
The central nervous system extensively expresses Ca2+-stimulated K+ channels, which serve to use Ca2+ to control their opening and closing. In this study, we explore the numerical computation of Hopf bifurcation in the Chay model based on the equilibrium point's stability and the center manifold theorem to illustrate the emergence of complicated neuronal bursting induced by variation of the conductance of the Ca2+-sensitive K+ channel. The results show that the formation and removal of various firing activities in this model are due to two subcritical Hopf bifurcations of equilibrium based on theoretical computation. Furthermore, the computational simulations are shown to support the validity of the conceptual approach. Consequently, the conclusion could be helpful to improve and deepen our understanding of the contribution of the Ca2+-sensitive K+ channel.
Citation: Danqi Feng, Yu Chen, Quanbao Ji. Contribution of a Ca2+-activated K+ channel to neuronal bursting activities in the Chay model[J]. Electronic Research Archive, 2023, 31(12): 7544-7555. doi: 10.3934/era.2023380
[1] | Hui Zhou, Bo Lu, Huaguang Gu, Xianjun Wang, Yifan Liu . Complex nonlinear dynamics of bursting of thalamic neurons related to Parkinson's disease. Electronic Research Archive, 2024, 32(1): 109-133. doi: 10.3934/era.2024006 |
[2] | Yapeng Zhang, Yu Chen, Quanbao Ji . Dynamical analysis of spontaneous Ca2+ oscillations in astrocytes. Electronic Research Archive, 2024, 32(1): 405-417. doi: 10.3934/era.2024020 |
[3] | Xiaofang Jiang, Hui Zhou, Feifei Wang, Bingxin Zheng, Bo Lu . Bifurcation analysis on the reduced dopamine neuronal model. Electronic Research Archive, 2024, 32(7): 4237-4254. doi: 10.3934/era.2024191 |
[4] | Bo Lu, Xiaofang Jiang . Reduced and bifurcation analysis of intrinsically bursting neuron model. Electronic Research Archive, 2023, 31(10): 5928-5945. doi: 10.3934/era.2023301 |
[5] | Qixiang Wen, Shenquan Liu, Bo Lu . Firing patterns and bifurcation analysis of neurons under electromagnetic induction. Electronic Research Archive, 2021, 29(5): 3205-3226. doi: 10.3934/era.2021034 |
[6] | A. M. Elaiw, E. A. Almohaimeed, A. D. Hobiny . Modeling the co-infection of HTLV-2 and HIV-1 in vivo. Electronic Research Archive, 2024, 32(11): 6032-6071. doi: 10.3934/era.2024280 |
[7] | Fanqi Zeng . Some almost-Schur type inequalities and applications on sub-static manifolds. Electronic Research Archive, 2022, 30(8): 2860-2870. doi: 10.3934/era.2022145 |
[8] | Ke Yin, Kewei Zhang . Some computable quasiconvex multiwell models in linear subspaces without rank-one matrices. Electronic Research Archive, 2022, 30(5): 1632-1652. doi: 10.3934/era.2022082 |
[9] | Li Li, Zhiguo Zhao . Inhibitory autapse with time delay induces mixed-mode oscillations related to unstable dynamical behaviors near subcritical Hopf bifurcation. Electronic Research Archive, 2022, 30(5): 1898-1917. doi: 10.3934/era.2022096 |
[10] | Chengchun Hao, Tao Luo . Some results on free boundary problems of incompressible ideal magnetohydrodynamics equations. Electronic Research Archive, 2022, 30(2): 404-424. doi: 10.3934/era.2022021 |
The central nervous system extensively expresses Ca2+-stimulated K+ channels, which serve to use Ca2+ to control their opening and closing. In this study, we explore the numerical computation of Hopf bifurcation in the Chay model based on the equilibrium point's stability and the center manifold theorem to illustrate the emergence of complicated neuronal bursting induced by variation of the conductance of the Ca2+-sensitive K+ channel. The results show that the formation and removal of various firing activities in this model are due to two subcritical Hopf bifurcations of equilibrium based on theoretical computation. Furthermore, the computational simulations are shown to support the validity of the conceptual approach. Consequently, the conclusion could be helpful to improve and deepen our understanding of the contribution of the Ca2+-sensitive K+ channel.
