Bogdanov-Takens bifurcations in the enzyme-catalyzed reaction comprising a branched network

  • Received: 02 July 2016 Accepted: 01 January 2017 Published: 01 October 2017
  • MSC : 34C23, 92C45

  • There have been some results on bifurcations of codimension one (such as saddle-node, transcritical, pitchfork) and degenerate Hopf bifurcations for an enzyme-catalyzed reaction system comprising a branched network but no further discussion for bifurcations at its cusp. In this paper we give conditions for the existence of a cusp and compute the parameter curves for the Bogdanov-Takens bifurcation, which induces the appearance of homoclinic orbits and periodic orbits, indicating the tendency to steady-states or a rise of periodic oscillations for the concentrations of the substrate and the product.

    Citation: Qiuyan Zhang, Lingling Liu, Weinian Zhang. Bogdanov-Takens bifurcations in the enzyme-catalyzed reaction comprising a branched network[J]. Mathematical Biosciences and Engineering, 2017, 14(5&6): 1499-1514. doi: 10.3934/mbe.2017078

    Related Papers:

    [1] Mohammad Sajid, Sahabuddin Sarwardi, Ahmed S. Almohaimeed, Sajjad Hossain . Complex dynamics and Bogdanov-Takens bifurcations in a retarded van der Pol-Duffing oscillator with positional delayed feedback. Mathematical Biosciences and Engineering, 2023, 20(2): 2874-2889. doi: 10.3934/mbe.2023135
    [2] Mengyun Xing, Mengxin He, Zhong Li . Dynamics of a modified Leslie-Gower predator-prey model with double Allee effects. Mathematical Biosciences and Engineering, 2024, 21(1): 792-831. doi: 10.3934/mbe.2024034
    [3] Hongqiuxue Wu, Zhong Li, Mengxin He . Dynamic analysis of a Leslie-Gower predator-prey model with the fear effect and nonlinear harvesting. Mathematical Biosciences and Engineering, 2023, 20(10): 18592-18629. doi: 10.3934/mbe.2023825
    [4] Kunlun Huang, Xintian Jia, Cuiping Li . Analysis of modified Holling-Tanner model with strong Allee effect. Mathematical Biosciences and Engineering, 2023, 20(8): 15524-15543. doi: 10.3934/mbe.2023693
    [5] Yingzi Liu, Zhong Li, Mengxin He . Bifurcation analysis in a Holling-Tanner predator-prey model with strong Allee effect. Mathematical Biosciences and Engineering, 2023, 20(5): 8632-8665. doi: 10.3934/mbe.2023379
    [6] Xiaoli Wang, Junping Shi, Guohong Zhang . Bifurcation analysis of a wild and sterile mosquito model. Mathematical Biosciences and Engineering, 2019, 16(5): 3215-3234. doi: 10.3934/mbe.2019160
    [7] Zhenliang Zhu, Yuming Chen, Zhong Li, Fengde Chen . Dynamic behaviors of a Leslie-Gower model with strong Allee effect and fear effect in prey. Mathematical Biosciences and Engineering, 2023, 20(6): 10977-10999. doi: 10.3934/mbe.2023486
    [8] Juan Li, Yongzhong Song, Hui Wan, Huaiping Zhu . Dynamical analysis of a toxin-producing phytoplankton-zooplankton model with refuge. Mathematical Biosciences and Engineering, 2017, 14(2): 529-557. doi: 10.3934/mbe.2017032
    [9] Juan Ye, Yi Wang, Zhan Jin, Chuanjun Dai, Min Zhao . Dynamics of a predator-prey model with strong Allee effect and nonconstant mortality rate. Mathematical Biosciences and Engineering, 2022, 19(4): 3402-3426. doi: 10.3934/mbe.2022157
    [10] Yuhong Huo, Gourav Mandal, Lakshmi Narayan Guin, Santabrata Chakravarty, Renji Han . Allee effect-driven complexity in a spatiotemporal predator-prey system with fear factor. Mathematical Biosciences and Engineering, 2023, 20(10): 18820-18860. doi: 10.3934/mbe.2023834
  • There have been some results on bifurcations of codimension one (such as saddle-node, transcritical, pitchfork) and degenerate Hopf bifurcations for an enzyme-catalyzed reaction system comprising a branched network but no further discussion for bifurcations at its cusp. In this paper we give conditions for the existence of a cusp and compute the parameter curves for the Bogdanov-Takens bifurcation, which induces the appearance of homoclinic orbits and periodic orbits, indicating the tendency to steady-states or a rise of periodic oscillations for the concentrations of the substrate and the product.


    1. Introduction

    Many differential equations have been proposed (see [8,11,13], [17]-[19], [21]-[22], [24,27] and references therein) to model the dynamic changes of substrate concentration and product one in enzyme-catalyzed reactions. Among those models, a typical form ([7]) is the following skeletal system

    {˙x=vV1(x,y)V3(x),˙y=q(V1(x,y)V2(y)), (1)

    where x and y denote the concentrations of the substrate and the product respectively, v and q are both positive constants, V1(x,y) and V2(y) denote the enzyme reaction rate and the output rate of the product respectively and satisfy that

    V1(0,y)=0,  V1/x>0,  V1/y>0,  V2(y)0,  x,y>0,

    and V3(x) denotes the branched-enzyme reaction rate. Figure 1 shows the scheme of the enzyme-catalyzed reaction which comprises a branched network from the substrate. In Figure 1, S and P represent the substrate and product, respectively, and E1,E2 and E3 are the three enzymes.

    Figure 1. Reaction scheme.

    The case that V3(x)0 in system (1), which represents an unbranched reaction, has been discussed extensively in [1,6,7,9,20]. Recently, more efforts were made to the case that V3(x)0. One of the efforts ([12,23]) is made for V1(x,y)=xmyn,V2(y)=y and V3(x)=lx and v=1, with which system (1) reduces to

    {˙x=1xmynlx,˙y=q(xmyny),

    called the multi-molecular reaction model sometimes, where m,n1 are integers and l0 is real. All local bifurcations of this system such as saddle-node bifurcation, Hopf bifurcation and Bogdanov-Takens bifurcation were discussed in [12] and [23]. Reference [15] is concerned with the case that V1(x,y)=γxmyn, V3(x)=βx, q=1 and V2(y) is a saturated reaction rate, i.e., V2(y)=v2y/(μ2+y), with which (1) reduces to

    {˙x=vγxmynβx,˙y=γxmynv2yμ2+y,

    where v,γ>0,μ2,v2 and β0. Results on existence and nonexistence of periodic solutions on Hopf bifurcation were obtained in [15] with n=1 and β=0. When V2(y) and V3(x) are both saturated reaction rates, system (1) was considered in [16] as

