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Cao2013 Modelo Numerico Del Las Redes de Rulkov
Cao2013 Modelo Numerico Del Las Redes de Rulkov
International Journal of Bifurcation and Chaos, Vol. 23, No. 12 (2013) 1330041 (20 pages)
c World Scientific Publishing Company
DOI: 10.1142/S0218127413300413
Based on the detailed bifurcation analysis and the master stability function, bursting types and
stable domains of the parameter space of the Rulkov map-based neuron network coupled by
the mean field are taken into account. One of our main findings is that besides the square-wave
bursting, there at least exist two kinds of triangle burstings after the mean field coupling, which
can be determined by the crisis bifurcation, the flip bifurcation, and the saddle-node bifurcation.
Under certain coupling conditions, there exists two kinds of striking transitions from the square-
wave bursting (the spiking) to the triangle bursting (the square-wave bursting). Stable domains
of fixed points, periodic solutions, quasiperiodic solutions and their corresponding firing regimes
in the parameter space are presented in a rigorous mathematical way. In particular, as a function
of the intrinsic control parameters of each single neuron and the external coupling strength, a
stable coefficient of the Neimark–Sacker bifurcation is derived in a parameter plane. These
results show that there exist complex dynamics and rich firing regimes in such a simple but
thought-provoking neuron network.
Keywords: Rulkov neuron model; bursting; bifurcation analysis; master stable function; mean
field coupling.
∗
Author for correspondence
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H. Cao & Y. Wu
biological neurons and collective behaviors of made by Rulkov [2001] and de Vries [2001], a
large-scale coupled map-based neuron networks more comprehensive bifurcation analysis is pre-
[Yang & Lu, 2005; Tanaka et al., 2006; Wang et al., sented from a single neuron to N identical neurons
2008; Cao & Sanjuán, 2009; Izhikevich & Edelman, coupled by the mean field. We obtain the analyti-
2008; Cao & Ibarz, 2010]. cal expressions of the flip bifurcation and the cri-
Usually, as a basic neuron model, the chaotic sis bifurcation of the coupled neurons, by which it
Rulkov map-based neuron model receives much is useful to examine different types of bursting of
attention from two aspects: one is the classifica- the coupled neurons in a rigorous way rather than
tion of different kinds of bursting, and the other is only by numerical simulations. Different from other
whether the bursting can be recovered or destroyed works, much attention is paid to possible bifurca-
in a population of such nonbursting or bursting tion conditions leading to other types of bursting
neurons when they are coupled via various net- not only the square-wave bursting.
work connections. Among them, it is well known We find there are at least two kinds of trian-
that the single chaotic Rulkov map-based neuron gle bursting types in the coupled Rulkov map-based
model can produce the square-wave bursting due to neuron model. The first kind of triangle bursting
the bistability and the suitable position of a slow can be determined by a crisis bifurcation, a supflip
nullcline [Rulkov, 2001; de Vries, 2001]. According bifurcation and a saddle-node bifurcation, and the
to the classification of bursting mappings [Izhike- second one can be determined by two saddle-node
vich & Hoppensteadt, 2004], this kind of square- bifurcations, a subflip bifurcation, and a supflip
wave bursting belongs to the fold/homoclinic type, bifurcation. The duration between the active phase
namely, the square-wave bursting can be predicted and the silent phase of the first triangle burst-
by a fold (saddle-node) bifurcation and a crisis ing is the longest. Interestingly, after the mean
(homoclinic) bifurcation [Rulkov, 2001; de Vries, field coupling, we find that there are at least two
2001]. Besides that, few have considered whether kinds of striking transitions: the first transition is
there exist other kinds of bursting. Although de from the square-wave bursting to the triangle burst-
Vries mentioned that other types of bursting like the ing, and the second one is from the spiking of
triangle bursting (tapered type) could exist even in the single neuron to the square-wave bursting after
a single Rulkov map-based neuron model [de Vries, couplings.
2001], there is still a lack of adequate arguments to Secondly, we carry out a comprehensive qualita-
support this possibility. As far as the second aspect tive analysis and a codimension-2 bifurcation anal-
is concerned, Rulkov reported how the synchroniza- ysis for the coupled neurons. The goal is to clarify
tion among chaotically bursting neurons can lead the stable domains in the parameter space among
to the onset of regular bursting by establishing a fixed points, periodic solutions, quasiperiodic solu-
neuron network model with a mean field coupling tions and their corresponding firing regimes of
[Rulkov, 2001]. de Vries revealed that the bursting the coupled neuron network in a two-dimensional
can be recovered in a population of such nonburst- parameter plane. In particular, we make a detailed
ing neurons when they are coupled via the mean computation of a coefficient of the Neimark–Sacker
field, which is called the emergent phenomenon, and bifurcation [Kuznetsov, 1999] in a parameter plane.
this kind of emergent bursting in the network is due To attain this goal, the normal form restricted on
to coupling alone and is very robust to changes in the center manifold is calculated. As a function of
the coupling strength by using the geometric bifur- the intrinsic control parameters of each single neu-
cation analysis [de Vries, 2001]. While, it is still ron and the external coupling strength, the stability
not fully understood in the transition mechanism coefficient demonstrates that there exist different
why the bursting can be recovered or destroyed in a parameter regions where complex dynamics and
population of such nonbursting or bursting neurons rich behaviors in biological neurons arise in the sim-
after couplings. ple chaotic Rulkov neuron network.
