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. 2016 Aug 9;5(1):1303. doi: 10.1186/s40064-016-2958-y

Delay-induced periodic phenomenon in a diffusive regulated logistic model

Kejun Zhuang 1,2,, Gao Jia 3
PMCID: PMC4978664  PMID: 27547677

Abstract

The diffusive logistic growth model with time delay and feedback control is considered. First, the well-posedness and permanence of solutions are discussed by using some comparison techniques. Then, the sufficient conditions for stability of nonnegative constant steady states are established, and the occurrence of Hopf bifurcation at positive steady state is performed. Next, the bifurcation properties are derived by computing the normal form on center manifold. Our results not only supplement but also generalized some existing ones. Finally, some numerical simulations show the feasibility of our theoretical analyses.

Keywords: Logistic model, Feedback control, Hopf bifurcation, Reaction–diffusion system, Delay

Background

The classic logistic model

dN(t)dt=rN(t)1-N(t)K,r,k(0,+) 1

was first proposed by Verhulst in 1838. It can be utilized to describe the single–species growth and has been the basis of varieties of models in population ecology and epidemiology. For system (1) and its generalized forms, the significant results involve the asymptotic properties (Berezansky et al. 2004; Röst 2011), permanence and stability (Fan and Wang 2010; Chen et al. 2006), periodicity (Sun and Chen 2007) and almost periodicity (Yang and Yuan 2008) of solutions, Hopf bifurcation (Sun et al. 2007; Song and Yuan 2007; Song and Peng 2006; Chen and Shi 2012), traveling wave front (Zhang and Sun 2014), free boundary problem (Gu and Lin 2014), and so on. In addition, the Hopf bifurcation analyses for some diffusive predator–prey systems were also done (see Yang 2015; Yang and Zhang 2016a, b).

In particular, Gopalsamy (1993) considered the controlled delay system in the following form

dN(t)dt=rN(t)1-a1N(t)+a2N(t-τ)K-cu(t),du(t)dt=bN(t-τ)-au(t), 2

where all the coefficients and time delay τ are positive constants, N(t) is the number of individuals at time t, and variable u(t) denotes an indirect control variable (see Aizerman and Gantmacher 1964; Lefschetz 1965). They have derived the sufficient conditions to guarantee that the positive equilibrium solution is globally asymptotical stable.

Strictly speaking, spatial diffusion can not be ignored in studying the natural biological system (Murray 2003; Ghergu and Radulescu 2012). In the real world, most populations are moving and the densities are dependent of time and space. Therefore, diffusion should be taken into account in studying the basic logistic equation. However, there have been very few results on the influence of time delay on the reaction–diffusion logistic model with feedback control.

Inspired by the previous discussions, we mainly consider the reaction–diffusion system as follows:

N(x,t)t=d1ΔN(x,t)+rN(x,t)1-a1N(x,t)+a2N(x,t-τ)K-cu(x,t),u(x,t)t=d2Δu(x,t)+bN(x,t-τ)-au(x,t), 3

where (x,t)Ω×[0,+), Ω=(0,lπ).

The model (3) is considered with the initial value conditions as follows

N(x,t)=η1(x,t)0,u(x,t)=η2(x,t)0,x[0,lπ]×[-τ,0]. 4

We also assume that the model (3) is closed and there is no emigration or immigration across the boundary. Hence, the boundary conditions are considered as

Nν=uν=0,(x,t)Ω×[0,+), 5

where /ν represents the outward normal derivative on the boundary Ω.

In this paper, we develop a reaction–diffusion logistic model with time delay and diffusion, which makes up perfectly for the deficiencies of the previous literatures. The main objective is to explore the dynamics of system (3) by regarding τ as the bifurcation parameter. The structure of this paper is arranged as follows. In section “Preliminaries”, we derive the well–posedness of solutions and the permanence of the system. In section “Occurrence of the Hopf bifurcation”, we establish the existence of Hopf bifurcation. In section “Bifurcation properties”, we get the formulae for determining the Hopf bifurcation properties. In section “Numerical simulations”, we illustrate our theoretical results by some numerical simulations. Finally, we give some discussions and conclusions.

