2(tτ))x2y]22(m+m1y(tτ)+m2x2(tτ))2(m+m1y+m2x2)2+r3x2yy(t)+yσ32+r3yx2(t)y(t)(d3(r2a)2σ32)(y(t)y)2+x22m12+(x2y)2(m+m1y+m2x2)2+(x2(tτ)x2)22m12+r3x2yy(t)+r3yx2(t)y(t)+yσ32,LV4=(x2(t)x2)22m12(x2(tτ)x2)22m12,LV5=r2ax2(tτ)y(tτ)m+m1y(tτ)+m2x2(tτ)d3y(t)r3x2(t)y(t),LV6=r2ax2(t)y(t)m+m1y(t)+m2x2(t)r2ax2(tτ)y(tτ)m+m1y(tτ)+m2x2(tτ),LV7=a2(r1x1(t)y(t)d1x1(t)cx1(t))+a4(cx1(t)d2x2(t)).Therefore, we have (18) dV(x1,x2,y)=LVdt+a1σ1(x1(t)x1)x1(t)dB1(t)+σ2(x2(t)x2)x2(t)dB2(t)+a3σ3(y(t)x1)y(t)dB3(t)+a3σ3y(t)dB3(t).(18) where LVa1(d1+cσ12)(x1(t)x1)2+a1r1x2yy(t)+a1r1yx2(t)y(t)+a1σ12x1(d2c2σ22)(x2(t)x2)2+c2(x1(t)x1)2+σ22x2a3(d3(r2a)2σ32)(y(t)y)2+a3x22m12+a3(x2y)2(m+m1y+m2x2)2+a3(x2(tτ)x2)22m12+a3(x2(t)x2)22m12a3(x2(tτ)x2)22m12+a3r3x2yy(t)+a3r3yx2(t)y(t)+a3yσ32+a5r2ax2(tτ)y(tτ)m+m1y(tτ)+m2x2(tτ)a5d3y(t)a5r3x2(t)y(t)+a4(cx1(t)d2x2(t))+a5r2ax2(t)y(t)m+m1y(t)+m2x2(t)a5r2ax2(tτ)y(tτ)m+m1y(tτ)+m2x2(tτ)+a2(r1x1(t)y(t)d1x1(t)cx1(t))a1(d1+c2σ12)(x1(t)x1)2(d2c2a32m12σ22)(x2(t)x2)2a3(d3(r2a)2σ32)(y(t)y)2+(a1r1x2y+a3r3x2ya5d3)y(t)+(a1r1y+a3r3y+a2r1a5r3)x2(t)y(t)+a1σ12x1+σ22x2+a3yσ32+a3x22m12+a3(x2y)2(m+m1y+m2x2)2+(a5r2ama4d2)x2(t)+(a2(d1+c)+a4c)x1(t).Choosing a1a2r1+a5r32r1y,max{a5r2acd2m1(d1+c),a5r3x2a5d3r1x2}a2a5r3r1,a3min{2(d2c2σ22)m12,a2r1+a5r32r3y},a5r2ad2m1a4a2(d1+c)c,a5=1.then details calculation process of a1a5 is given in Appendix, then we have d2c2a32m12σ220,a1r1x2y+a3r3x2ya5d30,a5r2am1a4d20,a2(d1+c)+a4c0,a1r1y+a3r3y+a2r1a5r30.We can obtain (19) LVa1(d1+c2σ12)(x1(t)x1)2(d2c2a32m12σ22)(x2(t)x2)2a3(d3(r2a)2σ32)(y(t)y)2+a1σ12x1+σ22x2+a3yσ32+a3x22m12+a3(x2y)2(m+m1y+m2x2)2.(19) Integrating both sides of (Equation18) from 0 to t and taking the expectation, we get (20) EV(t)EV(0)E0ta1(d1+c2σ12)(x1(t)x1)2dθE0t(d2c2a32m12σ22c2)(x2(t)x2)2dθE0ta3(d3(r2a)2σ32)(y(t)y)2dθ+[a1σ12x1+σ22x2+a3yσ32+a3r2ax22m2+a3r2a(x2y)2(m+m1y+m2x2)2]t.(20) Dividing both sides by t and taking the limit superior, we have (21) lim supt1tE0ta1(d1+c2σ12)(x1(t)x1)2dθ+lim supt1tE0t(d2c2a32m12σ22)(x2(t)x2)2dθ+lim supt1tE0ta3(d3(r2a)2σ32)(y(t)y)2dθa1σ12x1+σ22x2+a3yσ32+a3x22m2+a3(x2y)2(m+m1y+m2x2)2.(21) Obviously, (22) lim supt1tE0t(x1(t)x1)2+(x2(t)x2)2+(y(t)y)2dθqp,(22) where (23) p=min{a1(d1+c2σ12),(d2c2a32m12σ22),a3(d3(r2a)2σ32)},q=a1σ12x1+σ22x2+a3yσ32+a3x22m2+a3(x2y)2(m+m1y+m2x2)2.(23) We have completed the proof.