Thus, we intend to characterize mathemati- These results show that there exist complex
cally how the transition takes place among differ- dynamics and rich firing regimes in such a sim-
ent kinds of firing regimes with or without the ple but thought-provoking neuron network. On the
mean field coupling. In this paper, first of all, based one hand, these dynamics and firing regimes are
on the previous geometrical bifurcation analysis very useful for simulating collective behaviors in
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large-scale neural networks. On the other hand, Suppose that R(xn ) = xn+1 , then the fixed
these analyses and results can be easily extended points of Eq. (2) satisfy the following equation
to a lot of neuron networks.
The layout of this paper is as follows. In Sec. 2, R(xn ) = f (xn ) + γ + εxn = xn . (4)
we give a description of the chaotic Rulkov neuron
map with the mean field coupling. Bifurcation of the 3. Bifurcation of the Single
single parameter is presented in Sec. 3. Sections 4 Parameter
and 5 discuss firing regimes of the single neuron,
and the coupled neurons with the mean field cou- 3.1. Saddle-node bifurcation
pling, respectively. Stability analysis of N identi- Assume that there exists a bifurcation parameter
cal Rulkov neurons with the mean field coupling γ0 and a fixed point x0 satisfying the following four
includes the Neimark–Sacker bifurcation that is dis- conditions:
cussed in Sec. 6. Finally, we sum up our results with
some comments in Sec. 7. (i) R(x0 ) = x0 ;
(ii) ∂R
∂x (x0 , γ0 ) = 1;
2. Model Description ∂R
(iii) ∂γ (x0 , γ0 ) = 0;
We consider the following neuron network with the ∂2R
(iv) ∂x2 (x0 , γ0 ) = 0,
mean field coupling [Rulkov, 2001; de Vries, 2001]:
then a saddle-node bifurcation will occur [Wiggins,
N
xn+1,i = f (xn,i ) + yn,i + ε
xn,j ,
1990].
N (1) From Eq. (4), it is easy to know that ∂R∂γ = 1
j=1
for any√x and γ. There exist two zero points at
y 2 −1)
x = ± 33 such that ∂∂xR2 = 2α(3x
2
n+1,i = yn,i − η(xn,i − σi ), = 0, while
(1+x2 )3
where f (xn,i ) = α/(1 + x2n,i ), the additional sub- the two zero points are not roots of the equation
∂R −2αx
script i indicates the ith neuron, xn,i and yn,i repre- ∂x = (1+x2 )2 + ε = 1.
∂R
sent the transmembrane voltage and the slow gating It is noted that if x < 0, then ∂x > 0. Other-
2
process of the ith neuron, respectively, n is the dis- wise, if x > 0, then ∂R
= 2α(3x
< 0. From −1)
∂2R
2 3 ,
∂x
√ (1+x )∂x2
crete time (n = 1, 2, . . .). N is the total number
we have the following conclusion: if x > 3/3, then
of neurons, α, σ, and η are control parameters of
R√(x) > 0, and√R (x) increases monotonously; if
the single neuron, in which α and σ are O(1), while
− 3/3 < x < 3/3, then R (x) < 0,√and R (x)
0 < η 1. ε is the strength of the coupling. Due
decreases monotonously; when x < − 3/3, then
to 0 < η 1, xn,i is called the fast variable, while
R (x) √
> 0 and R (x) increases monotonously; if
yn,i is referred to as the slow variable.
x√= − 3/3, then R (x) takes
√ the maximum value
A synchronized manifold of Eqs. (1) is defined
3 3α/8 + ε, while if√x = 3/3, then R (x) take the
by Π = {(x, y) | x = x1 = x2 = · · · = xN = σ, y =
minimum value −3 3α/8 + ε. Therefore, we have
y1 = y2 = · · · = yN = σ − f (σ) − εσ}. Then,
the following conclusion.
the dynamics of Eqs. (1) restricted on the synchro-
√
nized manifold Π can be described by the follow- Proposition 1. If α ≥ (1 − ε)8 3/9, then there
ing two-dimensional system [Rulkov, 2001; de Vries, exists a saddle-node bifurcation.
2001]:
α √ R (x) is a continuous function of
In addition,
x, so if α > 8 3/9
xn+1 =
1 + x2n
+ yn + εxn ,
√ + ε, then the maximum value
(2) Max{R (x)} = 3 3α/8 + ε > 1. There exist two
yn+1 = yn − η(xn − σ). pairs
√ of points
√ (x01 , γ) and (x02 , γ), which x01 <
− 3/3, − 3/3 < x02 < 0, such that R (x) = 1,
By using the fast–slow decomposition technique then we have the following stability conditions.
