Ankit Kumar and Balram Dubey
Department of Mathematics,BITS Pilani,Pilani Campus,Pilani,333031,India
ABSTRACT In this paper,the impact of additional food and two discrete delays on the dynamics of a prey-predator model is investigated.The interaction between prey and predator is considered as Holling Type-II functional response.The additional food is provided to the predator to reduce its dependency on the prey.One delay is the gestation delay in predator while the other delay is the delay in supplying the additional food topredators.The positivity,boundedness and persistence of the solutions of the system are studied to show the system as biologically well-behaved.The existence of steady states,their local and global asymptotic behavior for the non-delayed system are investigated.It is shown that(i)predator’s dependency factor on additional food induces a periodic solution in the system,and(ii)the two delays considered in the system are capable to change the status of the stability behavior of the system.The existence of periodic solutions via Hopf-bifurcation is shown with respect to both the delays.Our analysis shows that both delay parameters play an important role in governing the dynamics of the system.The direction and stability of Hopf-bifurcation are also investigated through the normal form theory and the center manifold theorem.Numerical experiments are also conducted to validate the theoretical results.
KEYWORDS Additional food;Gestation delay;Hopf-bifurcation;Prey-predator
Living organisms on the surface of the earth adopt only the way that boosts their survival possibilities so that they can pass their genes to the next generation.There are several fundamental instincts in ecological communities and,predation is one of them that constitutes the building blocks for multispecies food webs.Initially,Lotka [1] and Volterra [2] studied the model for preypredator interaction and observed the uniform fluctuations in the time series of the system.On later the fluctuations were removed from the system by taking logistic growth of prey population [3,4].Many researchers have widely studied prey-predator interactions for the last century[5-12].They have considered several essential concepts over time that play a vital role in the dynamics of the system like functional response,time delay,harvesting and conservation policies of species,stage structure,fear induced by predators,etc.The idea of functional response was proposed by Holling [5].It is defined as the consumption rate of prey by predators.Holling considered it nonlinear function of prey species that saturates at a level.Further,it was considered a function of prey and predator both by several authors [6,10,13,14].
In last few decades,many authors have studied the qualitative dynamics of prey-predator systems in the presence of additional food resources for predators [15-20].Additional food is an important component for predators like coccinellid which shapes the life history of many predator species [17].Ghosh et al.[20] investigated the impact of additional food for predator on the dynamics of prey-predator model with prey refuge and they observed that predator extinction possibility in high prey refuge may be removed by providing additional food to predators.Again,to study the role of additional food in an eco-epidemiological system,a model was proposed and studied by Sahoo [21].The author found that the system becomes disease free in presence of suitable additional food provided to predator.Recently,a prey-predator model with harvesting and additional food is analyzed by Rani et al.[22] and they have shown some local and global bifurcation with respect to different parameters.To incorporate the additional food into the model,they modified the Holling type II functional response.
Delayed models exhibit much more realistic dynamics than non delayed models [4,23].In preypredator system,the impact of consumed prey individuals into predator population does not appear immediately after the predation,there is some time lag that is gestation delay [24].We incorporate the effect of time delay into the model with delay differential equations.A delay differential equation demonstrates much more complex character than ordinary differential equation.On the other hand predators do not consume the additional food as soon as it is provided.They take some time to consume and digest the food.Delayed models are widely studied by researchers [10,14,25-33].A delayed prey and predator density dependent system is investigated by Li et al.[10].The authors analyzed stability,Hopf-bifurcation and its qualititative properties by using Poincare normal form and the formulae given in Hassard et al.[34].Sahoo et al.[35]examined prey-predator model with effects of supplying additional food to predators in a gestation delay induced prey-predator system and habitat complexity.They have pointed out that Hopfbifurcation occurs in the system when delay crosses a threshold value that strongly depends on quality and quantity of supplied additional food.The effect of additional food along with fear induced by predators and gestation delay is discussed by Mondal et al.[36].There are several studies carried out with multiple delays [37-40].Li et al.[37] have done stability and Hopf-bifurcation analysis of a prey-predator model with two maturation delays.Gakkhar et al.[30] explored the complex dynamics of a prey-predator system with multiple delays.They established the presence of periodic orbits via Hopf-bifurcation with respect to both delays.Recently,Kundu et al.[40]have discussed about the dynamics of two prey and one predator system with cooperation among preys against predators incorporating three discrete delays.The authors have found that all delays are capable to destabilize the system.