Preliminaries

As we know, spatial diffusion and time delay do not change the number and locations of constant equilibria because of no-flux boundary conditions. Then system (3) has two nonnegative equlibria E0=(0,0) and E=(N,u), where

N=aKa(a1+a2)+bcK,u=baN=bKa(a1+a2)+bcK.

Well–posedness of solutions

Here, for problem (3)–(5), we devote ourselves to the existence, uniqueness, nonnegativity and boundedness of solutions.

Theorem 1

For any given initial data satisfying the conditions (4) and boundary conditions (5), system (3) has a unique global solution of system and the solution maintains nonnegative and uniformly bounded for allt0.

Proof

Using the similar methods in Hattaf and Yousfi (2015), Hattaf and Yousfi (2015), we can get the local existence and uniqueness of solution (N(xt), u(xt)) with xΩ¯ and t[0,T), where T is the maximal existence time of solution.

It is easy to find that 0=(0,0) and M=(M1,M2) are a pair of coupled lower–upper solutions to problem (3)–(5), where

M1=maxKa1,sup-τs0φ1(·,s)C(Ω¯,R),M2=maxbM1a,sup-τs0φ2(·,s)C(Ω¯,R).

By means of the comparison theorem, we can obtain that 0N(x,t)M1 and 0u(x,t)M2 for xΩ¯ and t[0,T). It is obvious that the upper bound of solution is independent of the maximal existence interval [0, T). It follows from the standard theory for semilinear parabolic systems (Wu 1996; Henry 1993) that the solution globally exists. The proof is complete.

Dissipativeness and permanence

In the following, we will show that system (3) is permanent, which means that any nonnegative solution of (3) is bounded as t+ for all xΩ.

Theorem 2

(Dissipativeness) The nonnegative solution (Nu) of system (3) satisfies

lim supt+N(x,t)Ka1,lim supt+u(x,t)bKaa1.

Proof

Based on the first equation in system (3), we get

N(x,t)t-d1ΔN(x,t)rN(x,t)1-a1KN(x,t)for(x,t)Ω×[0,+).

Then from the standard comparison principle of parabolic equations, we can easily get

lim supt+N(x,t)Ka1.

For an arbitrary ε1>0, we could get a positive constant T1 such that for any tT1,

N(x,t)Ka1+ε1.

Thus, for any T[T1+τ,+), we have

u(x,t)t-d2Δu(x,t)bKa1+ε1-au(x,t).

This implies

lim supt+u(x,t)bKaa1

by comparison principle of parabolic equations and the arbitrariness of ε1.

Theorem 3

Ifaa1>aa2+bcK, then system (3) is permanent.

Proof

From Theorem 2, for an arbitrary ε2>0, we can find a constant T>T1+T2, such that

u(x,t)bKaa1+ε2

in Ω×[T2,+). Moreover, we can obtain

N(x,t)t-d1ΔN(x,t)rN(x,t)1-a2KKa1+ε1-cbKaa1+ε2-a1KN(x,t),

the comparison principle shows that

lim inft+N(x,t)Ka1aa1-aa2-bcKaa1>0

due to the continuity as ε10 and ε20.

Similarly, we can also have

lim inft+u(x,t)bKaa1aa1-aa2-bcKaa1>0.

Combining the results in Theorem 2, we can easily conclude that system (3) is permanent.

Occurrence of the Hopf bifurcation

For system (3), we shall study the local stability of two constant steady states and the occurrence of Hopf bifurcation phenomenon through discussing the distribution of characteristic values.

Denote

u1(t)=N(x,t),u2(t)=u(x,t),U(t)=(u1(t),u2(t))T.

By defining the phase space C=C([-τ,0],X), we can rewritten system (3) as the semilinear functional differential equation:

U˙(t)=DΔU(t)+G(Ut), 6

where X={(u,v)H2(0,lπ)×H2(0,lπ)|ux=vx=0,x=0,lπ}, Ut(·)=U(t+·), D=diag{d1,d2}, Δ=diag{2/x2,2/x2}, and G(Ut):CX is defined by

G(Ut)=ru1(t)1-a1u1(t)+a2u1(t-τ)K-cu2(t)bu1(t-τ)-au2(t).