[Rinzel, 1985, 1987], Eqs. (2) can be further reduced
into the following one-dimensional fast subsystem if Proposition 2
we suppose that yn = γ:
(i) If x < x01 , then 0 < R (x) < 1, which means
xn+1 = f (xn ) + γ + εxn . (3) that the fixed points of Eq. (2 ) are stable;
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H. Cao & Y. Wu
(ii) If x01 < x < x02 , then R (x) > 1, which means ∂R −2αx
that the fixed points of Eq. (2 ) are unstable; ∂x = (1 + x2 )2 + ε = 1,
(iii) If x02 < x < 0, then 0 < R (x) < 1, and the
α
fixed points of Eq. (2 ) are stable.
x = + γ + εx,
1 + x2
By solving the first two equations given in the
conditions of the saddle-node bifurcation the analytical expression of the saddle-node bifur-
cation is obtained by Rulkov [2001] as follows:
−18γ(1 − ε)2 − 2γ 3 ± [2γ 2 − 6(1 − ε)2 ] γ 2 − 3(1 − ε)2
α= . (5)
27(1 − ε)2
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In particular, when ε = 0, the analytical expres- the flip bifurcation (the dashed line), respectively.
sion of the crisis bifurcation is obtained by Rulkov The range of α is divided into the following four
[2001] as: intervals by four different values of α1,2,3,4 .
−3γ ± γ 2 − 8 Proposition 6
α= . (11)
2 √
(i) When 8 3/9 < α < 2.58, there coexist two
To sum up the above bifurcation analyses, we have saddle-node bifurcation values γsn1,sn2 , and
the following conclusion: two flip bifurcation values γf p1,f p2 , and their
Proposition 5
order is γsn1 < γsn2 < γf p1 < γf p2 ;
√ (ii) When 2.58 < α < 4, there coexist two saddle-
(i) If (1 − ε)8 3/9 < α < 4 − 2ε, then there node bifurcation values γsn1,sn2 , and two flip
coexist the saddle-node bifurcation and the flip bifurcation values γf p1,f p2 , their order becomes
bifurcation; √ γsn1 < γf p1 < γsn2 < √ γf p2 ;
(ii) If 4 − 2ε < α < (1 + ε)8 3/3, then there (iii) When 4 < α < 8 3/3, there coexist two
coexist the saddle-node bifurcation, the flip saddle-node bifurcation values γsn1,sn2 , two flip
bifurcation, and the
√ crisis bifurcation; bifurcation values γf p1,f p2 , and two crisis bifur-
(iii) If α = (1 + ε)8 3/3, then the saddle-node cation values γcs1,2 , and their order is γsn1 <
bifurcation is equal to the crisis bifurcation. γf p1 < γcs1 < γcs2 < γsn2 < γf p2 .
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H. Cao & Y. Wu
2 5
1.5 4
xmax
3
1
γfp2 xmax
2
0.5
1 γfp1 γfp2
γfp1
xn
xn
0
xmin 0
γsn1
−0.5
−1
xmin
−1
−2
γ γ
sn2 sn2
−1.5 −3
γsn1
−2 −4
−2.5 −2 −1.5 −1 −0.5 0 0.5 1 −5 −4 −3 −2 −1 0 1
γ γ
(a) (b)
5 5
4 4
xmax
x
max
3 3
2 2
γ γfp1
xn
xn
(c) (d)
Fig. 2. Bifurcation diagrams of Eqs. (2) with ε = 0, σ = −1, η = 0.001, and (a) α = 2, (b) α = 3.9, (c) α = 4.15 and
(d) α = 4.7.
saddle-node bifurcation γsn2 and the√ flip bifurca- In this interval of α, since there is no crisis bifurca-
tion γf p1 intersect at α = 87 13 16 2
≈ 2.58. By tion, xn moves between the maximum values and
7 + 7
Proposition 6, when 2.58 < α < 4, there exist four the minimum values of xn . Therefore, the single
bifurcation values, and their relative positions are neuron gives rise to the triangle bursting. Once xn
γsn1 < γf p1 < γsn2 < γf p2 . Seen from Fig. 2(b), dur- moves toward left along the branch of fixed points
ing the silent phase, if iterates of the map are on or and falls to the lower stable branch of fixed points
near the stable fixed points on the bottom branch of at γsn1 , the single neuron becomes the silent state
the S-shaped curve, and all of them are below the again. To sum up, when 2.58 < α < 4, the single
y-nullcline xn = σ = −1, then yn increases slowly. neuron produces the triangle bursting between γsn1
When yn moves to γsn2 , namely, when yn > γsn2 , and γsn2 . This is a so-called fold-flip type bursting
then the single neuron gives rise to the active phase. [Izhikevich & Hoppensteadt, 2004]. The correspond-
During the active phase, because iterates lie above ing time evolution and the phase portrait of the
xn = σ = −1 on average, yn decreases slowly. tapered bursting are shown in Figs. 3(c) and 3(d).
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4 −0.95
xn
xn
0 −1
−4 −1.05
0 5000 10000 −2.004 −2 −1.996
n yn
(a) (b)
4 5
xn
xn
0 0
−4 −5
0 5000 10000 −5 −4 −3 −2
n yn
(c) (d)
4 5
xn
xn
0 0
−4 −5
0 5000 10000 −3 −2.8 −2.6
n yn
(e) (f)
4 5
xn
xn
0 0
−4 −5
0 5000 10000 −3 −2.9 −2.8
n yn
(g) (h)
Fig. 3. Time evolutions and phase portraits of Eqs. (2) with ε = 0, σ = −1, η = 0.001, and (a, e) α = 2, (b, f) α = 3.9,
(c, g) α = 4.15 and (d, h) α = 4.7.