To the best of our knowledge,an ecological model including (i) Effect of additional food supplies to predators,(ii) Dependency factor of supplied additional food,(iii) Holling Type II functional response,(iv) Gestation delay in predator have not been considered.Inspired by this,we establish three dimensional non delayed and delayed models in Section 2.We analyze the dynamics of non delayed model and validate it via some numerical simulations in Section 3.In Section 4,we analyze the dynamics of delayed model through Hopf-bifurcation.Direction and stability of Hopf-bifurcation are carried out in Section 5.Section 6 is devoted to the numerical simulations for delayed model.Conclusions and significance of this work are discussed briefly in Section 7.
We consider a habitat where two biological populations,prey population and predator population are surviving and interacting with each other.It is assumed that prey population grows logistically and the interaction between prey and predator follows Holling Type II functional response.We assume that the density of the additional food supplied to the predators is directly proportional to the density of predators present in the habitat.Keeping these in view,the dynamics of the system can be governed by the following system of differential equations:
In the above modelx(t),y(t)are number of prey and predator individuals at timetandAis quantity of additional food provided to predators.A0is dependency factor of predators on provided additional food resources.IfA0=1,then predators depend completely on additional food and prey population grows logistically.IfA0=0,then predators depend only on the prey population and in such a case additional food is not required.λis maximum supply rate of additional food resources.
In real situations,each organism needs an amount of time to reproduce their progeny.Due to this fact the increment in predators does not appear immediately after consuming prey.It is assumed that a predator individual takesτ1time for gestation.Therefore,it seems reasonable to incorporate a gestation delay in the system.Thus,the delayτ1is considered in the numeric response only.Again it is assumed that the additional food is provided to predators with another delayτ2.The generalized model involving these two discrete delays takes the following form
subject to the non negative conditionsx(s)=φ1(s)≥0,A(s)=φ2(s)≥0,y(s)=φ3(s)≥0,s∈[?τ,0],whereτ=max{τ1,τ2}andφi(s)∈C([?τ,0]→R+),(i=1,2,3).
The biological meaning of all parameters and variables in above models is provided in Tab.1.
First of all,we examine the boundedness and persistence of the system (1).
Theorem 3.1.The set
is a positive invariant set for all the solutions of model (1),initiating in the interior of the positive octant,whereδ=min{r,β,d}.
ProofThe model system (1) can be written in the matrix form
whereX=(x1,x2,x3)T=(x,A,y)T∈R3,andG(X)is given by
SinceG:R3→R3+is locally Lipschitz-continuous inΩandX(0)=X0∈R3+,the fundamental theorem of ordinary differential equation guarantees the local existence and uniqueness of the solution.Since [Gi(X)]xi(t)=0,X∈R3+≥0,it follows thatX(t)≥0 for allt≥0.In fact,from the first equation of model (1),it can easily be seen that|x=0≥0,|y=0≥0 and hencex(t)≥0,y(t)≥0 for allt≥0.Secondly,|A=0=λA0y≥0 for allt≥0 (asy(t)≥0 for allt≥0.) and henceA(t)≥0 for allt≥0.
From the first equation of model (1),we can write
which yields
Now,suppose
W(t)=c1x(t)+c2A(t)+y(t),
then we have
whereδ=min{r,β,d}.
Hence,it follows that
We also note that ifx(t)≥KandthenThis shows that all solutions of system (1) are bounded and remains inΩfor allt>0 if(x(0),A(0),y(0))∈Ω.
Theorem 3.2.Let the following inequalities are satisfied:
Then model system (1) is uniformly persistence,where,xais defined in the proof.
Proof:System (1) is said to be permanence or uniform persistence if there are positive constantsM1andM2such that each positive solutionX(t)=(x(t),A(t),y(t))of the system with positive initial conditions satisfies
Keeping the above in view,if we define
then from Theorem 3.1,it follows that
This also shows that for any sufficiently smallε>0,there exists aT>0 such that for allt≥T,the following holds:
Now from the first equation of model system (1),for allt≥T,we can write
Hence,it follows that
which is true for everyε>0,thus
wherer>α(1?A0)ys.