It is known that the Chay model, comprising the mixed Na+-Ca2+ channel current and K+ channel current, can be used to simulate and describe various neuronal firings of pancreatic β cells, sensory terminals and cold receptors [1,2,3]. Exploring the Chay model is prevalent to understand not only physical-mathematical associations but also physiological due to its simplicity and abundant dynamic behavior consisting of synchronization and oscillations set off by noise. Ca2+-sensitive K+ channels are significant to the initiation of action potential, contributing substantially to physiological processes and its dysfunction leads to abnormal action potential propagation [4,5,6]. Bursting is crucial in the exchange of information between neurons, which is characterized by alternations between resting and repetitive firing states. The most significant messenger in cells is Ca2+, which conveys vital information about nearly every action essential to the survival and proper operation of cells. The dynamic modulation of several elements is necessary for Ca2+ signaling, and the Na+/Ca2+ exchanger (NCX) plays a role in maintaining its homeostasis by extracting Ca2+ from cells [7]. In both healthy and ischemic brains, astrocyte NCX activation may perform various roles. Studies are amassing that demonstrate the importance of one of the major glial ion transporters, NCX, in the regulation of astrocytic, microglial and oligodendrocytic functions. We can conjecture that alterations in NCX activity in distinct brain regions or astrocytic places may be linked to learning, memory and information processing functions in the brain [8]. It has become clear that neuronal firing patterns are usually associated with abundant dynamical behaviors since it is affected by intrinsic and extrinsic mediators, for instance, variation of ion path permeability, time delay and noise perturbation, as well as depolarizing current and so on [9,10,11]. Although nervous systems are quite different, neurons share many common features, such as action potential as carriers of information, ion channels and rich nonlinear phenomena.
Experimental and theoretical investigations indicate that action potentials generated by Na+, Ca2+ and K+ currents are attributed to Ca2+-activated K+ channels [12,13,14]. Due to the importance of electrical activities associated with Ca2+-activated K+ channels, the dynamic mechanism underlying bursting in the Chay model should be investigated in detail. This model was studied extensively by Duan et al. [15,16,17], Xu et al. [18,19], Lu et al. [20,21] and Zhu et al. [22]. However, most studies are confined to numerical simulation of the Hopf bifurcation with variations in different bifurcation parameters [15,16,17,18]. Based on these previous works, the mechanisms and contributions involved in firing activities related to various ion channels are not well understood. Hence, we simulate the bursting dynamics associated with Ca2+-activated K+ ion channels in combination with the use of the Chay model.