In addition, it is noted that the duration of the a periodic way. It is the so-called homoclinic/fold
tapered bursting is decided by the distance ∆γ1 = type of the square-wave bursting [Izhikevich & Hop-
|γsn2 − γsn1 |. Figure 4(a) shows that the duration of pensteadt, 2004]. The corresponding time evolution
the tapered bursting increases as α increases. and the phase portrait of the square-wave burst-
ing are shown in Figs. 3(e) and 3(f). The duration
4.3. Square-wave bursting of square-wave bursting is decided by the distance
√ ∆γ2 = γsn2 − γcs2 . Contrary to the triangle burst-
When 4 < α < 8 3/3, there coexist the saddle- ing, Fig. 4(b) demonstrates that the duration of the
node bifurcation, the flip bifurcation and the crisis square-wave bursting decreases with the increasing
bifurcation. The corresponding bifurcation diagram of α.
is shown in Fig. 2(c). During the silent phase, iter-
ates of Eq. (2) changes in a very similar way as
the triangle bursting. When yn > γsn1 , the single 4.4. Spiking
√
neuron changes to the active phase. While, dur- When α > 8 3/3, there exist still two saddle-
ing the active phase, because iterates of Eq. (2) lie node bifurcations γsn1,sn2 , and two flip bifurcations
above xn = σ on average, yn decreases slowly. At γf p1,f p2 , but there exists only a crisis bifurcation
the right crisis bifurcation γcs2 , the minimum val- γcs1 as shown in Fig. 2(d). In this situation, both
ues of xn intersect with the middle branch of the the fixed points on the bottom and middle branches
unstable fixed point, the single neuron backs to the of the S-shaped curve are located within the chaotic
silent phase. It leads to the square-wave bursting in attractor. When yn > γsn2 , the single neuron goes
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H. Cao & Y. Wu
1.4 0.07
1.3
0.06
1.2
0.05
1.1
1 0.04
∆γ
∆γ
0.9 0.03
0.8
0.02
0.7
0.01
0.6
0.5 0
2.5 3 3.5 4 4.2 4.3 4.4 4.5 4.6
α α
(a) (b)
Fig. 4. The duration of the bursting when α is varied, where ε = 0, σ = −1, and η = 0.001: (a) The duration of the triangle
bursting and (b) the duration of the square-wave bursting.
into the chaotic oscillation and moves right. When parameter of Eq. (2) corresponding to two pairs of
the average values of xn are below xn = σ = −1, parameter values ε = 0.2, α = 3.8, and ε = 0.4,
iterates of Eq. (2) increase slowly and move right. α = 4.15, respectively.
Once iterates of Eq. (2) lie above xn = σ on aver- Seen from Figs. 5(a)–5(d), there coexist two flip
age, then iterates of Eq. (2) decrease and the single bifurcation points located at the upper branch of
neuron moves left. So the single neuron produces the fixed point curve. When xn moves along the
the spiking regime. The corresponding time evolu- lower stable branch of fixed points, the coupled neu-
tion and phase portrait of spiking regime are shown ron is in the resting state, and yn increases slowly.
in Figs. 3(g) and 3(h). When it moves past the right saddle-node bifurca-
tion γsn1 , the fixed point becomes unstable, and xn
moves to the vicinity of the upper branch of fixed
5. Firing Regimes of the Coupled
points between γf p1 and γf p2 . Since there is no crisis
Neurons with Mean Field bifurcation, xn moves between the maximum value
Coupling and the minimum value of iterates of xn . In this
In this section, we further explore the mechanism process, the coupled neuron gives rise to the trian-
responsible for the emergent phenomenon [de Vries, gle bifurcation. Once xn moves along the curve of
2001], and see what effect will take place after the fixed point and falls to the lower stable branch of
mean field coupling. fixed points at γsn2 , the coupled neuron goes back
to the resting state. So this is the mechanism for
the triangle bursting.
5.1. Triangle bursting Figure 5(e) shows time evolutions from n =
From Proposition 6, when 2.58 < α < 4, a single 1, . . . , 20 000 when ε = 0 to n = 20 001, . . . , 30 000
neuron can give rise to the triangle bursting, which when ε = 0.2. In this case, the single neuron and the
is determined by the saddle-node bifurcation and coupled neurons are both in the triangle bursting
the flip bifurcation. So when the coupled neurons regime, but the size of the triangle bursting is obvi-
have the saddle-node bifurcation and the flip bifur- ously larger than that of the single neuron; while in
cation, can the coupled neurons also produce the Fig. 5(f), the transition takes place from the square-
triangle bursting? In Figs. 5(a)–5(d), we give bifur- wave bursting of the single neuron with ε = 0
cation diagrams of the two-parameter and the single and α = 4.15 for n = 1, . . . , 20 000 to the triangle
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bursting of the coupled bursting with ε = 0.4 and square-wave bursting, which is determined by the
α = 4.15 for n = 20 001, . . . , 30 000. saddle-node bifurcation and the crisis bifurcation.