    {˙x=vV1(x,y)v3xu3+x,˙y=q(V1(x,y)v2yu2+y)

    with V1(x,y)=v1x(1+x)(1+y)2/[L+(1+x)2(1+y)2], where L is the allosteric constant of E1. Varying the parameter v2 but fixing the other parameters, Liu ([16]) investigated numerically how the enzyme saturation affects the emergence of dynamical behaviors such as the change from a stable oscillatory state to a divergent state. Later, Davidson and Liu ([3]) discussed the saddle-node bifurcation, Hopf bifurcation and the global bifurcation corresponding to the appearance of homoclinic orbit. When V2(y) and V3(x) are both saturated reaction rates, system (1) was also considered in [4] as

    {˙x=vv1xyv3xu3+x,˙y=q(v1xyv2yu2+y) (2)

    with V1(x,y)=v1xy. With a change of variables x=u3˜x, y=u2˜y and the time rescaling tv11μ12t, system (2) can be written as

    {˙x=axybx1+x,˙y=κy(xc1+y), (3)

    where we still use x, y to present ˜x, ˜y and take notations a:=v11u13u12v, b:=v11u13u12v3, c:=v11u13u12v2 and κ:=u12qu3 for positive constants. Actually, system (3) is orbitally equivalent to the following quartic polynomial differential system

    {˙x=(1+y){(1+x)(axy)bx},˙y=κ(1+x)y{(1+y)xc}, (4)

    in the first quadrant Q+:={(x,y):x0,y0} by a time scaling dτ=(x+1)(y+1)dt. In [4] Davidson, Xu and Liu discussed the case that k=1 and a<c, where the system has at most two equilibria, giving the existence of limit cycles (by the Poincaré-Bendixson Theorem seen in [10] or [26]) and the non-existence of periodic orbits (by the Dulac Criterion seen in [10] or [26]), proving the uniqueness of limit cycles (by reducing to the form of Liénard system) with some restrictions, and illustrating with the software AUTO saddle-node bifurcation, transcritical bifurcation and Hopf bifurcation for fixed κ=1,b=1.5 and c=5. Recently, the general case that κ,a,b,c>0 was discussed in [27], where all codimension-one bifurcations such as saddle-node, transcritical and pitchfork bifurcations were investigated and the weak focus was proved to be of at most order 2.

    In this paper we continue the work of [27] to give conditions for the existence of a cusp and compute the parameter curves for the Bogdanov-Takens bifurcation, which induces the appearance of homoclinic orbits and periodic orbits, indicating the tendency to steady-states or a rise of periodic oscillations for the concentrations of the substrate and product.


    2. Condition for cusp

    It is proved in [27] that system (4) has at most 3 equilibria, i.e., E0:(a/(ba),0), E1:(p1,c/p11) and E2:(p2,c/p21), where

    p1:=12{(abc+1)[(abc+1)24(ac)]1/2},p2:=12{(abc+1)+[(abc+1)24(ac)]1/2}. (5)

    Moreover, if a=a:=c+(b1/21)2, then E1 and E2 coincide into one, i.e., the equilibrium E:(b1/21,c(b1/2+1)/(b1)1). There are found in [27] totally 6 bifurcation surfaces

    TE0:={(a,b,c,κ)R4+|a=bc/(1+c),b(c+1)2}:=4i=1T(i)E0,PE0:={(a,b,c,κ)R4+|a=bc/(1+c),b=(c+1)2},HE1:={(a,b,c,κ)R4+|κ=κ1,bc/(1+c)<a<c,0<b1}{(a,b,c,κ)R4+|κ=κ1,bc/(1+c)<a<c+(b1/21)2,1<b<(c+1)2},SNE:={(a,b,c,κ)R4+|a=a, 1<b<(c+1)2,κκ}:=4i=1SN(i)E,B1:={(a,b,c,κ)R4+|a=c},B2:={(a,b,c,κ)R4+|a=b},

    which divide R4+:={(a,b,c,κ):a>0,b>0,c>0,κ>0} into 8 subregions

    R1:={(a,b,c,κ)R4+|c<a<a,1<b<c,c>1,  or  b<a<a,c<b<(c+1)2/4,c>1},R2:={(a,b,c,κ)R4+|b<a<c,0<b<c}R3:={(a,b,c,κ)R4+|bc/(1+c)<a<b,0<b<c  or  bc/(1+c)<a<c,c<b<c+1},R4:={(a,b,c,κ)R4+|0<a<bc/(1+c),0<b<c+1  or  0<a<c,b>c+1},R5:={(a,b,c,κ)R4+|c<a<bc/(1+c),b>c+1},R6:={(a,b,c,κ)R4|c<a<b,c<b<(c+1),c>3  or  bc/(1+c)<a<b,c+1<b<(c+1)2/4,c>3  or  bc/(1+c)<a<a,(c+1)2/4<b<(c+1)2,c>3  or  c<a<b,c<b<(c+1)2/4,1<c3  or  c<a<a,(c+1)2/4<b<c+1,1<c3  or  bc/(1+c)<a<c+(b1/21)2,(c+1)<b<(c+1)2,c3  or  c<a<a,1<b<c+1,c1},R7:={(a,b,c,κ)R4+|c+(b1/21)2<a<b,(c+1)2/4<b<(c+1)2,c>1  or  bc/(1+c)<a<b,b>(c+1)2  or  c<a<b,c<b<1,c1  or  c+(b1/21)2<a<b,1<b<(c+1)2,c1},R0:=R4+{PE0SNETE0(2i=1Bi)B(7i=1Ri)},

    where

    T(1)E0:={(a,b,c,κ)R4+|a=bc/(1+c),0<b<c+1},T(2)E0:={(a,b,c,κ)R4+|a=bc/(1+c),c+1<b<(c+1)2},T(3)E0:={(a,b,c,κ)R4+|a=bc/(1+c),b>(c+1)2},T(4)E0:={(a,b,c,κ)R4+|a=bc/(1+c),b=c+1},SN(1)E:={(a,b,c,κ)R4+|a=a, 1<b<(c+1)2/4,c>1,κκ},SN(2)E:={(a,b,c,κ)R4+|a=a, b=(c+1)2/4,c>1,κκ},SN(3)E:={(a,b,c,κ)R4+|a=a, (c+1)2/4<b<(c+1)2,c>1,κκ},SN(4)E:={(a,b,c,κ)R4+|a=a, 1<b<(c+1)2,c1,κκ},κ1:=p21{(p1+1)(cp1)}1c{p1(cp1)+a},κ:=(cb1/2+1)1(b1/21)2c2. (6)

    The following lemma is a summary of Theorems 1, 2 and 3 of [27].