We use the two-pool model by Chay [23] and develop upon it as a demonstration of stability and bifurcation analysis. This model is formed from three dynamic indexes: The membrane voltage (V), the concentration of Ca2+ within the cell (C) and the odds of triggering the voltage-sensitive K+ channel (n). The model comprises three equations:
{ dVdt=g∗L(VL−V)+g∗K, Vn4(VK−V)+g∗Im3∞h∞(VI−V)+g∗K, CC1+C(VK−V), dCdt=ρ[m3∞h∞(VC−V)−kCC], dndt=n∞−nτn, | (1) |
where
m∞=αmαm+βm, n∞=αnαn+βn, h∞=αhαh+βh,αm=0.1(25+V)1−e−0.1V−2.5, βm=4e−(V+50)18, αh=0.07e−0.05V−2.5,βh=11+(e−0.1V−2), αn=0.01(20+V)1−e−0.1V−2, βn=0.125e−(V+30)80,τn=1λn(αn+βn). |
The details of each parameter can be found in [23]. Here, gkc* is designated as the bifurcation parameter. It represents the highest conductance of Ca2+-sensitive K+ channel. Let x = V, y = C, z = n, r = gkc*. In order to simplify the calculation process, system (1) is transformed into the subsequent expression:
{˙x=126.0e−0.05x−2.5(x−100)(0.1x+2.5)3(e−0.1x−2.5−1)3(0.07e−0.05x−2.5+1e−0.1x−2+1)(4e−x18−259−0.1x+2.5e−0.1x−2.5−1)3−1700z4(x+75) −ry(x+75)y+1−7x−280,˙y=0.0189e−0.05x−2.5(x−100)(0.1x+2.5)3(e−0.1x−2.5−1)3(0.07e−0.05x−2.5+1e−0.1x−2+1)(4e−x18−259−0.1x+2.5e−0.1x−2.5−1)3−0.0495y,˙z=−(28.75e−x80−38−230(0.01x+0.2)e−0.1x−2−1)(z+0.01x+0.2(e−0.1x−2−1)(0.125e−x80−38−0.01x+0.2e−0.1x−2−1)). | (2) |
The existence of equilibrium points can be determined by analyzing the differential equations of model. Suppose system (2) has three roots x0, y0, z0. Let x1 = x – x0, y1 = y – y0, z1 = z – z0, we get the following representations:
{˙x1=126.0e−2.5−0.05(x0+x1)((x0+x1)−100)(2.5+0.1(x0+x1))3(0.07e−0.05(x1+x0)−2.5+1e−0.1(x1+x0)−2+1)(e−0.1(x1+x0)−2.5−1)3(4e−x1+x018−259−0.1(x1+x0)+2.5e−0.1(x1+x0)−2.5−1)3 −1700(z1+z0)4((x1+x0)+75)−(y0+y1)r((x0+x1)+75)y1+y0+1−7(x0+x1)−280,˙y1=0.0189e−0.05(x1+x0)−2.5((x1+x0)−100)(0.1(x1+x0)+2.5)3(0.07e−0.05(x1+x0)−2.5+1e−0.1(x1+x0)−2+1)(e−0.1(x1+x0)−2.5−1)3(4e−x1+x018−259−0.1(x1+x0)+2.5e−0.1(x1+x0)−2.5−1)3 −0.0495(y1+y0),˙z1=−(28.75e−x1+x080−38−230(0.01(x1+x0)+0.2)e−(x0+x1)0.1−2−1) (z1+z0+0.2+(x0+x1)0.01(e−0.1(x1+x0)−2−1)(0.125e−x1+x080−38−0.01(x1+x0)+0.2e−0.1(x1+x0)−2−1)). | (3) |
Equilibrium is (0, 0, 0) and system (3) has the same properties in system (2). It is clear that the Jacobian matrix (aij)3×3 of system (3) and the characteristic equations satisfy: λ3 + Q1λ2 + Q2 λ + Q1 = 0, where Q3 = a31a13a22 + a12a21a33 + a32a23a11 – a11a22a33 – a12a23a31 – a13a21a32, Q2 = a11a22 + a11a33 + a22a33 – a13a31 – a12a21 – a32a23, Q1 = – (a11 + a22 + a33). Examine the Hurwitz matrix in the context of the coefficients Qi (i = 1, 2, 3) of the characteristic polynomial:
H1=[Q1],H2=(Q11Q3Q2),H3=[Q110Q3Q2100Q3]. |
The eigenvalues are negative when the determinant values of Hurwitz matrix are bigger than zero:
det(Hi)>0,i=1,2,3. |
The robustness of system (3) is taken into account as varying the values of gkc* using the criteria of the Routh's array:
Q1>0,Q3>0,Q1Q2>Q3. |
It is easy to see that:
1) r < -41.647, system (3) contains a stable node;
2) r = -41.647, the system possesses a non-hyperbolic stationary state, which is O1 = (-17.5904, 4.7463, 0.4334);
3) -41.647 < r < 27.25, the system includes a saddle point;
4) r = 27.25, the system contains a non-hyperbolic stationary state, which is O2 = (24.7680, -0.8969, 0.6095);
5) r > 27.25, the system is stable.