Remark 3.1

Theorem 3.1 shows that if the initial condition (Equation5) holds, the solution of system (Equation4) oscillates around the equilibrium point (x1,x2,y) of system (Equation3) under certain conditions, and the upper bound of the solution of system (Equation4) is positively correlated with the intensity of σi2,(i=1,2,3).

4. Persistence

In nature, whether the population has the conditions for persistence is an issue of great concern to us. Before discussing the persistence of stochastic system, we give the following assumption: (24) H2:μ=max{σ1,σ2,σ3}<min{x1p2q0,x2p2q0,yp2q0},a3<min{x12p2q1,x22p2q1,y2p2q1},(24) where q0=a1x1+x2+a3y, q1=x22m2+(x2y)2(m+m1y+m2x2)2.

Theorem 4.1

For any given initial condition (Equation5), if assumptions (H0), (H1) and (H2) hold, the system (Equation4) is persistent, that is (25) lim supt1tE0tx1(θ)dθ>0,lim supt1tE0tx2(θ)dθ>0,lim supt1tE0ty(θ)dθ>0.(25)

Proof.

According to (Equation15), we have (26) lim supt1tE0t(x1(t)x1)2dθqp,lim supt1tE0t(x2(t)x2)2dθqp,lim supt1tE0t(y(t)y)2dθqp.(26) As we know, x1(t)0 and x1>0, from 2x1(t)x1(x1)2(x1(t)x1)2, we yield (27) x1(t)x12(x1(t)x1)22x1.(27) By the condition of μ<x1p2q0,a3<x12p2q1, we have (28) lim supt1tE0tx1(θ)dθx12lim supt1tE0t(x1(t)x1)22x1dθx12q2px1x12μ2q02px1a3q12px1>0.(28) Similarly, when μ<min{x2p2q0,yp2q0},a3<min{x22p2q1,y2p2q1}, lim supt1tE0tx2(θ)dθ>0,lim supt1tE0ty(θ)dθ>0.This completes the proof of Theorem 4.1.

5. Numerical simulations

In this section, we will verify theory results and discuss the effect of environmental noise. We use the Milstein's method to simulate the stochastic model (Equation4) [Citation9], the numerical scheme for stochastic model (Equation4) is given by (29) {x1(n+1)=x1(n)+Δt(r1x2(n)y(n)d1x1(n)cx1(n))+σ1x1(n)Δtι(k)+σ122x1(n)(ι2(k)1)Δt,x2(n+1)=x2(n)+Δt(cx1(n)d2x2(n))+σ2x2(n)Δtη(k)+σ222x2(n)(η2(k)1)Δt,y(n+1)=y(n)+Δt(r2ax2(n+1τΔt)y(n+1τΔt)m+m1y(n+1τΔt)+m2x2(n+1τΔt)d3y(n)r3x2(n)y(n))+σ3y(n)Δtξ(k)+σ322y(n)(ξ2(k)1)Δt,(29) where ι(k),η(k),ξ(k)(k=1,2,,n) are the Gaussian random variables N(0,1), and time increment Δt>0.

We adopt the following parameter values: (30) r1=0.96,r2=0.25,r3=0.3,d1=0.2,d2=0.4,d3=0.1,m=0.1,m1=0.2,m2=0.01,a=0.45,c=0.65,τ=4.(30) We fix the initial value (x1(0),x2(0),y(0))=(2,1,3) except for the other specification, we obtain E(9.813,15.946,0.545). Next we take into account the effect of environmental noise intensities on the system by comparing the following different situations. Use matlab to plot diagrams of those cases.