    Lemma 2.1. (ⅰ) System (4) has a saddle-node E0 if (a,b,c,κ)TE0PE0. Moreover, as (a,b,c,κ) crosses either T(1)E0 from R3 to R4, T(2)E0 from R6 to R5, or T(4)E0 from R6 to R4, a saddle E0 and a stable (resp. unstable) node E1 merge into a stable node E0 on the boundary of the first quadrant Q+ for κ<κ1(resp. κ>κ1) through a transcritical bifurcation; as (a,b,c,κ) crosses T(3)E0 from R5 to R7, a stable node E0 and a saddle E2 merge into a saddle E0 on the boundary of Q+ through a transcritical bifurcation; as (a,b,c,κ) crosses PE0 from R7 to R5, a saddle E0 changes into a stable node E0, a saddle E2 through a pitchfork bifurcation at E0 on the boundary of Q+.

    (ⅱ) System (4) has a weak focus E1 of at most order 2 for (a,b,c,κ)HE1, which is of order exactly and produces at most limit cycles through Hopf bifurcations as (a,b,c,κ)H()E1, =1,2, where H(1)E1:=HE1H(2)E1 and

    H(2)E1:={(a,b,c,κ)HE1:2p1(p1+1)a3+{(p21+p1+1)c2+p1(2p21+p12)c3p31(p1+1)}a2(cp1){(p31+3p21+p1+1)c2+2p21(p21+3p1+3)c+3p41(p1+1)}a+p21{(p1+2)c+p21}{cp1(p1+1)}(cp1)2=0}.

    (ⅲ) System (4) has a saddle-node E if (a,b,c,κ)SNE. Moreover, as (a,b,c,κ) crosses either SN(1)E from R0 to R1, SN(2)E from R0 to R6, or SN(3)ESN(4)E from R7 to R6, a stable (resp. unstable) node E1 and a saddle E2 arise through a saddle-node bifurcation for κ<κ1(resp. κ>κ1).

    The above Lemma 2.1 does not consider parameters in the set

    B:={(a,b,c,κ)R4+|a=a, 1<b<(c+1)2,κ=κ}, (7)

    where a is given below (5) and κ is given in (6). B is actually the intersection of the saddle-node bifurcation surface SNE and the Hopf bifurcation surface HE1, which are described by the curves ^SNE and ^HE1 respectively on the section {(a,b,c,κ)R4+|b=2, c=1} in Figure 2. The intersection of ^SNE and ^HE1 indicates B.

    Figure 2. Bifurcation surfaces projection on the (a,κ)-plane.

    This paper is devoted to bifurcations in B. For (a,b,c,κ)B, equilibrium E is degenerate with two zero eigenvalues. In the following lemma we prove that E is a cusp.

    Lemma 2.2. If (a,b,c,κ)BC, where

    C:={(a,b,c,κ)B|c=ς(b):=14b1/2(b1/21){b1/2+2+(17b12b1/2+4)1/2}},

    then equilibrium E is a cusp in system (4).

    Proof. For simplicity in statements, we use the notation

    p:=b1/21. (8)

    For (a,b,c,κ)B, system (4) can be transformed into the form

    {˙x=y+c(p2+cp+c)p3x2+1p+1xypc2(p+1)y2c(p2+c)p4x3p2+2pc+2cp2c(p+1)x2y2p+1c2(p+1)2xy2c2p4x42p2(p+1)x3y1c2(p+1)2x2y2,˙y=c3(p+1)p3x2c2(p+1)p2(cp)xy1cpy2(p+1)(p2+c)p5(cp)x3c(p2+2pc+2c)p3(cp)x2y2p+1p(p+1)(cp)xy2c4(p+1)p5(cp)x42c2p3(cp)x3y1p(p+1)(cp)x2y2, (9)

    by translating E to the origin O and Jordanizing the linear part of system (4). For convenience, introducing new variables (x,y)(u,v), where u=x and v denotes the right-hand side of the first equation in (9), we change (9) into the Kukles form

    {˙u=v,˙v=c3(p+1)p3u2+c{(2p+2)c2(p2+3p)c2p3}p3(cp)uv+c2p1(p+1)(cp)v2+c3(p2+c)p4(cp)u3c{(p+1)(p+3)c2+p(p23p3)cp3(3p+2)}p4(p+1)(cp)u2v(5p2+8p+4)c+2p2(p+1)cp2(p+1)2uv21c2(p+1)v3c2(c2+2p2cp3)p5(cp)u4+1p5(p+1)2(cp){(p+4)(p+1)2c3+p(7p3+7p23p4)c2p3(8p2+15p+8)c2p5(p+1)}u3v+(3p3+6p2+6p+2)c2+p(2p+1)(2p2+2p1)cp3(p+1)(7p+4)cp3(p+1)3(cp)u2v2(3p+4)c23p(p+2)c2p3c3p(p+1)2(cp)uv32c3pc4(p+1)2(cp)v4+O(|u,v|5). (10)

    Since the linear part is nilpotent, by Theorem 8.4 in [14] system (10) is conjugated to the Bogdanov-Takens normal form, i.e., the right-hand side of the second equation is a sum of terms of the form auk+buk1v. Hence, one can use the transformation uu, vvc2p1(p+1)(cp)uv together with the time-rescaling dt=(1c2p1(p+1)(cp)u)dτ to change system (10) into the following

    {˙u=v,˙v=c3(p+1)p3u2+c{(2p+2)c2(p2+3p)c2p3}p3(cp)uv+O(|u,v|3), (11)

    where the term of v2 is eliminated and terms of degree 2 are normalized. The term of u2 exists since c3(p+1)/p30. For the existence of the term of uv, we need to discuss on the quadratic equation

    c2p2+3p2(p+1)cp3p+1=0, (12)

    which comes from the numerator of the coefficient of uv. Since the constant term is negative for p>0, the quadratic equation (12) has exactly one positive root

    c=14(p+1)1p{p+3+(17p2+22p+9)1/2},

    which defines the function ς(b) as shown in Lemma 2.2 with the replacement (8). It implies by Theorem 8.4 of [14] that for cς(b), i.e., (a,b,c,κ)SC, O is a cusp of system (11). The proof of this lemma is completed.


    3. Bogdanov-Takens bifurcation

    In this section we discuss in the case that (a,b,c,κ)BC, in which system (4) is of codimension 2. We choose a,κ as the bifurcation parameters and unfold the Bogdanov-Takens normal forms of codimensions 2 when (a,κ) is perturbed near the point (a,κ), where a is given below (5) and κ is given in (6).