To sum up, there exists the transition from the So what effect will take place if the coupled neurons
square-wave bursting to the triangle bursting after have also the crisis bifurcation and the saddle-node
the mean field coupling. bifurcation?
It can be seen from Figs. 6(a)–6(d), there coex-
ist six bifurcations values γsn1,sn2 , γf p1,f p2 , and
5.2. Square-wave bursting γcs1,cs2 , with the increase of the parameter α,
√ previous bifurcation analysis, when 4 <
From the γcs2 → γsn2 . Based on these bifurcation analyses,
α < 8 3/3, a single neuron will give rise to the two arrays of time evolutions are presented in
10 3
9
γ 2
γ sn
8 cs
1 γ γfp2
7
γ γsn1 fp1
fp
6 0
5
xn
−1
α
4
3 −2
γ
2 sn2
−3
1
−4
0
−1 −5
−5 −4 −3 −2 −1 0 1 −5 −4 −3 −2 −1 0 1
γ γ
(a) (b)
10 3
γ
9 cs γ 2
sn
8
γfp 1 γfp1
γ
7 fp2
0
6 γsn1
−1
5
xn
−2
α
4
3
−3 γsn2
−4
2
1 −5
0 −6
−1 −7
−5 −4 −3 −2 −1 −5 −4 −3 −2 −1 0 1
γ γ
(c) (d)
Fig. 5. (a) The two-parameter bifurcation diagram for the coupled neurons with ε = 0.2, α = 3.8, (b) when α = 3.8, the
corresponding single parameter bifurcation diagram for the coupled neurons with ε = 0.2, (c) the two-parameter bifurcation
diagram for the coupled neurons with ε = 0.4, α = 4.15, (d) when α = 4.15, the corresponding single parameter bifurcation
diagram for the coupled neurons with ε = 0.4, (e) time evolutions from n = 1, . . . , 20 000 when ε = 0 to n = 20 001, . . . , 30 000
when ε = 0.2. In this case, the single neuron and the coupled neurons are both in the triangle bursting regimes and (f) the
transition from the square-wave bursting of the single neuron with ε = 0 and α = 4.15 for n = 1, . . . , 20 000 to the triangle
bursting of the coupled bursting with ε = 0.4 and α = 4.15 for n = 20 001, . . . , 30 000.
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H. Cao & Y. Wu
6 6
4 4
2 2
xn
xn
0 0
−2 −2
−4 −4
−6 −6
(e) (f)
Fig. 5. (Continued)
Figs. 6(e) and 6(f), respectively. Seen from Fig. 6(e), square-wave bursting of the coupled bursting with
the square-wave bursting of the single neuron with ε = 0.2 and α = 4.7.
ε = 0 and α = 4.4 can be continually maintained In addition, the distance between γcs2 and γsn2
after the coupling with ε = 0.2 and α = 4.4. While, is obtained by numerical simulations shown in Fig. 7
there exists another transition from the spiking of corresponding to α = 4.4, while ε is varied, in which
the single neuron with ε = 0 and α = 4.7 to the the crisis bifurcation γcs2 is always less than the
10
3
9
γ
8 sn
2
7 γcs
γfp 1 γfp2
6 γ
fp1
5 0
xn
γsn1
α
4
−1 γ
cs1
3
γcs2
2 −2
1 γ
−3 sn2
0
−1 −4
−5 −4 −3 −2 −1 0 1 −5 −4 −3 −2 −1 0 1
γ γ
(a) (b)
Fig. 6. (a) The two-parameter bifurcation diagram for the coupled neurons with ε = 0.2, α = 4.4, (b) when α = 4.4, the
corresponding single parameter bifurcation diagram for the coupled neurons with ε = 0.2, (c) the two-parameter bifurcation
diagram for the coupled neurons with ε = 0.2, α = 4.7, (d) when α = 4.4, the corresponding single parameter bifurcation
diagram for the coupled neurons with ε = 0.2, (e) the square-wave bursting of the single neuron with ε = 0 and α = 4.4 for
n = 1, . . . , 20 000 is kept after the coupling with ε = 0.2 and α = 4.4 for n = 20 001, . . . , 30 000 and (f) the transition from the
spiking of the single neuron with ε = 0 and α = 4.7 for n = 1, . . . , 20 000 to the square-wave bursting of the coupled bursting
with ε = 0.2 and α = 4.7 for n = 20 001, . . . , 30 000.