The linear system of (6) at E0(0,0) is

U˙(t)=DΔU(t)+LE0(Ut), 7

where

LE0(φ)=rφ1(0)0bφ1(-τ)-aφ2(0)

for φ(θ)=Ut(θ), φ=(φ1,φ2)TC. The characteristic equation of (7) is

λy-DΔy-LE0(eλ·y)=0, 8

where ydom(Δ)\{0}, domΔX and eλ·(θ)y=eλθy for θ[-τ,0]. We know that the operator Δ in Ω with homogeneous Neumann boundary condition has the eigenvalues -n2/l2 and the corresponding eigenfunctions cos(nx/l), nN0={0,1,2,}. By using the Fourier expansion in (8),

y=n=0αnγncos(nx/l),

where αn, γnC. Therefore, the characteristic equation (8) can be transferred into

λ+d1n2l2-r0-be-λτλ+d2n2l2+a=0,nN0.

We then obtain the characteristic values as follows

λ1,n=-d1n2l2+r,λ2,n=-d2n2l2-a,nN0.

It is obvious that λ1,0=r>0, and we can establish the instability of E0.

Theorem 4

The trivial equilibriumE0of system (3) is always unstable.

Next, we will focus on the occurrence of Hopf bifurcation phenomenon.

Linearizing system (3) at E=(N,u), we get

U˙(t)=DΔU(t)+L(Ut), 9

where L:CX is given by

L(φ)=-ra1KNφ1(0)-ra2KNφ1(-τ)-cNφ2(0)bφ1(-τ)-aφ2(0)

with φ(θ)=Ut(θ), φ=(φ1,φ2)TC. Similar to the previous discussion, we can obtain the characteristic equation

λ2+Anλ+Bn+e-λτ(Cλ+Dn)=0,nN0, 10

where

An=(d1+d2)n2l2+a+ra1KN>0,Bn=d1d2n4l4+a+ra1KNd1n2l2+raa1KN>0,C=ra2KN>0,Dn=ra2KNd1n2l2+raa2KN+bcN>0.

For τ=0, Eq. (10) can be reduced to

λ2+(An+C)λ+Bn+Dn=0

with An+C>0 and Bn+D>0. On the basis of Routh–Hurwitz stability criterion, we can obtain the local stability of E when τ=0.

Lemma 1

The positive equilibrium is always locally asymptotically stable without time delay.

Remark 1

From Lemma 1, we can find that there is no Turing instability without time delay.

For τ0, let us suppose that λ=iω(ω>0) satisfies Eq. (10).

First, plugging λ=iω into Eq. (10) and then segregating the real and imaginary components with the help of Euler’s formula, we can get the following two equations of ω

ω2-Bn=Dncosωτ+Cωsinωτ,-ωAn=Cωcosωτ-Dnsinωτ.

Second, solving these equations, we can obtain

cosωτ=(Dn-AnC)ω2-BnDnC2ω2+Dn2,sinωτ=Cω3+(AnDn-BnC)ωC2ω2+Dn2. 11

Third, squaring both sides of those two equations and then adding them up, we get the following equation

ω4+An2-2Bn-C2ω2+Bn2-Dn2=0, 12

where

An2-2Bn-C2=d12+d22n4l4+2ra1KNd1+ad2n2l2+r2a12-a22K2N2,Bn2-Dn2=(Bn+Dn)d1d2n4l4+a+ra1KNd1n2l2+raa1KN-ra2KNd1n2l2-raa2KN-bcN.

Lemma 2

Forτ>0, we have

(i) Ifa1>a2+bcKar, then Eq. (10) does not have purely imaginary root.

(ii) Ifa2<a1<a2+bcKar, then there existsN0N0, such that Eq. (10) does not have purely imaginary root whenn>N0, and has a pair of conjugate purely imaginary eigenvalues when0nN0.

Proof

We can easily verify that An2-2Bn-C2>0 and Bn2-Dn2>0 when a1>a2+bcKar. This means that Eq. (12) has no positive root. In other words, there could be no purely imaginary root in Eq. (10) for any τ>0.

On the contrary, if a2<a1<a2+bcKar, then B02-D02<0 and there exists NN0 such that

Bn2-Dn2<0,n=0,1,2,,N0,Bn2-Dn20,n=N0+1,N0+2,.

That is to say, Eq. (12) has no positive root when n>N0 and has the unique positive root ωn when 0nN0, where

ωn=-(An2-2Bn-C2)+(An2-2Bn-C2)2-4(Bn2-Dn2)212.