    Theorem 3.1. If (a,b,c,κ)BC, where B is defined in (7) and C is defined as in Lemma 2.2, then there are a neighborhood U of the point (a,κ) in the (a,κ)-parameter space and four curves

    SN+:={(a,κ)U|a=a, κ>κ,0<c<ς(b)}{(a,κ)U|a=a, κ<κ,c>ς(b)},SN:={(a,κ)U|a=a, κ<κ,0<c<ς(b)}{(a,κ)U|a=a, κ>κ,c>ς(b)},H:={(a,κ)U|a=a((2b1/2+1)c2((b1/21)2+3(b1/21))c             2(b1/21)3)2b1/2(b1/21)6(cb1/2+1)4(κκ)2+O(|κκ|3),             κ>κ,0<c<ς(b)}{(a,κ)U|a=a((2b1/2+1)c2((b1/21)2+3(b1/21))c             2(b1/21)3)2b1/2(b1/21)6(cb1/2+1)4(κκ)2+O(|κκ|3),             κ<κ,c>ς(b)},L:={(a,κ)U|a=a49/25((2b1/2+1)c2((b1/21)2+3(b1/21))c             2(b1/21)3)2b1/2(b1/21)6(cb1/2+1)4(κκ)2+O(|κκ|3),             κ>κ,0<c<ς(b)}{(a,κ)U|a=a49/25((2b1/2+1)c2((b1/21)2+3(b1/21))c             2(b1/21)3)2b1/2(b1/21)6(cb1/2+1)4(κκ)2+O(|κκ|3),             κ<κ,c>ς(b)},

    such that system (4) produces a saddle-node bifurcation near E as (a,c) acrosses SN+SN, a Hopf bifurcation near E as (a,κ) acrosses H, and a homoclinic bifurcation near E as (a,κ) acrosses L, where κ and ς(b) are given in (6) and Lemma 2.2 respectively.

    The above bifurcation curve H is exactly the same as HE1 given in Lemma 2.1, and the union SN+SN+ is exactly the bifurcation curves SNE given in Lemma 2.1.

    Proof. Let p=b1/21 and

    ε1:=aa,ε2:=κκ, (13)

    and consider |ε1| and |ε2| both to be sufficiently small. Expanding system (4) at E, we get

    {˙x=c(p+1)pε1+(c2(p+1)p2+cpε1)x+(c(p+1)+(p+1)ε1)yc(cp)p2x2+(c(2+3p)p+ε1)xyp(p+1)y2+O((x,y)3),˙y=(c3(p+1)p4+c(p+1)(cp)p2ε2)x+(c2(p+1)p2+(p+1)(cp)ε2)y+(c3p4+c(cp)p2ε2)x2+(c3(2+3p)c2p(2p+1)(cp)p3+c(3p+2)p(2p+1)pε2)xy+(c2(p+1)(cp)p+p(p+1)ε2)y2+O((x,y)3). (14)

    Introducing new variables (x,y)(ξ1,η1), where ξ1=x and η1 denotes the right-hand side of the first equation in (14), we change (14) into the Kukles form, whose second order truncation is the following

    {˙ξ1=η1,˙η1=E00(ε1,ε2)+E10(ε1,ε2)ξ1+E20(ε1,ε2)ξ21+F(ξ1,ε1,ε2)η1+E02(ε1,ε2)η21, (15)

    where F(ξ1,ε1,ε2):=E01(ε1,ε2)+E11(ε1,ε2)ξ1 and Eij s (i,j=0,1,2) are given in Appendix. Notice that (a,b,c,κ)BC implies that cς(b). From (12) we see that the quadratic equation has exactly one positive root c=ς(b). Thus, for cς(b) we can check that

    F(0,0,0)=0,   Fξ1(0,0,0)=E11(0,0)=(2p+2)(c2p2+3p2(p+1)cp3p+1)0.

    By the Implicit Function Theorem, there exists a function ξ1=ξ1(ε1,ε2) defined in a small neighborhood of (ε1,ε2)=(0,0) such that ξ1(0,0)=0 and F(ξ1(ε1,ε2),ε1,ε2)=0. Thus, from the definition of F we obtain ξ1(ε1,ε2)=E01(ε1,ε2)/E11(ε1,ε2) near (0,0). Then, we use a parameter-dependent shift

    ξ2=ξ1ξ1(ε1,ε2),η2=η1

    to vanish the term proportional to η2 in the equation for η2 from system (15), which leads to the following system

    {˙ξ2=η2,˙η2=ψ1(ε1,ε2)+ψ2(ε1,ε2)ξ2+E20(ε1,ε2)ξ22+E11(ε1,ε2)ξ2η2+E02(ε1,ε2)η22, (16)

    where

    ψ1(ε1,ε2):=E00(ε1,ε2)+E10(ε1,ε2)ξ1(ε1,ε2)+E20(ε1,ε2)ξ21(ε1,ε2),ψ2(ε1,ε2):=E10(ε1,ε2)+2ξ1(ε1,ε2)E20(ε1,ε2).

    In order to eliminate the η22 term, one can use the transformation

    ξ3=ξ2,   η3=η2E02(ε1,ε2)ξ2η2

    together with the time-rescaling dt=(1E02(ε1,ε2)ξ2)dτ to change system (16) into the following

    {˙ξ3=η3,˙η3=ζ1(ε1,ε2)+ζ2(ε1,ε2)ξ3+˜E20(ε1,ε2)ξ23+E11(ε1,ε2)ξ3η3, (17)

    where

    ζ1(ε1,ε2):=ψ1(ε1,ε2),   ζ2(ε1,ε2):=ψ2(ε1,ε2)ψ1(ε1,ε2)E02(ε1,ε2),˜E20(ε1,ε2):=E20(ε1,ε2)E10(ε1,ε2)E02(ε1,ε2).

    Further, in order to reduce coefficient of ξ23 to 1, we apply the transformation

    u=˜E20(ε1,ε2)E211(ε1,ε2)ξ3,v=sign(E11(ε1,ε2)˜E20(ε1,ε2))˜E220(ε1,ε2)E311(ε1,ε2),

    where ˜E20(0,0)=c3(p+1)p3<0, and the time-scaling dt=|E11(ε1,ε2)˜E20(ε1,ε2)|dτ to system (17) and obtain

    {˙u=v,˙v=ϕ1(ε1,ε2)+ϕ2(ε1,ε2)u+u2+ϑuv, (18)

    where ϑ=sign(E11(0,0)˜E20(0,0)),

    ϕ1(ε1,ε2):=E411(ε1,ε2)˜E320(ε1,ε2)ζ1(ε1,ε2)={(2p+2)c2(p2+3p)c2p3}4ε1ϕ11(ε1,ε2)p4(cp)4ϕ212(ε1,ε2), ϕ2(ε1,ε2):=E211(ε1,ε2)˜E220(ε1,ε2)ζ2(ε1,ε2)=2{(2p+2)c2(p2+3p)c2p3}ϕ21(ε1,ε2)c3/2(cp)2(p+1)1/2pϕ3/212(ε1,ε2),

    and polynomials ϕij s are given in the Appendix.