  • In case 1, we take σ1=0.05,σ2=0.05,σ3=0.01, we conclude that μ=0.01<min{x1p2q0,x2p2q0,yp2q0}=min{7.299,0.738,0.52}, and a3=0.00029<min{0.00054,0.0014,0.0041}, it is obvious that the conditions of H0, H1, and H2 in Theorem 4.1 are satisfied, the system is persistence. It is easy to see that the solution of stochastic model oscillate around the equilibrium point of the deterministic model and maintain persistent in Figure , when the white noise is small. To satisfy the conditions of Theorem 4.1, we should select the value of a3 as smaller as possible, so there is no need to verify a3 in the following cases because we just take it sufficient small.

  • Then, to analyse the effect of σ1, we change σ1=0.4 and σ1=1.4 in case 2 and case 3, respectively, the value of μ=0.4<min{15.0358,24.4331,0.8348}=0.8348 in case 2, and μ=1.4>min{15.0358,24.4331,0.8348}=0.8348 in case 3, we can conclude that case 2 satisfy Theorem 4.1 but case 3 do not. After comparing Figures  and , we find out that the change of σ1 will affect the fluctuation of the whole system. More specifically, when σ1 increases appropriately, the solution of the leaf-cutter ants and the fungus remain persistent below the equilibrium point of the deterministic model, and all three populations of the stochastic model are rapidly extinct when σ1 is large enough, which indicates that a random variation of the death rate of immature leaf-cutter ants will turn the persistent population into extinction.

  • Furthermore, we chose σ2=0.2 and σ2=0.5 in case 4 and case 5 to explore the effect of  σ2 on the system (Equation4), μ=0.2<min{10.5349,17.1181,0.5849}=0.5849 can satisfy Theorem 4.1, but μ=0.5<min{8.4490,13.7297,0.4691}=0.4691 cannot satisfy Theorem 4.1. Which can be concluded by comparing Figures  and  is that the small changes of σ2 will have a great impact on the whole system, and even lead to the extinction of the fungus (see Figure  c,d).

  • At last, comparing Figures  and   that plotted by case 6 which σ3=0.4 and case 7 which σ3=0.9, we can make inquiry for the impact of σ3 on system (Equation4), μ=0.4<min{15.0358,24.4331,0.8348}=0.8348 can satisfy Theorem 4.1, but μ=0.9<min{15.0358,24.4331,0.8348}=0.8348 cannot satisfy Theorem 4.1, we can conclude that σ3 have the same impact as σ1 on the system, that is to say when σ3 increases appropriately, the solution of the leaf-cutter ants and the fungus remain persistent below the equilibrium point of the deterministic model, and all populations are rapidly extinct when σ3 is large enough, which indicates that a random variation of the death rate of immature leaf-cutter ants will turn the persistent population into extinction.

6. Conclusion

In this paper, we propose a stochastic delay mutualistic model of the leaf-cutter ants with stage structure and their fungus garden. First of all, we analyse the existence and uniqueness of the positive solution, and consider the asymptotic property of the positive solution, then we give the sufficient conditions for the persistence of the population. By exploring some numerical simulations, we conclude the impact of the impact of environmental disturbances on the system. In nature, the secretion of leaf-cutter ants contains many unknown antibodies, so it is of great scientific significance to maintain the persistence of leaf-cutter ants and their fungus. The research results of this paper provide a good solution for preventing the extinction of species. That is, to artificially reduce the impact of environmental disturbances on the system, such as watering the fungus nursery during drought.

Figure 1. The evolution of (x1(t),x2(t),y(t)) for model (Equation4) and its corresponding deterministic model (Equation3) with initial value (x1(0),x2(0),y(0))=(2,1,3): (a) time series diagram of x1(t),x2(t),y(t) in case 1: σ1=0.05,σ2=0.05,σ3=0.01 and (b) phase diagram of x1(t),x2(t),y(t) in case 1: σ1=0.04,σ2=0.03,σ3=0.02.

Figure 1. The evolution of (x1(t),x2(t),y(t)) for model (Equation4(4) {dx1(t)=(r1x2(t)y(t)−d1x1(t)−cx1(t))dt+σ1x1(t)dB1(t),dx2(t)=(cx1(t)−d2x2(t))dt+σ2x2(t)dB2(t),dy(t)=[r2ax2(t−τ)y(t−τ)m+m1y(t−τ)+m2x2(t−τ)−d3y(t)−r3x2(t)y(t)]dt+σ3y(t)dB3(t),(4) ) and its corresponding deterministic model (Equation3(3) {dx1(t)dt=r1x2(t)y(t)−d1x1(t)−cx1(t),dx2(t)dt=cx1(t)−d2x2(t),dy(t)dt=r2ax2(t−τ)y(t−τ)m+m1y(t−τ)+m2x2(t−τ)−d3y(t)−r3x2(t)y(t),(3) ) with initial value (x1(0),x2(0),y(0))=(2,1,3): (a) time series diagram of x1(t),x2(t),y(t) in case 1: σ1=0.05,σ2=0.05,σ3=0.01 and (b) phase diagram of x1(t),x2(t),y(t) in case 1: σ1=0.04,σ2=0.03,σ3=0.02.