By direct computation, we have

AnDn-BnC=ra2KNd22n4l4+bcNd1+2ra2KNd2+bcNd2n2l2+ra2a2KN+abcN+ra1bcKN2>0.

Moreover, Eq. (10) has characteristic values ±iωn with

τj(n)=τ0(n)+2jπωn,0nN0,j=0,1,2,,

where

τ0(n)=1ωnarccos(Dn-AnC)ωn2-BnDnDn2+C2ωn2.

This completes the proof.

We now check the transversality condition.

Lemma 3

Ifa2<a1<a2+bcKar, thendRe(λ)dττ=τj(n)>0forjN0andn{0,1,2,,N0} .

Proof

By taking the derivatives on both sides of (10) with respect to τ, we can get

2λdλdτ+Andλdτ+Ce-λτdλdτ+(Cλ+Dn)e-λτ-λ-τdλdτ=0,

and

dλdτ-1=2λ+An+Ce-λτ-τe-λτ(Cλ+Dn)λe-λτ(Cλ+Dn)=(2λ+An)eλτ+Cλ(Cλ+Dn)-τλ.

On the basis of (11) and (12), we get

dλdττ=τj(n)-1=(2iωn+An)cosωnτj(n)+isinωnτj(n)+Ciωn(iCωn+Dn)-τj(n)iωn=C+Ancosωnτj(n)-2ωnsinωnτj(n)-Cωn2+iDnωn-τj(n)iωn+i2ωncosωnτj(n)+Ansinωnτj(n)-Cωn2+iDnωn.

Further simplification will lead to

Redλdττ=τj(n)-1=Dnωn2ωncosωnτj(n)+Ansinωnτj(n)Cωn22+(Dnωn)2-Cωn2C+Ancosωnτj(n)-2ωnsinωnτj(n)Cωn22+(Dnωn)2=ωn4+Dn2-Bn2Cωn22+Dnωn2>0.

The proof is complete.

According to Lemmas 13 and the Hopf bifurcation theory developed by Wu (1996), the following conclusions can be drawn.

Theorem 5

Define

τ0=minn{0,1,2,,N0},jN0τj(n).

(i) Ifa1>a2+bcKar, then for anyτ>0, the positive equilibriumEis always locally asymptotically stable.

(ii) Ifa2<a1<a2+bcKar, thenEis locally asymptotically stable whenτ[0,τ0), and is unstable whenτ(τ0,+).

(iii) System (3) has a Hopf bifurcation fromEatτj(n)withn{0,1,2,,N0}andjN0. Ifn=0, the periodic solutions bifurcating positive equilibrium are all spatially homogeneous. Otherwise, these bifurcating periodic solutions are spatially inhomogeneous.

Bifurcation properties

In Theorem 5, we have demonstrated that there exist some spatially homogeneous or inhomogeneous periodic solutions when time delay crosses through some particular values. We are now in the position to investigate the bifurcation properties.

In general, we use τ to denote an arbitrary value of τj(n) with jN0 and n{0,1,2,,N0}. And we also use ±iω to denote the corresponding simply purely imaginary roots ±iωn.

Set N~(·,t)=N(·,τt), u~(·,t)=u(·,τt), U~(t)=(N~(·,t),u~(·,t)), and τ=τ+α with αR. For simplicity we drop the tilde and rewrite system (3) as follows,

dU(t)dt=τDΔU(t)+L(α)(Ut)+f(Ut,α), 13

where φ=(φ1,φ2)TC, L(α)(·):CX and f:C×RX are respectively denoted by

L(α)(φ)=(τ+α)-ra1KNφ1(0)+ra2KNφ1(-1)-cNφ2(0)bφ1(-1)-aφ2(0)

and

f(φ,α)=(τ+α)-2ra1Kφ12(0)-ra2Kφ1(0)φ1(-1)-rcφ1(0)φ2(0)0.

Note that α=τ-τ, we can find that system (13) may causes a Hopf bifurcation when α=0.

For the following linear differential equation:

U˙(t)=τDΔU(t)+L(α)(Ut), 14

we can easily deduce that the corresponding characteristic equation has characteristic values ±iωτ when α=0.