    Let

    μ1=ϕ1(ε1,ε2),μ2=ϕ2(ε1,ε2), (19)

    where ϕ1 and ϕ2 are defined just below (18). Clearly, ϕ1(0,0)=ϕ2(0,0)=0. Compute the Jacobian determinant of (19) at the point (0,0)

    |ϕ1(ε1,ε2)ε1ϕ1(ε1,ε2)ε2ϕ2(ε1,ε2)ε1ϕ2(ε1,ε2)ε2|(ε1,ε2)=(0,0)={(2p+2)c2(p2+3p)c2p3}5p6c4(cp)4(p+1)0, (20)

    implying that (19) is a locally invertible transformation of parameters. This transformation makes a local equivalence between system (18) and the versal unfolding system

    {˙˜u=˜v,˙˜v=μ1+μ2˜u+˜u2+ϑ˜u˜v, (21)

    where ϑ is given in (18). As indicated in Section 7.3 of [10], system (21) has the following bifurcation curves

    SN+:={(μ1,μ2)V0 | μ1=0, μ2>0},SN:={(μ1,μ2)V0 | μ1=0, μ2<0},H:={(μ1,μ2)V0 | μ1=μ22, μ2>0},L:={(μ1,μ2)V0 | μ1=4925μ22+o(|μ2|2), μ2>0}, (22)

    where V0 is a small neighborhood of (0,0) in R2.

    In what follows, we present above bifurcation curves in parameters ε1 and ε2 in explicit forms. For this purpose, we need the relation between (ε1,ε2) and (μ1,μ2). Note that ϕ1 and ϕ2 defined just below (18) are Ck near the origin (0, 0)(k is an arbitrary integer). By condition (20), the well-known Implicit Function Theorem implies that there are two Ck functions

    ε1=ω1(μ1,μ2),    ε2=ω2(μ1,μ2) (23)

    in a small neighborhood of (0,0,0,0) such that ω1(0,0)=ω2(0,0)=0 and

    μ1=ϕ1(ω1(μ1,μ2),ω2(μ1,μ2)),   μ2=ϕ2(ω1(μ1,μ2),ω2(μ1,μ2)). (24)

    Substitute the second order formal Taylor expansions of ω1 and ω2 in (24) while expand ϕ1 and ϕ2 in (24) to the second order

    ϕ1(ε1,ε2)={(2p+2)c2(p2+3p)c2p3}4ε1/{p6c2(cp)4(p+1)}{(2p+2)c2(p2+3p)c2p3}4(24p2c4+42c4p+21c48p3c354c3p244c3p36c2p412p3c2+27p2c2+8p5c+32cp4+16p6)ε21/{2c4p8(cp)6(p+1)2}{(2p+2)c2(p2+3p)c2p3}4ε1ε2/{(c4p4(cp)3(p+1)}+o(|ε1,ε2|2), (25)
    ϕ2(ε1,ε2)={(2p+2)c2(p2+3p)c2p3}ε1/{2c2(p32cp+p2+c2p+c22cp2)p4}{(2p+2)c2(p2+3p)c2p3}ε2/c2{(2p+2)c2(p2+3p)c2p3}(243p3c3+832p3c4+513p2c4+455p4c3594p5c21347p3c51209p2c5+165p4c4+1138p5c3324p6c2424p7c200p5c4+382p6c3+512p7c2520cp8396c5p48p9+108c648p10+384c6p3+414c6p104cp9+264c2p8+594c6p2672c5p4+96c6p4136c5p544c4p676c3p7)ε21/{4c3(p+1)2(cp)4p6}{(2p+2)c2(p2+3p)c2p3}(8p2c4+23c4p+12c4+30p3c3+8c3p222c3p58c2p485p3c2+6p2c28p5c+46cp4+24p6)ε1ε2/{4c4p2(p+1)(cp)2}+(cp)p2{(2p+2)c2(p2+3p)c2p3}ε22/c4+o(|ε1,ε2|2). (26)

    Then, comparing the coefficients of terms of the same degree in (24), we obtain the second order approximations

    ε1=c2p6(cp)4(p+1)μ1/{(2p+2)c2(p2+3p)c2p3}4+c2p10(cp)6(p+1)(32p2c4+56c4p+27c416p3c379c3p259c3p48c2p419p3c2+36p2c2+12p5c+50cp4+24p6)μ21/{2{(2p+2)c2(p2+3p)c2p3}8}+c2p8(cp)5(p+1)μ1μ2/{(2p+2)c2(p2+3p)c2p3}5+o(|μ1,μ2|2), (27)
    ε2=c2p2(cp)2(8p512cp418cp3+8c3p211p2c29c2p+14c3p+6c3)μ1/{2{(2p+2)c2(p2+3p)c2p3}4}c2μ2/{(2p+2)c2(p2+3p)c2p3}+c2p6(cp)4(1314c7p2+630pc7270p3c4+2068p3c5+612p2c5+677p4c41134p5c3+4387p5c41056p6c31804p7c23741c6p3+756c5p4+1160c3p82268c6p4+1176c7p31272c5p6352c6p5+384c7p4320p11+108c7704cp10+224c2p92046c5p5+4258c4p6+832p7c41464p8c22289c6p2+1544p7c3450c6p1344cp9)μ21/{8{(2p+2)c2(p2+3p)c2p3}8}+c2p4(cp)2(40p2c4+61c4p+24c478p3c3158c3p268c3p14c2p4+43p3c2+48p2c2+32p5c+62cp4+24p6)μ1μ2/{4{(2p+2)c2(p2+3p)c2p3}5}+c2p2(cp)μ22/{(2p+2)c2(p2+3p)c2p3}2+o(|μ1,μ2|2). (28)

    Then we are ready to express those bifurcation curves in parameters ε1 and ε2.

    For curves SN±, we need to consider μ1=0. From the first equality of (19) we see that μ1=0 if and only if ε1=0 because in the expression of ϕ1(ε1,ε2) we have ϕ11(0,0)/ϕ212(0,0)=1/p2c2(p+1)0. Thus, for μ1=0 we obtain from (28) that

    ε2=c2(2p+2)Ψ(c)μ2+O(|μ2|2), (29)

    where Ψ(c) is the same quadratic polynomial as given in (12). It follows that the inequality μ2>0 (or <0) together with the sign of Ψ(c) determines the sign of ε2. From the analysis of the quadratic equation (12) we see that Ψ(c)<0 (or >0) if 0<c<ς(b) (or c>ς(b)), where ς(b) is defined in Lemma 2.2. Hence from (22) we obtain that

    SN+:={(ε1,ε2) | ε1=0,ε2>0,0<c<ς(b)}{(ε1,ε2) | ε1=0,ε2<0,c>ς(b)},SN:={(ε1,ε2) | ε1=0,ε2<0,0<c<ς(b)}{(ε1,ε2) | ε1=0,ε2>0,c>ς(b)}.

    For curve H, we need to consider μ1=μ22, which is equivalent to Υ(ε1,ε2):=ϕ1(ε1,ε2)+ϕ22(ε1,ε2)=0 by (19). Clearly, Υ(0,0)=0 and

    Υε1|(ε1,ε2)=(0,0)={(2p+2)Ψ(c)}4/{p6c2(cp)4(p+1)}0.