The system (3) reaches equilibrium at (x0, y0, z0) as r = r0, x1 = x - x0, y1 = y - y0, z1 = z - z0, r1 = r - r0. Next, we introduce a new variable denoted as r1 in the application of center manifold theorem with respect to the parameter gkc*. Let dr1/dt = 0, we obtain:
{˙x1=126.0e−2.5−0.05(x0+x1)((x0+x1)−100)(2.5+0.1(x0+x1))3(0.07e−0.05(x1+x0)−2.5+1e−0.1(x1+x0)−2+1)(e−0.1(x1+x0)−2.5−1)3(4e−x1+x018−259−0.1(x1+x0)+2.5e−0.1(x1+x0)−2.5−1)3 −1700(z0+z1)4((x0+x1)+75)−(y0+y1)((x1+x0)+75)(r0+r1)y1+y0+1−7(x1+x0)−280,˙y1=0.0189e−2.5−0.05(x0+x1)((x0+x1)−100)(2.5+0.1(x0+x1))3(0.07e−0.05(x1+x0)−2.5+1e−0.1(x1+x0)−2+1)(e−0.1(x1+x0)−2.5−1)3(4e−x1+x018−259−0.1(x1+x0)+2.5e−0.1(x1+x0)−2.5−1)3 −0.0495(y1+y0),˙z1=−(28.75e−x1+x080−38−230(0.01(x1+x0)+0.2)e−2−0.1(x0+x1)−1) (z0+z1+0.01(x0+x1)+0.2(e−0.1(x1+x0)−2−1)(0.125e−x1+x080−38−0.01(x1+x0)+0.2e−0.1(x1+x0)−2−1)),˙r1=0. | (4) |
As r1 = 0, system (4) achieves equilibrium O (x1, y1, z1, r1) = (0, 0, 0, 0), which has the identical property in system (2). When r0 = -41.647, we consider the characteristic values of balanced state O1 = (0, 0, 0, 0) in system (4): ξ1 = -0.0005, ξ2 = 1.6305i, ξ3 = -1.6305i, ξ4 = 0.
Suppose
(x1y1z1r1)=U(uvws),whereU=(0.12631.00000−0.08290.9920000.00220.00210.00010.0051−0.00140000.9966), |
system (4) has the following form
(˙u˙v˙w˙s)=(−0.000500000−1.6305001.6305000000)(uvws)+(g1g2g3g4), | (5) |
where
g1=1.008064516f2−0.002225307983f4+0.0005u,g2=f1−0.1273185484f2+0.08346387799f4+1.6305w,g3=−0.01960784314f1−0.4125889469f2+196.0784314f3+0.2747260781f4−1.6305v,g4=0,f1=126.0e−0.05g11−2.5(g11−100)(0.1g11+2.5)3(0.07e−0.05g11−2.5+1e−0.1g11−2+1)(e−0.1g11−2.5−1)3(4e−g1118−259−0.1g11+2.5e−0.1g11−2.5−1)3 −1700g134(g11+75)−g14g12(g11+75)g12+1−7g11−280,f2=0.0189e−0.05g11−2.5(g11−100)(0.1g11+2.5)3(0.07e−0.05g11−2.5+1e−0.1g11−2+1)(e−0.1g11−2.5−1)3(4e−g1118−259−0.1g11+2.5e−0.1g11−2.5−1)3 −0.0495g12,f3=−(28.75e−g1180−38−230(0.01g11+0.2)e−0.1g11−2−1) (g13+0.01g11+0.2(e−0.1g11−2−1)(0.125e−g1180−38−0.01g11+0.2e−0.1g11−2−1)),f4=0,g11=x1+x0=0.1263u−0.0829s+1.0v−17.5904,g12=y1+y0=0.0022s+0.992u+4.7463,g13=z1+z0=0.0021u−0.0014s+0.0001v+0.0051w+0.4334,g14=0.9966s−41.647. |
Based on center manifold theorem, a center manifold exists. The specific form is as follows:
Wcloc(O1)={(u,v,w,s)∈R4|u=h∗(v,w,s),h∗(0,0,0)=0,Dh∗(0,0,0)=0}, |
Assume h (v, w, s) = av2 + bw2 + cs2 + dvw + evs + fws +…, the center manifold is
N(h)=Dh⋅[˙v˙w˙s]+0.0005h−g1≡0. | (6) |