Next, we discuss the following differential equation:

Y˙(t)=-τDn2Y(t)+L(α)(Yt). 15

We can use Riesz representation theorem here, which tells us that there is a 2×2 matrix function η(θ,α)(-1θ0) with bounded variation elements satisfying

-τDn2l2φ(0)+L(α)(φ)=-10d[η(θ,α)]φ(θ),

where

η(θ,α)=(τ+α)-d1n2l2-ra1KN-cN0-d2n2l2-a,θ=0,0,θ(-1,0),(τ+α)-ra2KN0b0,θ=-1.

For ΦC1([-1,0],R2), ΨC1([0,1],R2), we define

A1(Φ(θ))=dΦ(θ)dθ,θ[-1,0),-10[dη0(θ)]Φ(θ),θ=0,A1(Ψ(s))=-dΨ(s)ds,s(0,1],-10[dη0(θ)]Ψ(-θ),s=0.

Then the formal adjoint, A1, of A1 is given by

(Ψ,Φ)0=Ψ¯(0)Φ(0)--10ζ=0θΨ¯(ζ-θ)d[η(θ,0)]Φ(ζ)dζ=Ψ¯(0)Φ(0)+τ-10Ψ¯(ζ+1)-ra2KN0b0Φ(ζ)dζ.

By calculation, we can find that q(θ)=(1,ξ)Teiωθτ and q(s)=M(1,η)eiωsτ are eigenvectors of A1 and A1 associated with iωτ, respectively, where

θ[-1,0],s[0,1],

and

ξ=be-iωτiω+a,η=cNiω+a,M=1+ξ¯η+τbη-ra2KNeiωτ-1.

Then P=span{q(θ),q(θ)¯}, P=span{q(s),q(s)¯} are the center subspace of system (3).

Define h·fn=h1βn1+h2βn2, fn=βn1,βn2 and βn1=cosnxl,0T, βn2=0,cosnxlT. The complex-valued L2 inner product on Hilbert space XC are

U1,U2=1lπ0lπ(u1v1¯+u2v2¯)dx, 16

for U1=(u1,u2),U2=(v1,v2)XC. And β0i,β0i=1, βni,βni=12, i=1,2, n=1,2,,

Φ,fn=Φ,βn1,Φ,βn2, 17

where ΦC([-1,0],X). We can establish the center subspace of system (14) at α=0 as follows

PCNL=(q(θ)z+q(θ)¯z¯)·fn,zC. 18

Based on the conclusions drawn by Wu (1996) and Hassard et al. (1981), the solutions of (13) are

Ut=(q(θ)z(t)+q(θ)¯z¯(t))·fn+W(z(t),z¯(t),θ),

where

W(z,z¯,θ)=W20z22+W11zz¯+W02z¯22+. 19

Moreover, for UtC0 of (13) at τ=τ, we have z˙=iωτz+g(z,z¯), where

g(z,z¯)=q(0)¯f(Ut,0),fn=g20z22+g11zz¯+g02z¯22+g21z2z¯2+. 20

By (16)–(20), we can compute

g20=0,n=1,2,,-2τM¯ra1K+ra2Ke-iωτ+rcξ,n=0,g11=0,n=1,2,,-2τM¯ra1K+rcRe{ξ}+ra2KRe{eiωτ},n=0,g02=g20¯,g21=-2M¯τlπ0lπ2ra1K(W11(1)(0)+W20(1)(0))cos2nxldx+0lπrcW11(2)(0)+12W20(2)(0)+12ξ¯W20(1)(0)+ξW11(1)(0)cos2nxldx+0lπra2Ke-iωτW11(2)(0)+12eiωτW20(2)(0)+12ξ¯W20(1)(-1)cos2nxldx+0lπra2KξW11(11)(-1)cos2nxldx.

Then we should compute W20(θ) and W11(θ) to determine g21. Following the formulas in Wu (1996), We can obtain that

W20(θ)=ig20ωτq(θ)+ig02¯3ωτq(θ)¯·fn+E1e2iωτθ,W11(θ)=-ig11ωτ1(θ)+ig11¯q(θ)¯ωτ·fn+E2,E1=E1×-2ra1K-2ra2Ke-iωτ-2rcξ0cos2nxl,E1=2iω+d1n2l2+ra1KN+ra2KNe-2iωτcN-be-2iωτ2iω+a+d2n2l2-1,

and

E2=E2×-2ra1K-2rcRe{ξ}-2ra2KRe{eiωτ}0cos2nxl,E2=d1n2l2+ra1KN+ra2KNcN-ba+d2n2l2-1.