    By the Implicit Function Theorem, there exists a unique Ck function ε1=ϵ1(ε2) such that ϵ1(0)=0 and Υ(ϵ1(ε2),ε2)=0. Similarly to (27) and (28), expanding Υ at (ε1,ε2)=(0,0) and substituting with a formal expansion of ϵ1(ε2) of order 2, we obtain by comparison of coefficients that

    ε1=ϵ1(ε2)=p6(cp)44(p+1)Ψ2(c)ε22+o(|ε2|2). (30)

    Further, replacing μ1 with μ1=μ22 in (28), we get

    ε2=c2(2p+2)Ψ(c)μ2+o(|μ2|).

    Similarly to (29), from (22) we obtain that

    H:={(ε1,ε2) | ε1=p6(cp)44(p+1)Ψ2(c)ε22+o(|ε2|2),  ε2>0,0<c<ς(b)}{(ε1,ε2) | ε1=p6(cp)44(p+1)Ψ2(c)ε22+o(|ε2|2),  ε2<0,c>ς(b)}.

    For curve L, we need to consider μ1=4925μ22+o(|μ2|2), i.e., ϕ1(ε1,ε2)=4925ϕ22(ε1,ε2)+o(|ϕ2|2). Similarly to H, we apply the Implicit Function Theorem to obtain

    ε1=49p6(cp)4100(p+1)Ψ2(c)ε22+o(|ε2|2).

    Similarly to (29), from (22) we obtain that

    L:={(ε1,ε2) | ε1=49p6(cp)4100(p+1)Ψ2(c)ε22+o(|ε2|2),  ε2>0,0<c<ς(b)}{(ε1,ε2) | ε1=49p6(cp)4100(p+1)Ψ2(c)ε22+o(|ε2|2),  ε2<0,c>ς(b)}.

    Finally, with the replacement (13) we can rewrite the above bifurcation curves SN±,H and L expressed in parameters (ε1,ε2) in expressions in the original parameters (a,b,c,κ) as shown in Theorem 3.1.


    4. Conclusions

    In this paper we analyzed the dynamics of system (4) near the equilibrium E when parameters lie near BC. We proved that E is a cusp when parameters lie on BC. We investigated the Bogdanov-Takens bifurcation near the cusp and compute in Theorem 3.1 the four bifurcation curves SN+, SN, H and L in the practical parameters. Those bifurcation curves can be observed in Figure 3 in the case that c>1 and b=(c+1)2/4 (which implies p=(c1)/2). They display the merge of equilibria and the rise of homoclinic orbits and periodic orbits.

    Figure 3. Bifurcation diagrams of system (4) for the case that c>1 and b=(c+1)2/4.

    More concretely, in this case,

    a=(c+1)24, κ=8c2(c+1)(c1)2.

    Moreover, the four bifurcation curves divide the neighborhood U of (a,κ) into the following regions:

    DI:={(a,κ)U| a<(c+1)24, κ8c2(c+1)(c1)2}{(a,κ)U| a<(c+1)2449(c1)6(c+1)33200(2c2+c+1)2{κ8c2(c+1)(c1)2}2+O(|κ8c2(c+1)(c1)2|3), κ>8c2(c+1)(c1)2},DII:={(a,κ)U| (c+1)2449(c1)6(c+1)33200(2c2+c+1)2{κ8c2(c+1)(c1)2}2+O(|κ8c2(c+1)(c1)2|3)<a<(c+1)24(c1)6(c+1)3128(2c2+c+1)2{κ8c2(c+1)(c1)2}2+O(|κ8c2(c+1)(c1)2|3), κ>8c2(c+1)(c1)2},DIII:={(a,κ)U| (c+1)24(c1)6(c+1)3128(2c2+c+1)2{κ8c2(c+1)(c1)2}2+O(|κ8c2(c+1)(c1)2|3)<a<(c+1)24, κ>8c2(c+1)(c1)2},DIV:={(a,κ)U| a>(c+1)24}.

    Theorem 3.1 gives dynamical behaviors of system (4) near E in the first quadrant in Table 4. The coordinates of equilibria E0:(x0,0), E1:(p1,q1) and E2:(p2,q2) are given by x0:=a/(b1) and

    p1:=12{(abc+1){(abc+1)24(ac)}1/2},p2:=12{(abc+1)+{(abc+1)24(ac)}1/2}

    as in [27]. E0 exists in the first quadrant when (a,κ)DILDIIHDIII but disappears when (a,κ)DIV (appearing in other quadrants) or (a,κ)SN+{(a,κ)}SN (not existing).

    Table 4. Dynamics of system (4) in various cases of parameter (a,κ).
    Parameters (a,κ) Equilibria Limit cycles and homoclinic orbits Region in bifurcation diagram
    E0 E1 E2 E
    DI saddle unstable focus saddle DI
    L saddle unstable focus saddle one homoclinic rrbit L
    DII saddle unstable focus saddle one limit cycle DII
    H saddle stable focus saddle H
    DIII saddle stable focus saddle DIII
    SN+ saddle-node SN+
    DIV DIV
    (a,κ) cusp (a,κ)
    SN saddle-node SN
     | Show Table
    DownLoad: CSV

    The appearance of limit cycle displays a rise of oscillatory phenomenon in system (4). Choosing parameters a=3.99999,b=4,c=3 and κ=4.495 in DII, we used the command ODE45 in the software Matlab Version R2014a to simulate the orbit initiated from (x0,y0)=(1.00432,1.98662845) numerically, which plots an attractive limit cycle in Figure 4 and shows a dynamic balance and permanence of the substrate and the product in the enzyme-catalyzed reaction. The homoclinic loop actually gives a boundary for the break of the dynamic balance and permanence.

    Figure 4. An attracting limit cycle.

    In this paper we only considered parameters in BC. When parameters lie in C, higher degeneracy may happen at E. Although efforts have been made for higher degeneracies, for example, versal unfolding was discussed in [5] for a normal form of cusp system of codimension 3, it is still difficult to compute bifurcation curves in original parameters in the case of codimension 3. Such a computation with original parameters is indispensible for practical systems and for system (4) it will be our next work.