Here a = -0.0000036406, b = -0.000003093017389, c = 0.00008567728077, d = 0.00000180521, e = 0.00000016, f = -0.0000005491. If the system is limited by center manifold, the following conditions are satisfied:
(˙v˙w)=(0−1.63051.63050)(vw)+(f1(v,w)f2(v,w)), | (7) |
where
f1(v,w)=0.580313865s−7.0v+1.6305w−0.0000001404922113sv+0.0000004820211492sw −0.000001584700346vw+⋯,f2(v,w)=−0.01133350044s−1.493245098v+0.000000006016773356sv−0.00000002064322272sw +0.00000006786698519vw+⋯. |
Then, we have
a=116[f1vvv+f1vww+f2vvw+f2www]|(0,0)+116×0.0204[f1vw(f1vv+f1ww)−f2vw(f2vv+f2ww)−f1vvf2vv+f1wwf2ww)]|(v=0,w=0,s=0)=3.01538319 > 0,d=d(Re(ξ(s))ds|(v=0,w=0,s=0)=−0.00001623167121<0. | (8) |
According to previous numerical computations, we have:
Conclusion 1: At r0 = -41.647, a supercritical Hopf bifurcation is obtained. When the value of r is less than r0, the equilibrium O1 turns to be stable. The equilibrium state loses stability as r > r0 and a stable periodic solution appears, which causes the oscillation of system (2). As r0 = 27.25, the characteristic values are ξ1 = 0.0036, ξ2 = 3.8251i, ξ3 = -3.8251i and ξ4 = 0, respectively.
The simplified form based on the center manifold system (4) is depicted as
(˙v˙w)=(0−3.82513.82510)(vw)+(f1(v,w)f2(v,w)), | (9) |
where
f1(v,w)=0.62108s−7.0v+3.8251w−0.000007826615347sv+0.00001148249661sw −0.0001076807683vw+⋯,f2(v,w)=−0.1979232s−2.9851v+0.000001122019432sv−0.000001646124635sw +0.00001543705793vw+⋯. |
The next conclusion can be obtained as a = 0.32334838 > 0 and d = 0.01462214783 > 0.
Conclusion 2: At r0 = 27.25, the system exhibits a subcritical Hopf bifurcation transition. As r is less than r0, the equilibrium O2 is in an unstable state. As r > r0, the equilibrium O2 returns to be stable.
The generation processes of the parameter gkc* are presented so as to study the bifurcation phenomenon underlying various firing activities. Figures 1(a), (b) show the bifurcations that illustrate the stationary states of system (2) in the (V, gkc*) and (C, gkc*) planes, respectively. Each point of the solid line in the curve denotes equilibrium stability, and the dotted curve is an unsteady stationary state. Additionally, gkc* passes through two Hopf bifurcations labeled with HB1 and HB2, where gkc* 1 = -41.647 μM/s and gkc* 2 = 27.25 μM/s. As gkc* is increased, the balanced stationary initially loses its steady state at HB1, only to regain the balanced state at HB2.