Now from the third equation of model system (1),we obtain
which implies
which is true for everyε>0,thus
for persistence,we must have
Second equation of model system (1) yields
Hence
TakingM1=min{xa,Aa,ya},the theorem follows.
Remark.Theorem 3.2 shows that threshold values for the persistence of the system are dependent on the parameterA0.
System (1) has four equilibrium points,trivial equilibriumE0(0,0,0),axial equilibriumE1(K,0,0),prey free equilibriumE2(0,,) and interior equilibriumE?(x?,A?,y?).E0andE1always exist.
? Existence ofE2(0,,):The prey free equilibriumE2is positive solution of the following system:
From the second equation of above system,we have
Putting the value ofAin the first equation of system (3),we get
Above equation has two positive roots if
System (1) has two prey free equilibrium under conditions given in (5):andAgain,Ifc2φλA0<φd+βe,then Eq.(4) does not have any positive root.Therefore,E2does not exist in this case.
? Existence of interior equilibriumE?(x?,A?,y?):It may be seen thatx?,A?andy?are the positive solution of the following system of algebraic equations:
From the second equation of system (6),we have
Putting this into the first and third equation of system (6),we obtain the following system:
We note the following points from Eq.(7):
1.Wheny=0,thenx=<0 orx=K>0.
2.Whenx=0 theny=>0.
Similarly,from Eq.(8),we note the following:
1.Wheny=0,thenx=
Hence,we can state the following theorem.
Theorem 3.3.The system (1) has a unique positive equilibriumE?(x?,A?,y?) if (9)and (10) hold.
Remark.The number of positive equilibrium for the system (1) depends on values of parameters,which we have chosen.Several possibilities are depicted in Fig.1.
The local behavior of a system in the vicinity of any existing equilibrium is very close to the behavior of its Jacobian system.So,we compute the Jacobian matrix to see the local behavior of the system around its equilibrium and we observe that
? The trivial equilibriumE0(0,0,0) is always a saddle point having stable manifold along the
Aandy-axes and unstable manifold along thex-axis.
? The axial equilibriumE1(K,0,0) is locally asymptotically stable ifIfthenE1is a saddle point having stable manifold along thexandA-axes and unstable manifold along they-axis.
? The Jacobian matrix evaluated at prey free equilibriumis given by
Characteristic equation is given by
The roots of Eq.(11) have negative real part if
Eq.(11) have at least one positive and one negative root if
Remark.By replacingbyandby,similar analysis holds good for the stability behavior of
Figure 1:Four possibilities of the prey and predator zero growth isoclines.(a) Interior equilibrium does not exist for the parametric values a=0.08,d=0.01,(b) interior equilibrium exists uniquely for the values of parameters a=0.1,d=0.235,(c) two interior equilibria for parameter values a=0.1,d=0.137,(d) three interior equilibria for parameter values a=0.105,d=0.1.Rest of the parameters are same as that in (25)
? In order to analyze the local stability of unique interior equilibriumE?(x?,A?,y?),we evaluate the Jacobian matrix atE?and it is given by
Characteristic equation corresponding to above matrix is given by
where
Now using the Routh-Hurwitz criterion,all eigenvalues ofJ|E?have negative real part iff
Thus we can state the following theorem.
Theorem 3.4The system (1) is stable in the neighborhood of its positive equilibrium iff inequalities in (15) hold.
It is also noted that inequalities in (15) hold if
Infect,the above two conditions imply thatA1>0 andA3>0.The third conditionA1A2>A3is also satisfied.
Remark.The system (1) is stable around its positive equilibriumE?if inequalities in (16) hold.
In the following theorem we give a criterion for global asymptotic stability of interior equilibriumE?(x?,A?,y?) of the system (1).
Theorem 3.5.The interior equilibriumE?(x?,A?,y?) of the system (1) is globally asymptotically stable under the following conditions:
Proof.We Choose a suitable Lyapunov function aboutE?as
whereγ1andγ2are positive constants,to be specified later.Now,differentiatingVwith respect totalong the solutions of system (1),we get
Choosingγ2=andγ1=we get
Applying Sylvester criterion,is negative definite if conditions in (17) hold.HenceE?is globally stable under conditions in (17).