    Appendix. Some coefficients

    The functions in system (15) are

    E00:={(2p+2)c2(p2+3p)c2p3}4ε1/{c2(p+1)p6(cp)4},E10:={(2p+2)c2(p2+3p)c2p3}2ε1{(6c3p4c3p24p3c2+3p2c2+4cp4+4c4p+3c4)(p2c23c3p3c2p+cp2+2cp32p4)ε1(p3c22cp4+p5+4c2p45p5cp3c3+2p6)ε2}/{(p+1)p4c3(cp)4},E20:={(2c6(p+1)2(cp)2)+(9c3p2+4c2p413c4p+4p5c2+6p3c3+9c5p15p2c42p4c3+4p2c54p3c4+6c5)ε1(2p7c6p7c26p6c22p5c4+6p6c3+2cp82p4c4+6p5c3)ε2+(6p5c22p4c36p6c6p7c+6p6c22p5c3+2p7+2p8)ε1ε2+(6p3c34p2c42c2p410p3c29c4p2cp4+17c3p22p5c+13c3p9p2c26c4)ε21}/{2c3p2(cp)2(p+1)},E01:={(2p+2)c2(p2+3p)c2p3}{2c3ε1+(cp42p3c2+c3p2)ε2+(2p46cp3+4p2c2)ε1ε2+(12c26cp)ε21}/{p2(cp)2c3},E11:={(3c3p28p2c4p4c3+2c5+2c2p4+4c5p+2p2c55c4p+2p5c2+2p3c33p3c4)+(3c2p4+3c3p+p2c2+2p5c+3p2c4+3p3c2+2c4p+2cp44p3c3+c3p2)ε1+(5p6c22p7c3p6c+7p5c2+2c2p4p5c5p4c3p3c3+p3c4+p4c44p5c3)ε2(5p6c4p5c2+p4c35c2p4p3c2+7p5c+p3c3+2cp42p7p53p6)ε1ε2+(13cp28cp4+9c3p238p2c2+5cp3+10p4+10p5+19c3p+10c313p3c225c2p)ε21}/{c2(p+1)(p+c)},E02:={(c2p1)+(5c32c2p)ε1(3p3c22c3p2cp4)ε2+(p4cp3)ε1ε2(2cpc2)ε21}/{(p+1)2(cp)2}.

    The functions below system (18) are

    ϕ11:=24c6p5+4c8p216c7p4+4c8p316c5p6+4p7c4+24c6p416c7p316c5p5+4c4p6+(9p4c416p6c3+40c3p7+68p5c426p3c5+3c86c8p+42c6p3+36c6p494c5p4+6c7p24c4p616c2p856c5p58c8p2+8c7p3+28c6p214c7p)ε1+(4c7p4+40c5p74c2p94c2p10+20c3p8+20c3p920c6p5+40c5p620c6p640c4p8+4c7p540p7c4)ε2(40p2c5+12p4c3+32c7p2+8p5c3+12c7+92p3c5+8p6c232p3c412p6c328p7c2+4c5p488c6p256p4c4+36c7p+48p5c460c6p3+16cp832c6p)ε21+(12cp924p7c48c7p46c7p388c5p532c2p8+20cp1024c5p4+6c6p3+2c3p724p6c3+36c6p5+72c4p6+96c3p876c2p9+36p5c4+40c6p444c5p6+6p7c2)ε1ε2+(8p7c9p2c416p5c2+6p3c3c2p4+11p4c4+6p3c5+10p4c316p5c318p2c5+12p3c49c6p2+4p6c4p8)ε31+(34c4p6+2c3p7+4p928cp916cp8+6p4c4+8p102p7c+32c2p8+32p7c232p6c314p5c3+10p6c26c6p4+26c5p5+12p5c4)ε21ε2+(4c3p7c6p6+44c3p941c2p10c4p6+4cp9+2c5p64p11+28c3p832c2p9+8c5p726c4p8+20p11c12p7c4p104p12+18cp106c2p8)ε1ε22,ϕ12:=(2p5c34p3c4+2p4c3+2p3c5+2p2c54p4c4)+(9c3p2+4c2p413c4p+4p5c2+6p3c3+9c5p15p2c42p4c3+4p2c54p3c4+6c5)ε1+(2p7c+6p7c2+6p6c2+2p5c46p6c32cp8+2p4c46p5c3)ε2+(6p3c34p2c42c2p410p3c29c4p2cp4+17c3p22p5c+13c3p9p2c26c4)ε21+(6p5c22p4c36p6c6p7c+6p6c22p5c3+2p7+2p8)ε1ε2,ϕ21:=(6c10+12c8p5+69c8p477c9p3+20c7p6+9c6p433c7p3+18c5p634c6p5+45c8p226c7p4+102c8p327c9p80c9p2+27c7p5+6c5p7+8c4p812c5p855c6p612c6p7+8c4p9+22c10p224c9p4+8c10p3+20c10p)ε1+(4p10c4+20p9c610p10c54p5c9+2p11c42c9p42p6c9+10p7c8+20p6c8+10c8p520p9c5+2c4p940p7c720c7p810c5p820c7p6+40c6p8+20c6p7)ε2+(12c9+12c3p947c8p4+10c9p386c6p419c7p3+102c5p6220c6p5+60c8p2+159c7p440c8p3+61c5p5+2c4p616p7c418c9p+3c9p2+92c7p5+12c3p8+26c5p714c4p8+53c8p+35c6p376c7p279c6p6)ε21+(2p5c934c8p5+2c3p9+19c8p410c9p3+151c7p617c5p6+39c6p545c7p4+26c8p32c3p10+3p7c46c9p2+23c7p5+77c5p726c4p8+145c5p885c6p6227c6p731c4p92c9p4103c6p8+83p7c7+51p9c52p10c44p11c327p6c8)ε1ε2+(4p7c82p6c830p10c460p11c4+40p9c52p8c8+12p11c3+24c7p8+12c7p960p9c6+12p13c330p10c6+40p11c530p12c44p13c22p14c22p12c2+24p12c330c6p8+80p10c5+12p7c7)ε22+(30c8+69p3c516p4c4212p5c4+58p6c3+331c5p4232c6p4+79c7p3+117c5p665c6p521c8p23c7p4+5c8p3+379c5p5187c4p6+4p7c494c3p8+44c2p8+44c2p9+91c7p53c8p263c6p3+163c7p2106c6p238c3p7)ε31+(36c2p1010c8p5199c3p941c8p4+9c7p6+18c2p11166c6p4+84c7p3+165c5p6297c6p518c8p2+193c7p448c8p3110c3p10+164c5p576c4p6+47p7c4+123c7p578c3p8+18c2p9208c5p7+351c4p8219c5p862c6p6+79c6p7+233c4p9+12c3p7)ε21ε2+(2p14c+c2p104c3p98c7p6+21c2p112p12c72c3p104p13c4c5p7+6c4p8102c5p8+c6p6+45c6p7+118c4p9+58p9c6114p10c5+121p11c4+102c6p822p7c7212p9c5+233p10c4138p11c3+40p12c270p12c3+20p13c214c7p8+p7c8+p6c8)ε1ε22(176p3c4+41p4c3+769p3c5+293p2c5388p4c4+27p5c3+28p6c2178p5c458p6c3+4p7c2+20cp8+20cp9+603c5p4192c6p4+75c7p3+127c5p5+34c4p6+72c7234c6p24c2p8+210c7p616c6p3+213c7p2658c6p244c3p7)ε41+(286p5c4+154p6c332p7c2+136c2p1056cp10198c3p924cp9+262c5p432cp11330c6p4+70c7p3+438c5p6284c6p5+68c7p4+636c5p5580c4p6210p7c4+22c7p5154c3p8+30c2p8+198c2p9+64c5p7+84c4p8122c6p3+24c7p276c6p6+198c3p7)ε31ε2+(4p12+102c2p10cp10158c3p9c7p6+198c2p1164p12c33cp11c5p6313c3p1032p13c+4p7c4+8p136c3p8+4c2p957c5p7+132c4p8126c5p8+10c6p6+26c6p7+272c4p9+4p14+16c6p8p7c770p9c5+144p10c4161p11c3+100p12c2)ε21ε22.