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10 4
9
3
8
7 2
γfp2
6 1 γfp1
5
xn
0
α
4 γsn1
−1
γcs1
3
2 −2 γsn2
1
−3
0 γ
cs2
−1 −4
−5 −4 −3 −2 −1 0 1 −5 −4 −3 −2 −1 0 1
γ γ
(c) (d)
3 3
2 2
1 1
xn
xn
0 0
−1 −1
−2 −2
−3 −3
1 1.5 2 2.5 3 1 1.5 2 2.5 3
n x 10
4 n x 10
4
(e) (f)
Fig. 6. (Continued)
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H. Cao & Y. Wu
55
50 γfp1
γ
0 fp2
45
γsn1
40
γ
fp
35 −5 γ
sn2
30
n
α
x
25 −10
20 γ
γcs sn
15
−15
10
5
−20
0
−5 −4 −3 −2 −1 0 −5 −4 −3 −2 −1 0
γ γ
(a) (b)
2 γ (d)
0
fp1
0 γfp2 −5
γsn1
n
x
−2 −10
γsn2 −15
−4
−20
−6 0 1 2 3 4
n 4
xn
x 10
−8
−10 (e)
0
−12
−5
n
−14
x
−10
−16
−15
−5 −4 −3 −2 −1 0
γ 0 1 2 3 4
n 4
x 10
(c)
Fig. 8. (a) The two-parameter bifurcation diagram for the coupled neurons with ε = 0.8, the corresponding single parameter
bifurcation diagrams and time evolutions for the coupled neurons with (b) ε = 0.8, α = 4.7, (c) ε = 0.8, α = 4.15, (d) ε = 0.8,
α = 4.7 and (e) ε = 0.8, α = 4.15.
Seen from Figs. 8(a)–8(c), γsn2 > γf p2 . The that the duty cycle of the first type of the triangle
square-wave bursting of the single neuron may pro- bursting is the longest one among any type of burst-
duce the first kind of triangle bursting after cou- ing shown in paper.
pling shown in Fig. 8(d), in which the active phase
of bursting is determined by γf p1 and γf p2 , while
the silent phase of bursting is determined by γsn1 5.4. Spiking
√
and γsn2 . Interestingly, there exists another kind of From Fig. 2, when α > 8 3/3, a single neuron
triangle bursting shown in Fig. 8(e), where it means √ produce the spiking. In particular, when α =
will
that the spiking neuron can produce the second kind 8 3/3, the saddle-node bifurcation γsn2 intersects
of triangle bursting after coupling, whose character- √ crisis bifurcation γcs2 . Therefore, when
with the
istic is the active phase of bursting determined by α > 8 3/3, or when the saddle-node bifurcation
γcs2 and γf p2 , while the silent phase of bursting is γsn1 intersects with the crisis bifurcation γcs1 , the
determined by γcs2 and γsn2 . In addition, it is noted single neuron may give rise to the spiking. Under
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5.5 3
γcs
5
2
4.5
1
4 γ
γ fp1
sn 0
3.5
xn
γsn1
α
3 −1 γcs1
γ
2.5 fp
−2
γsn2
2
−3
1.5
1 −4
−5 −4 −3 −2 −1 0 1 2 −5 −4 −3 −2 −1 0 1
γ γ
(a) (b)
3 (d)
2
2 xn
0
1
γfp1 −2
0 1 1.5 2 2.5 3
n
γ
4
xn
x 10
−1
sn1 γ
cs1
(e)
−2 2
γ
sn2
xn
−3 0
−4 −2
−5 −4 −3 −2 −1 0 1
γ 1 1.5 2 2.5 3
n 4
x 10
(c)
Fig. 9. (a) The two-parameter bifurcation diagram for the coupled neurons with ε = −0.2, α = 4.15, (b) the correspond-
ing single parameter bifurcation diagrams for the coupled neurons with α = 4.15, ε = −0.2, (c) α = 4.7, ε = −0.2, time
evolutions: (d) α = 4.15 and ε = 0 for n = 1, . . . , 20 000, and ε = −0.2 for n = 20 001, . . . , 30 000. In this case, the single
neuron without couplings and the coupled neurons are both in the square-wave bursting regimes and (e) α = 4.7 and ε = 0 for
n = 1, . . . , 20 000, and ε = −0.2 for n = 20 001, . . . , 30 000. In this case, the single neuron without couplings and the coupled
neurons are both in the spiking regime.
the effect of the mean field coupling, what will hap- single neuron without couplings can also become
pen when γcs2 ≈ γsn2 or when γcs2 disappears? the spiking after the negative coupling.
In Figs. 9(a)–9(e), we give the two-parameter
bifurcation diagram, the corresponding single
6. Stability Analysis of N Identical
parameter bifurcation diagrams, and time evolu-
tions with α = 4.15, ε = −0.2 and α = 4.7,
Rulkov Neurons with Mean Field
ε = −0.2, respectively. Seen from Fig. 9(d), the Coupling
square-wave bursting of the single neuron without In this section, we discuss the stability condition
couplings is still maintained after the negative cou- of Eq. (1), the corresponding variational equation
pling. While, seen from Fig. 9(e), the spiking of the of Eqs. (1) restricted to Π is δxn+1 = Jδxn . Here,
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H. Cao & Y. Wu
the Jacobian matrix J of Eqs. (1) at a fixed point and ∆ = (t + 1 − ε)2 − 4η, then Eq. (15) becomes
Φ(x, y) on Π for 1 ≤ i ≤ N can be written
1 − t + ε ± (t + 1 − ε)2 − 4η
J = IN ⊗ F (x) + εG ⊗ H, (12) λN ± = . (16)
2
where IN is a N × N identity matrix, We will take into account the conditions such
that |λN ± | < 1 according to the sign of ∆,
f (x) 1 1 0 respectively.