Because of the fluctuation in the parameter gkc*, the two Hopf bifurcation points are plainly visible. It can be seen that gkc* has a stable equilibrium between gkc* = -80 pS and -41.647 pS, as well as between gkc* = 27.25 pS and 100 pS. There are two unstable equilibria in the range of gkc* = -41.647 pS to 27.25 pS. Firing activities can be obtained from the ISI bifurcation with variation of the parameter gkc*. As gkc* is increased approximately to 30 pS, simple firing activities occur (Figure 1(c)). Then, in Figure 1(d), a partially enlarged picture of the ISI bifurcation in the (gkc*, C) plane is shown. As gkc* is increased to 31 pS, bursts are observed.
The corresponding time series of system (2) are presented in Figure 2. Figure 2(a1), (b1), (c1), (d1) shows the time evolution of parameter gkc* with different values. Figure 2(a2), (b2), (c2), (d2) represents distinct state trajectories in the three-dimensional phase space under different gkc* values.
Figure 2 depicts the time-varying membrane voltage for various values of the parameter gkc*. On the left, the time evolution of V is compared with gkc*. The state trajectories in the 3D phase portrait V-C-n space are shown in the middle panels, and the right panels are the time-frequency of parameter gkc*. For instance, there is a single peak in the oscillation when gkc* = 21 pS in Figure 2(a1). The one-to-one correspondence of the three-dimensional phase space is visualized in Figure 2(a2). Moreover, Figure 2(a3) shows a temporal-spectral pattern with the parameter gkc* = 21 pS. When the value of the bifurcation parameter gkc* drops, the total number of peak counts and magnitudes begins to increase. Similarly, six peaks were produced when gkc* = 11 pS, as shown in Figure 2(b1). Then, in Figure 2(c2), chaos appears. Furthermore, the number of peak counts in Figure 2(d1) tends to oscillate in a periodic fashion. Figure 2(a3), (b3), (c3), (d3) illustrates the frequency spectrogram of the temporal evolution pattern, making the change observation more apparent.
The membrane voltage of neurons exhibits spontaneous oscillations in Figure 2(a1). The temporal evolution Figure 2(a1), which corresponds to the 3D phase Figure 2(a2), appears as an inflection point, denoted by a red hollow circle. Figure 2(b1) displays a multi-peak oscillation phenomenon as the parameter gkc* lowers continually, in contrast to the single-peak oscillation phenomenon in Figure 2(a1). Among these, Figure 2(a1), (b1), (d1) represents a regular burst of membrane potential (V). Figure 2(c1) depicts chaos. In the third experimental situation, the evolution diagram of time shows significant irregularity, that is, the irregular spike sequence of spontaneous oscillations of membrane voltage (V). These phenomena are plentiful and warrant further investigation.
Mathematical modeling and numerical simulation are two effective methods to help us understand the internal workings of the neuronal system. We investigated the properties of primitive hippocampal neurons in the Chay model using the bifurcation parameter gkc*. We analyzed the theoretical stability of equilibrium and bifurcation and explored the variations in the conductance of the Ca2+-sensitive K+ channel. As the parameter gkc* varies, two supercritical Hopf critical nodes were found.
The Chay model exhibited a bi-stability phenomenon, namely the coexistence of chaotic attractors and stationary point attractors. This phenomenon was numerically revealed through time evolution, local bifurcation, phase planes and spectrum diagrams. Numerical calculations of Hopf bifurcations confirmed the theoretical analysis. It is concluded that the conductance of Ca2+-sensitive K+ channels leads to the emergence of complex neuronal bursting. Thus, some dynamic behaviors of system (1) are schematically presented. Measurements corroborated the numerical results, displaying dynamic behaviors. Validation of theoretical results was achieved through the use of numerical methods, which were employed after conducting a thorough theoretical analysis. Other complex dynamical behaviors of the presented Chay model should be further studied. We aim to conduct more comprehensive research on the relationship between the synchronization of oscillatory patterns and bifurcation in our upcoming investigations.
The authors declare they have not used Artificial Intelligence (AI) tools in the creation of this article.