    [1] [ A. Betz,E. Sel'kov, Control of phosphofructokinase [PFK] activity in conditions simulating those of glycolysing yeast extract, FEBS Lett., 3 (1969): 5-9.
    [2] [ S. N. Chow and J. K. Hale, Methods of Bifurcation Theory, Springer, New York, 1982.
    [3] [ F. A. Davidson,J. Liu, Global stability of the attracting set of an enzyme-catalysed reaction system, Math. Comput. Model., 35 (2002): 1467-1481.
    [4] [ F. A. Davidson,R. Xu,J. Liu, Existence and uniqueness of limit cycles in an enzyme-catalysed reaction system, Appl. Math. Comput., 127 (2002): 165-179.
    [5] [ F. Dumortier,R. Roussarie,J. Sotomayor, Generic 3-parameter families of vector fields on the plane, unfolding a singularity with nilpotent linear part. The cusp case of codimension 3, Ergod. Theor. Dyn. Syst., 7 (1987): 375-413.
    [6] [ D. Erle, Nonuniqueness of stable limit cycles in a class of enzyme catalyzed reactions, J. Math. Anal. Appl., 82 (1981): 386-391.
    [7] [ D. Erle,K. H. Mayer,T. Plesser, The existence of stable limit cycles for enzyme catalyzed reactions with positive feedback, Math. Biosci., 44 (1979): 191-208.
    [8] [ A. Goldbeter, Biochemical Oscillations and Cellular Rhythms: The Molecular Bases of Periodic and Chaotic Behaviour, Cambridge University Press, Cambridge, 1996.
    [9] [ A. Goldbeter,G. Dupont, Allosteric regulation, cooperativity and biochemical oscillations, Biophy. Chem., 37 (1990): 341-353.
    [10] [ J. Guckenheimer and P. Holmes, Nonlinear Oscillations Dynamical Systems and Bifurcations of Vector Fields, Springer, New York, 1990.
    [11] [ B. Hassard,K. Jiang, Unfolding a point of degenerate Hopf bifurcation in an enzyme-catalyzed reaction model, SIAM J. Math. Anal., 23 (1992): 1291-1304.
    [12] [ X. Hou,R. Yan,W. Zhang, Bifurcations of a polynomial differential system of degree n in biochemical reactions, Comput. Math. Appl., 43 (2002): 1407-1423.
    [13] [ J. P. Kernévez,G. Joly,M. C. Duban,B. Bunow,D. Thomas, Hysteresis, oscillations, and pattern formation in realistic immmobilized enzyme systems, J. Math. Biol., 7 (1979): 41-56.
    [14] [ Y. A. Kuznetsov, Elements of Applied Bifurcation Theory, Appl. Math. Sci., 112, Springer, New York, 1995.
    [15] [ Z. Leng,B. Gao,Z. Wang, Qualitative analysis of a generalized system of saturated enzyme reaction, Math. Comput. Model., 49 (2009): 556-562.
    [16] [ J. Liu, Coordination restriction of enzyme-catalysed reaction systems as nonlinear dynamical systems, Proc. R. Soc. Lond. A, 455 (1999): 285-298.
    [17] [ A. G. Marangoni, Enzymes Kinetics: A Modern Approach, Wiley-Interscience, Hoboken, NJ, 2003.
    [18] [ L. Michaelis,M. L. Menten, Die kinetik der invertinwirkung, Biochem. Z., 49 (1913): 333-369.
    [19] [ J. D. Murray, Mathematical Biology Ⅰ: An Introduction, Interdisciplinary Applied Mathematics 17, Springer, Berlin, 2002.
    [20] [ H. G. Othmer,J. A. Aldridge, The effects of cell density and metabolite flux on cellular dynamics, J. Math. Biol., 5 (1978): 169-200.
    [21] [ I. Stoleriu,F. A. Davidson,J. Liu, Effects of priodic input on the quasi-steady state assumptions for enzyme-catalyzed reactions, J. Math. Biol., 50 (2005): 115-132.
    [22] [ Y. Tang,D. Huang,W. Zhang, Direct parametric analysis of an enzyme-catalyzed reaction model, IMA J. Appl. Math., 76 (2011): 876-898.
    [23] [ Y. Tang,W. Zhang, Bogdanov-Takens bifurcation of a polynomial differential system in biochemical reaction, Comput. Math. Appl., 48 (2004): 869-883.
    [24] [ R. Varón,M. García-Moreno,F. García-Molina,M. E. Fuentes,E. Arribas,J. M. Yago,M. Ll. Amo-Saus,E. Valero, Two new regulatory properties arising from the transient phase kinetics of monocyclic enzyme cascades, J. Math. Chem., 38 (2005): 437-450.
    [25] [ Y. -Q. Ye et al., Theory of Limit Cycles, Transl. Math. Monogr. 66 American Mathematical Society, Providence, RI, 1986.
    [26] [ Z. -F. Zhang, T. -R. Ding, W. -Z. Huang and Z. -X. Dong, Qualitative Theory of Differential Equations, Transl. Math. Monogr., 101 Amer. Math. Soc., Providence, RI, 1992.
    [27] [ Q. Zhang, L. Liu and W. Zhang, Local bifurcations of the enzyme-catalyzed reaction comprising a branched network, Int. J. Bifur. Chaos, 25 (2015), 155081 (26 pages).
  • This article has been cited by:

    1. Juan Su, Zhaoxia Wang, Global Dynamics of an Enzyme-Catalyzed Reaction System, 2020, 43, 0126-6705, 1919, 10.1007/s40840-019-00780-2
    2. Juan Su, Bifurcation Analysis of an Enzyme Reaction System with General Power of Autocatalysis, 2019, 29, 0218-1274, 1950079, 10.1142/S0218127419500792
    3. Juan Su, Bing Xu, Local bifurcations of an enzyme-catalyzed reaction system with cubic rate law, 2018, 94, 0924-090X, 521, 10.1007/s11071-018-4375-y
  • Reader Comments
  • © 2017 the Author(s), licensee AIMS Press. This is an open access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/4.0)
通讯作者: 陈斌, bchen63@163.com
  • 1. 

    沈阳化工大学材料科学与工程学院 沈阳 110142

  1. 本站搜索
  2. 百度学术搜索
  3. 万方数据库搜索
  4. CNKI搜索

Metrics

Article views(3121) PDF downloads(617) Cited by(3)

Article outline

Figures and Tables

Figures(4)  /  Tables(1)

Other Articles By Authors

/

DownLoad:  Full-Size Img  PowerPoint
Return
Return

Catalog