F (x) = , H= ,
−η 1 0 0
where sk = 0 (k = 1, . . . , N − 1), sN = 1 are the From Proposition 7, four bifurcation curves can
eigenvalues of G. Then, the eigenvalues of the com- be obtained by the following mathematical analyt-
plete system of Eqs. (1) are now obtained from each ical expressions.
block at xi = σ in Eqs. (1) as follows: √ (1 + σ 2 )2
α1,2 = (±2 η − 1 + ε) ,
2σ
f (σ) + 1 + εsk ± [f (σ) − 1 + εsk ]2 − 4η
λk± = . (1 + σ 2 )2
2 α3 = (1 + 0.5η + ε) , (17)
(15) 2σ
(1 + σ 2 )2
The stability of the fixed point on Π requires α4 = ( 1 + 2η + ε) .
2σ
that |λk± | < 1. Thus, as a function of the maximum
absolute value of the eigenvalues of Mk , neuron It is noted that αi (i = 1, . . . , 4) are odd func-
parameters α, σ, η, the coupling strength ε, and the tions about σ in the significant domain of σ, and
eigenvalues sk of the coupling matrix G, the mas- their geometrical presentations of αi are symmetri-
ter stability function [Pecora & Carroll, 1998] deter- cal about the origin O(0, 0) in the parametric plane
mine the stability of the neuron network around the (σ, α). Therefore, in the following discussion, we
fixed point. only consider the left-half plane of the parameter
Due to the fact that s1 = s2 = · · · = sN −1 = 0, plane (σ, α).
and sN = 1, therefore, the stability of the fixed The parameter space (σ, α) is divided into three
point on Π is mainly determined by |λN ± | < 1. domains by α1 , α2 , σ-axis, and α-axis as shown in
Therefore, in the following part of this section, we Fig. 10. The three domains in the parameter plane
will focus on the dynamics of the fixed points of (σ, α) are entitled “Sink”, “Source”, and “Saddle”
Eqs. (2). as shown in Figs. 10(a) (ε = 0) and 10(b) (ε = 0.4),
For the sake of simplicity, in the following part respectively, which visualize stable sinks, unstable
of this section, we suppose that t = 2ασ/(1 + σ 2 )2 , sources, and unstable saddles when the parameter
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5 5
4 Source 4
Source
3 α 3
1
2 α2 2
1 α3 1 α1
α4 Sink α2
0 0
α
α
α3
−1 −1 Sink
α4
−2 −2
−3 Saddle −3
−4 −4 Saddle
−5 −5
−2 −1.5 −1 −0.5 0 −2 −1.5 −1 −0.5 0
σ σ
(a) (b)
Fig. 10. When ∆ > 0, fixed points and their types in the parameter plane (σ, α): (a) ε = 0 and (b) ε = 0.4.
values of (σ, α) are located at the three different Eqs. (2) as λ1,2 = [1 − t + ε ± i 4η − (t + 1 − ε)2 ]/
subregions, respectively. It is noted that ∆ < 0 2, then we have the following conclusion.
when the pair of parameter values (σ, α) is located
on the region between α1 and α2 , in which some Proposition 9
very rich nonlinear dynamical phenomena such as
√
the saddle-node bifurcation, the Neimark–Sacker (i) If η − 1 + ε < t < 2 η − 1 + ε, then the fixed
bifurcation may occur. Apart from this region where point Φ is a stable focus;
√
∆ < 0, most parameter values in the (σ, α) plane (ii) If −2 η − 1 + ε < t < η − 1 + ε, then the fixed
correspond to ∆ ≥ 0. In particular, when the pair point Φ is an unstable focus.
of parameter values (σ, α) is chosen on the bound-
aries of α1,2 , then ∆ = 0. In addition, it is hard From Proposition 9, the parameter region
to distinguish α4 from α3 due to α3 ≈ α4 when where ∆ < 0 is divided into two different subre-
0 < η 1. gions by α9 , where α5 is in fact the Neimark–Sacker
bifurcation line defined by
6.2. ∆ = 0
(1 + σ 2 )2
When ∆ = 0, i.e. 4η = (t + 1 − ε)2 ,
then the pair α5 = (η − 1 + ε) . (18)
2σ
of parameter values (σ, α) is just located on α1,2 ,
on which there exist two identical real eigenvalues As shown in Fig. 11(a), when the pair of para-
of Eqs. (2) as λN ± = (1 − t + ε)/2. We obtain the metric values of (σ, α) is chosen in the light gray
following conclusion: blue region enclosed by α1 and α5 , then Φ is a sta-
ble focus, while in the dark gray region enclosed by
Proposition 8 α5 and α1 , Φ is a unstable focus. Their correspond-
(i) Φ is a stable sink if the pair of parameter values ing time evolutions and phase portraits are given in
(σ, α) is located on α1 ; Figs. 11(b)–11(e).
(ii) Φ is a unstable source when the pair of param-
eter values (σ, α) is just located on α2 .