Figure 2. The evolution of (x1(t),x2(t),y(t)) for model (Equation4) and its corresponding deterministic model (Equation3) with initial value (x1(0),x2(0),y(0))=(2,1,3): (a) time series diagram of x1(t),x2(t),y(t) in case 2:σ1=0.4,σ2=0.05,σ3=0.01; (b) phase diagram of x1(t),x2(t),y(t) in case 2:σ1=0.4,σ2=0.05,σ3=0.01; (c) time series diagram of x1(t),x2(t),y(t) in case 3:σ1=1.4,σ2=0.05,σ3=0.01; (d) phase diagram of x1(t),x2(t),y(t) in case 3:σ1=1.4,σ2=0.05,σ3=0.01.

Figure 2. The evolution of (x1(t),x2(t),y(t)) for model (Equation4(4) {dx1(t)=(r1x2(t)y(t)−d1x1(t)−cx1(t))dt+σ1x1(t)dB1(t),dx2(t)=(cx1(t)−d2x2(t))dt+σ2x2(t)dB2(t),dy(t)=[r2ax2(t−τ)y(t−τ)m+m1y(t−τ)+m2x2(t−τ)−d3y(t)−r3x2(t)y(t)]dt+σ3y(t)dB3(t),(4) ) and its corresponding deterministic model (Equation3(3) {dx1(t)dt=r1x2(t)y(t)−d1x1(t)−cx1(t),dx2(t)dt=cx1(t)−d2x2(t),dy(t)dt=r2ax2(t−τ)y(t−τ)m+m1y(t−τ)+m2x2(t−τ)−d3y(t)−r3x2(t)y(t),(3) ) with initial value (x1(0),x2(0),y(0))=(2,1,3): (a) time series diagram of x1(t),x2(t),y(t) in case 2:σ1=0.4,σ2=0.05,σ3=0.01; (b) phase diagram of x1(t),x2(t),y(t) in case 2:σ1=0.4,σ2=0.05,σ3=0.01; (c) time series diagram of x1(t),x2(t),y(t) in case 3:σ1=1.4,σ2=0.05,σ3=0.01; (d) phase diagram of x1(t),x2(t),y(t) in case 3:σ1=1.4,σ2=0.05,σ3=0.01.

Figure 3. The evolution of (x1(t),x2(t),y(t)) for model (Equation4) and its corresponding deterministic model (Equation3) with initial value (x1(0),x2(0),y(0))=(2,1,3): (a) time series diagram of x1(t),x2(t),y(t) in case 4: σ1=0.05,σ2=0.2,σ3=0.01; (b) phase diagram of x1(t),x2(t),y(t) in case 4:σ1=0.05,σ2=0.2,σ3=0.01; (c) time series diagram of x1(t),x2(t),y(t) in case 5: σ1=0.05,σ2=0.5,σ3=0.01 and (d) phase diagram of x1(t),x2(t),y(t) in case 5: σ1=0.05,σ2=0.5,σ3=0.01.

Figure 3. The evolution of (x1(t),x2(t),y(t)) for model (Equation4(4) {dx1(t)=(r1x2(t)y(t)−d1x1(t)−cx1(t))dt+σ1x1(t)dB1(t),dx2(t)=(cx1(t)−d2x2(t))dt+σ2x2(t)dB2(t),dy(t)=[r2ax2(t−τ)y(t−τ)m+m1y(t−τ)+m2x2(t−τ)−d3y(t)−r3x2(t)y(t)]dt+σ3y(t)dB3(t),(4) ) and its corresponding deterministic model (Equation3(3) {dx1(t)dt=r1x2(t)y(t)−d1x1(t)−cx1(t),dx2(t)dt=cx1(t)−d2x2(t),dy(t)dt=r2ax2(t−τ)y(t−τ)m+m1y(t−τ)+m2x2(t−τ)−d3y(t)−r3x2(t)y(t),(3) ) with initial value (x1(0),x2(0),y(0))=(2,1,3): (a) time series diagram of x1(t),x2(t),y(t) in case 4: σ1=0.05,σ2=0.2,σ3=0.01; (b) phase diagram of x1(t),x2(t),y(t) in case 4:σ1=0.05,σ2=0.2,σ3=0.01; (c) time series diagram of x1(t),x2(t),y(t) in case 5: σ1=0.05,σ2=0.5,σ3=0.01 and (d) phase diagram of x1(t),x2(t),y(t) in case 5: σ1=0.05,σ2=0.5,σ3=0.01.