From the previous expressions of g20, g11, g02 and g21, we can further compute

c1(0)=i2ωτg20g11-2|g11|2-13|g02|2+g212,μ2=-Re(c1(0))Re(λ(τ)),β2=2Re(c1(0)),T2=-1ωτ(Im(c1(0))+μ2Im(λ(τ))).

On account of preceding calculations, we arrive at the following conclusion on the bifurcation properties.

Theorem 6

The bifurcation direction is supercritical ifμ2>0, which means that the periodic solution exists forτ>τ0. On the contrary, the bifurcation direction is subcritical ifμ2<0, which means that the periodic solution exists forτ<τ0.

Moreover, the periodic solution is orbitally asymptotically stable ifβ2<0, or unstable ifβ2>0. The period of periodic solution is monotonically increasing at the time delayτwhenT2>0, or is monotonically decreasing at the time delayτwhenT2<0.

Numerical simulations

In this section, we give some numerical examples to test the preceding results with assistance of MATLAB.

For system (3), let Ω=(0,2π) and choose

d1=1,d2=0.5,r=0.6,a=b=c=1,a1=a2=2,K=1,

and the initial values N(x,0)=0.5 and u(x,0)=0.9. Then we can get the positive equilibrium E=(0.2,0.2). By direct computation, we have N0=0, ω00.348266, and τ0(0)5.81966, then the Hopf bifurcation values are given by

τj(n)=τj(0)=τ0(0)+2jπω0,j=0,1,2,

Concretely, τ0=τ0(0)5.81966, τ1(0)23.861, τ2(0)41.9024, ... From Fig. 1, we can see the asymptotical stability of positive equilibrium E when time delay is slightly smaller than the first bifurcation value τ0.

Fig. 1.

Fig. 1

The equilibrium E is stable when τ=2<τ0

Moreover, we can obtain c1(0)-1.4328+1.53343i. From Theorem 6, the Hopf bifurcation is supercritical, that is, the periodic solutions exist for τ>τ0, and they are orbitally asymptotically stable (see Fig. 2).

Fig. 2.

Fig. 2

Spatially periodic solution exists when τ=10

In the light of these simulations, we can find that spatially periodic solutions still exist even when τ=50(τ2(0),τ3(0)) and τ=130(τ6(0),τ7(0)) (see Figs. 3, 4).

Fig. 3.

Fig. 3

The spatially periodic solution still exists when τ=50

Fig. 4.

Fig. 4

The spatially periodic solution still exists even when τ=130

Discussions and conclusions

In this paper, we considered the reaction–diffusion regulated logistic growth model. We have investigated the basic properties and Hopf bifurcation under the Neumann boundary conditions. It is shown that the logistic model may undergo Hopf bifurcation when time delay varies. We further give the formulae for determining the bifurcation properties, such as the direction of bifurcation, the stability of periodic solution and the monotonicity of period of periodic solution.

Here, we only discussed the single–species diffusive model with feedback control. In fact, how spatial diffusion and time delay affect the dynamic behaviors of multi–species controlled model remains unclear. We will focus on these novel and interesting models in the future.

Furthermore, from the numerical simulations in section “Numerical simulations”, we conjecture that the Hopf bifurcation induced by time delay is global. This means that the periodic solutions due to Hopf bifurcation still exist even if the time delay is sufficiently large.

Authors' contributions

KZ carried out the genetic studies and drafted the manusctipt. GJ designed the structure of this paper and helped to draft the manuscript. Both authors read and approved the final manuscript.

Acknowledgements

This work is supported by the National Natural Science Foundation of China (11171220) and the Hujiang Foundation of China (B14005). This work is also sponsored by Key Project for Excellent Young Talents Fund Program of Higher Education Institutions of Anhui Province (gxyqZD2016100).

Competing interests

The authors declare that they have no competing interests.

Contributor Information

Kejun Zhuang, Phone: +86-13305525048, Email: zhkj123@163.com.

Gao Jia, Email: gaojia89@163.com.

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