6.4. Neimark–Sacker bifurcation
When ∆ < 0, namely, 4η > (1 + t − ε)2 , then there
6.3. ∆ < 0 exists a pair of conjugate complex eigenvalues λN ±
Under this condition, i.e. 4η > (1 + t − ε)2 , there of Eqs. (2). The Neimark–Sacker (NS) bifurcation
exists a pair of conjugate complex eigenvalues of will occur [Wiggins, 1990; Kuznetsov, 1999] if the
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December 23, 2013 18:44 WSPC/S0218-1274 1330041
H. Cao & Y. Wu
1
xn
−1 −1
and the pair of conjugate complex roots can be
−2
−2
0 5000 10000 −1.4 −1.3 −1.2 rewritten as λN ± = (2 − η ± i 4η − η 2 )/2.
yn
n In a very straightforward way, it is easy to prove
(b) (c) that λnN ± = 1 (n = 1, 2, 3, 4). In the following, we
will determine the coefficient of the normal form
1 derived on the center manifold.
1
At first, the nonzero fixed point Φ(σ, σ − f (σ)−
0 0 εσ) on the synchronized manifold Π can be trans-
formed into the origin O(0, 0) by the following
xn
xn
−1 −1 transformation
−2 −2
−4 −2 0 un+1 = xn+1 − σ,
0 5000 10000 (20)
yn
n
vn+1 = yn+1 − σ + f (σ) + εσ.
(d) (e)
Fig. 11. (a) When ∆ < 0, the parameter domain of (σ, α) in Then Eqs. (2) becomes
the light gray region corresponds to stable foci, while in the
dark gray region corresponds to unstable foci. Time evolu-
α α
tions and phase portraits of Eqs. (2) when ε = 0.4, σ = −1,
un+1 = εun + vn + 2
− ,
1 + (un + σ) 1 + σ2
η = 0.001: (b)–(c) in a limit cycle state with α = 1.3 and
(d)–(e) in a silent state with α = 1.15.
vn+1 = vn − ηun .
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December 23, 2013 18:44 WSPC/S0218-1274 1330041
2ασ α(1 − 3σ 2 ) 2 4ασ(1 − σ 2 ) 3
u ε− u+v− u + u + O(4)
→
(1 + σ 2 )2 (1 + σ 2 )3 (1 + σ 2 )4 .
(22)
v
−ηu + v
Secondly, there exists a matrix M , such that
the Jacobian matrix J can be transformed into the Thirdly, Eqs. (26) can be reduced into the following
following form: one-dimensional complex map after letting:
2 z 1 i x
1− η −
4η − η = , (28)
2 2 z 1 −i y
M −1 J M = , (23)
then we have
4η − η 2 η
1−
2 2 zn+1 = (Re λ + i Im λ)zn + m + ig. (29)
where Fourthly, expanding m + ig in a Taylor expansion
with zn and zn , we obtain the following normal form
1 0
zn+1 = λ(η)zn + ξ20 z 2n + ξ11 zn zn
M =
η 4η −
η 2 ,
−
2 2 + ξ02 z 2n + ξ21 z 2n zn + · · · , (30)
(24)
where the bar of zn denotes the conjugate complex
1 0
variable of zn , and
M −1 =
η 2 .
− ξ20 =
1
[m − myy + 2g xy
4η − η 2 4η − η 2 8 xx
Letting + i(g xx − g yy − 2mxy )](0,0) ,
u x 1
→M , (25) ξ11 = [m + myy + i(g xx + g yy )](0,0) ,
v y 4 xx
1 (31)
and substituting M and M −1 into Map. (22), then ξ02 = [m − myy − 2g xy
8 xx
we have
+ i(g xx − g yy + 2mxy )](0,0) ,
2
1− η −
4η − η
xn+1 xn ξ21 =
1
[m + m
xyy + g xxy + g yyy
2 2
→ 16 xxx
yn+1 yn
4η − η 2 η + i(g
1− xxx + g xyy − mxxy − myyy )](0,0) ,
2 2
where the superscript and the subscript denote the
m second or the third derivative of m or g about x,
+ , (26) and y, respectively.
g
We have
where −2α + 6ασ
mxx (0, 0) = = a1 ,
α 2ασ (1 + σ 2 )3
m=− + − ε xn
1 + σ2 (1 + σ 2 )2 η
g xx (0, 0) = a1 = b1 ,
α 4η − η 2
+ , (27)
1 + (xn + σ)2
mxy (0, 0) = gxy (0, 0) = myy (0, 0)
η
g= f.
4η − η 2 = gyy (0, 0) = 0,
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H. Cao & Y. Wu
3(a21+ b21 ) 1 θ0 θ0
=− + csc 2b2 cos
64 64 2 2
θ0 3θ0
− 2a2 sin − 2(2a1 b1 + b2 ) cos
2 2
−5
3θ0 5θ0 −5 0 5
+ 2(a21 b21
− + a2 ) sin + 2a1 b1 cos σ
2 2
Fig. 12. A contour plot of the stable coefficient a(0) as a
5θ0
− (a21 − b21 ) sin . (37) function of σ and α in the range of (σ, α) ∈ [−5, 5] × [−5, 5]
2 for ε = 0.4.
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December 23, 2013 18:44 WSPC/S0218-1274 1330041
(a) (b)
Fig. 13. Isoperiodic diagrams in the range of (σ, α) ∈ [−1.5, −0.1] × [−3, 3]. (a) ε = 0 and (b) ε = 0.4.
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December 23, 2013 18:44 WSPC/S0218-1274 1330041
H. Cao & Y. Wu
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