Figure 4. The evolution of (x1(t),x2(t),y(t)) for model (Equation4) and its corresponding deterministic model (Equation3) with initial value (x1(0),x2(0),y(0))=(2,1,3): (a) time series diagram of x1(t),x2(t),y(t) in case 6:σ1=0.05,σ2=0.05,σ3=0.4; (b) phase diagram of x1(t),x2(t),y(t) in case 6:σ1=0.05,σ2=0.05,σ3=0.4; (c) time series diagram of x1(t),x2(t),y(t) in case 7:σ1=0.05,σ2=0.05,σ3=0.9; (d) phase diagram of x1(t),x2(t),y(t) in case 7:σ1=0.05,σ2=0.05,σ3=0.9.

Figure 4. The evolution of (x1(t),x2(t),y(t)) for model (Equation4(4) {dx1(t)=(r1x2(t)y(t)−d1x1(t)−cx1(t))dt+σ1x1(t)dB1(t),dx2(t)=(cx1(t)−d2x2(t))dt+σ2x2(t)dB2(t),dy(t)=[r2ax2(t−τ)y(t−τ)m+m1y(t−τ)+m2x2(t−τ)−d3y(t)−r3x2(t)y(t)]dt+σ3y(t)dB3(t),(4) ) and its corresponding deterministic model (Equation3(3) {dx1(t)dt=r1x2(t)y(t)−d1x1(t)−cx1(t),dx2(t)dt=cx1(t)−d2x2(t),dy(t)dt=r2ax2(t−τ)y(t−τ)m+m1y(t−τ)+m2x2(t−τ)−d3y(t)−r3x2(t)y(t),(3) ) with initial value (x1(0),x2(0),y(0))=(2,1,3): (a) time series diagram of x1(t),x2(t),y(t) in case 6:σ1=0.05,σ2=0.05,σ3=0.4; (b) phase diagram of x1(t),x2(t),y(t) in case 6:σ1=0.05,σ2=0.05,σ3=0.4; (c) time series diagram of x1(t),x2(t),y(t) in case 7:σ1=0.05,σ2=0.05,σ3=0.9; (d) phase diagram of x1(t),x2(t),y(t) in case 7:σ1=0.05,σ2=0.05,σ3=0.9.

Acknowledgments

We are grateful to the editor and reviewers for their valuable comments and suggestions that greatly improved the presentation of this paper.

Disclosure statement

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

Additional information

Funding

This work is supported by National Natural Sciences Foundation of China (11971405, 22072057).

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Appendix

If Equation (Equation19) holds, the following equations should be satisfied: (A1) d2c2a32m12σ220,(A1) (A2) a1r1x2y+a3r3x2ya5d30,(A2) (A3) a5r2am1a4d20,(A3) (A4) a2(d1+c)+a4c0,(A4) (A5) a1r1y+a3r3y+a2r1a5r30.(A5) From Equation (EquationA1), we can obtain a32(d2c2σ22)m12.

It is easy to conclude that Equation (EquationA5) holds, if we have the following equations: a2a5r3r1,a1a2r1+a5r32r1y,a3a2r1+a5r32r3y.Considering Equations (EquationA2) and (EquationA5), if we have a2r1x2a5r3x2a5d3,a2a5r3x2a5d3r1x2,then Equation (EquationA2) holds.

From Equations (EquationA4) and (EquationA5), a5r2ad2m1a4a2(d1+c)c, at last we need a2a5r2acd2m1(d1+c), combine with a2a5r3r1, so we need r3r1r2acd2m1(d1+c), that is r1r2acr3d2m1(d1+c)0.

In summary, we have a1a2r1+a5r32r1y,max{a5r2acd2m1(d1+c),a5r3x2a5d3r1x2}a2a5r3r1,a3min{2(d2c2σ22)m12,a2r1+a5r32r3y},a5r2ad2m1a4a2(d1+c)c,a5=1.