Hopf-bifurcation is a local phenomenon where a system’s stability switches and a periodic solution arises around its equilibrium point by varying a parameter.In system (1),the parameterA0seems crucial,therefore we analyze the Hopf-bifurcation by takingA0as bifurcation parameter,then we have someA0=A?0.The necessary and sufficient conditions for occurrence Hopf-bifurcation atA0=A?0are
(a)A1|A?0>0,A3|A?0>0,
(b)f(A?0)≡(A1A2?A3)|A?0=0,
FromA1A2?A3=0,we get an equation inA0and assume that it has at least one positive rootA?0.Then for someε>0,there is an interval containingA?0,(A?0?ε,A?0+ε)such thatA?0?ε>0 andA2>0 forA0∈(A?0?ε,A?0+ε).Thus,Eq.(14) cannot have any real positive root forA0∈(A?0?ε,A?0+ε).
Therefore,atA0=A?0,Eq.(14) becomes
(Λ+A1)(Λ2+A2)=0,
this gives us three roots
Λ1,2=±iρ,Λ3=μ,
whereρ=andμ=?A1.
ForA0∈(A?0?ε,A?0+ε),roots can be taken as
Λ1,2=k1(A0)±ik2(A0),Λ3=?A1(A0).
Now,we have to verify the transversality condition.Differentiating Eq.(14) with respect to the bifurcation parameterA0,we obtain
whereR=A1A2?A3and,i=1,2,3 denote the derivative ofAiwith respect to time.Thus
Thus,we can state the following theorem.
Theorem 3.6.The system undergoes Hopf-bifurcation near interior equilibriumE?(x?,A?,y?)under the necessary and sufficient conditions (a),(b) and (c).Critical value of bifurcation parameterA0is given by the equationf(A?0)=0.
In order to see the stability and direction of Hopf-bifurcation,we use center manifold theorem [34] and some concepts used in [41].Now,consider the following transformation
x1=x?x?,x2=A?A?,x3=y?y?.
Using this transformation,system (1) takes the following form
whereX=(x1,x2,x3)T,
Letv1andv2be the eigenvectors corresponding to eigenvaluesiρa(bǔ)ndμofE?atA0=A?0.Thenv1andv2are given by
and
Let
where
Δ=p11(p22p33?p23p32)+p12(p23p31?p21p33)+p13(p21p32?p22p31)/=0,
q11=p22p33?p23p32,q12=p21p33?p23p31,q13=p21p32?p22p31,
q21=p12p33?p13p32,q22=p11p33?p13p31,q23=p11p32?p31p12,
q31=p12p23?p13p22,q32=p11p23?p13p21,q33=p11p22?p12p21.
Now letX=HYorY=H?1X,whereY= (y2,y2,y3)T.Using this transformation,system (18) can be written as
where
So,we can write system (19) as
Thus,system (20) takes the following form
whereU=(y1,y2)T,V=(y3),C=(μ),f=(f1,f2) andg=(f3).The eigenvalues ofBandCmay have zero real part and negative real parts,respectively.f,gvanish along with their first partial derivative at the origin.
Since the center manifold is tangent toWC(y=0) we can represent it as a graph
WC={(U,V):V=h(U)}:h(0)=h′(0)=0,
whereh:U→R2is defined on some vicinityU?R2of the origin [42,43].
We consider the projection of vector field onV=h(U)ontoWC:
Now we state the following theorem to approximate the center manifold.
Theorem 3.7.LetΦbe aC1mapping of a neighborhood of the origin inR2intoRwithΦ(0)=0 andΦ′(0)=0.If for someq>1,(NΦ)(U)=o(|U|q) asU→0,thenh(U)=Φ(U)+o(|U|q) asU→0,where(NΦ)(U)=Φ′(U)[BU+f(U,Φ(U))]?CΦ(U)?g(U,Φ(U)).
In order to approximateh(U),we consider
where h.o.t.stands for high order terms.Using (23),we get from (22)After simplification,we get
where
Equating both the sides of Eq.(24),we get
Using Crammer’s rule,
We can find the behavior of the solution of system (21) from the following theorem.
Theorem 3.8.If the zero solution of (22) is stable (asymptotically stable/unstable),then the zero solution of (21) is also stable (asymptotically stable/unstable).
Now from Eq.(22),we have
where
We determine the direction and stability of bifurcation periodic orbit of the system (21) by the following formula [44]
In above expression,ifν>0(<0),then the Hopf-bifurcation is supercritical (subcritical)and bifurcation periodic solution exists forA0=A?0.The bifurcating periodic solution is stable(unstable) if
The bifurcating direction of periodic solution of the system (1) is same as the system (21).
To validate our theoretical findings of model (1),we perform some numerical simulations using MATLAB R2018b.We have chosen the following dataset
For the above set of parameters,condition for existence of prey free equilibrium (5) and conditions for existence and uniqueness of interior equilibrium (9) and (10) are satisfied.Therefore,the system (1) has five equilibrium points.The behavior of these equilibrium points are given in Tab.2.
Table 2:Existing equilibria and their stability nature
The eigenvalues of the Jacobian matrix atE0andE1are (3,?0.32,?0.3) and(?3,?0.32,0.4113),respectively.ThereforeE0andE1both are saddle points.SimilarlyE2and ?E2are also saddle points.Again all the inequalities in (15) are satisfied.So according to Theorem 3.4,the interior equilibriumE?is locally asymptotically stable.The stability of system in the vicinity of the positive equilibriumE?is illustrated by Fig.2.In Fig.2a,time evolution of species is shown and it is noted that they converge to their equilibrium levels after some oscillations.In Fig.2b,phase diagram is drawn inxAy-space which shows the asymptotic stability behavior of positive equilibriumE?.
In this study,we found that predators dependency factorA0on additional food plays an important role in the dynamics of the system.If it is less than a threshold value then it can be the cause of destabilizing the system.The threshold value can be calculated by solvingf(A?0)=0(Theorem 3.6).By our computer simulation we obtain it asA?0=0.482.All the conditions of Theorem 3.6 are satisfied,so the system undergoes a Hopf-bifurcation atA?0=0.482.If we keep the value of parameterA0below its threshold value,then the system (1) always remains unstable.The instable behavior of solutions and presence of stable limit cycle atA0=0.45 In the model (1),consumption rate of additional foodφis also a vital parameter.We have noted that if system is stable for parameterA0(A0∈[0.482,1]) then it is stable for all range of parameterφ.But ifA0∈[0.2361,0.482) then system undergoes a Hopf-bifurcation with respect to parameterφ.In Fig.5,we have shown the bifurcation diagram whenA0=0.4 and other parameters are same as given in (25).The Hopf-bifurcation point isφ?=0.02847. Figure 2:Time series evolution (a) and phase portrait (b) of species for the set of parameters chosen in (25).Positive equilibrium E?is locally asympeoeically stable As the system (1) shows Hopf-bifurcation with respect to parametersA0andφ,and direction of Hopf-bifurcation is opposite for both the parameters.Therefore,we can divide theA0φ-plane into two regions Region of stability (green)S1={(A0,φ):system (1) is locally asymptotically stable}, Region of instability (white)S2={(A0,φ):system (1) is unstable}. Both the regions are drawn in Fig.6.The curve which separates both the regions is called Hopf-bifurcation curve. The number of interior equilibrium points depend on the values of parameters.In the table below,we have shown dependence of total number of interior equilibrium on parametersaanddand the nature of their stability.It is observed that whena=0.105 andd=0.1 (other parameters are as in (25)),then three interior equilibrium exist for the system (1),E?1(0.5099,1.398,20.9174),E?2(8.2355,1.409,32.9068) andE?3(42.309,1.4136,43.0591).E?1andE?3are locally asymptotically stable andE?2is unstable.Since there are two locally asymptotically stable equilibrium in the system,so it shows bistability.Bistability is a phenomenon where a system converges to two different equilibrium points for the same parametric values based on the variation of the initial conditions.In Fig.7,we initiated two trajectories from two nearby points and they converse to different interior equilibria.The black dotted curve is separatrix,which divides thexy-plane into two regions in such a way that if a solution is initiated from the left of the separatrix,it converses toE?1and if a solution is initiated from the right of the separatrix,it converses toE?3.In other words,left region is region of attraction forE?1and right region is region of attraction forE?3. Remark.For the best representation of bistability phenomenon and separatrix curve,the Fig.7 is drawn in thexy-plane.But initial conditions and interior equilibrium points are written as they are.Computer Modeling In Engineering&Sciences2021年2期