This work was supported by the National Natural Science Foundation of China (No. 12062004), the Natural Science Foundation of Guangxi Minzu University (No. 2022KJQD01), the Guangxi Natural Science Foundation (2020GXNSFAA297240), Guangxi Science and Technology Program (Grant No. AD23023001) and Xiangsi Lake Young Scholars Innovation Team of Guangxi Minzu University (No. 2021RSCXSHQN05).
The authors declare there is no conflict of interest.
[1] |
K. Koyama, H. Ando, K. Fujiwara, Functional improvement in β cell models of type 2 diabetes using on-demand feedback control, AIP Adv., 13 (2023), 045317. https://doi.org/10.1063/5.0124625 doi: 10.1063/5.0124625
![]() |
[2] |
Z. Yang, Q. Lu, L. Li, The genesis of period-adding bursting without bursting-chaos in the Chay model, Chaos, Solitons Fractals, 27 (2006), 689–697. https://doi.org/10.1016/j.chaos.2005.04.038 doi: 10.1016/j.chaos.2005.04.038
![]() |
[3] |
Z. Yang, Q. Lu, H. Gu, W. Ren, Integer multiple spiking in the stochastic Chay model and its dynamical generation mechanism, Phys. Lett. A, 299 (2002), 499–506. https://doi.org/10.1016/S0375-9601(02)00746-6 doi: 10.1016/S0375-9601(02)00746-6
![]() |
[4] |
M. Gu, Y. Zhu, X. Yin, D. M. Zhang, Small-conductance Ca2+-activated K+ channels: insights into their roles in cardiovascular disease, Exp. Mol. Med., 50 (2018), 1–7. https://doi.org/10.1038/s12276-018-0043-z doi: 10.1038/s12276-018-0043-z
![]() |
[5] |
X. Chen, Y. Feng, R. J. Quinn, D. L. Pountney, D. R. Richardson, G. D. Mellick, et al., Potassium channels in parkinson's disease: potential roles in its pathogenesis and innovative molecular targets for treatment, Pharm. Rev., 75 (2023), 758–788. https://doi.org/10.1124/pharmrev.122.000743 doi: 10.1124/pharmrev.122.000743
![]() |
[6] |
J. P. Adelman, J. Maylie, P. Sah, Small-conductance Ca2+-activated K+ channels: form and function, Annu. Rev. Physiol., 74 (2012), 245–269. https://doi.org/10.1146/annurev-physiol-020911-153336 doi: 10.1146/annurev-physiol-020911-153336
![]() |
[7] |
M. Al-Khannaq, J. Lytton, Regulation of K+-Dependent Na+/Ca2+-Exchangers (NCKX), Int. J. Mol. Sci., 24 (2023), 598. https://doi.org/10.3390/ijms24010598 doi: 10.3390/ijms24010598
![]() |
[8] |
S. Song, L. Luo, B. Sun, D. Sun, Roles of glial ion transporters in brain diseases, Glia, 68 (2020), 472–494. https://doi.org/10.1002/glia.23699 doi: 10.1002/glia.23699
![]() |
[9] |
Y. Li, R. Wang, T. Zhang, Nonlinear computational models of dynamical coding patterns in depression and normal rats: from electrophysiology to energy consumption, Nonlinear Dyn., 107 (2022), 3847–3862. https://doi.org/10.1007/s11071-021-07079-7 doi: 10.1007/s11071-021-07079-7
![]() |
[10] |
L. Li, Z. Zhao, White-noise-induced double coherence resonances in reduced Hodgkin-Huxley neuron model near subcritical Hopf bifurcation, Phys. Rev. E: Stat. Nonlinear Soft Matter Phys., 105 (2022), 034408. https://doi.org/10.1103/PhysRevE.105.034408 doi: 10.1103/PhysRevE.105.034408
![]() |
[11] |
P. Crotty, K. Segall, D. Schult, Biologically realistic behaviors from a superconducting neuron model, IEEE Trans. Appl. Supercond., 33 (2023), 1–6. https://doi.org/10.1109/TASC.2023.3242901 doi: 10.1109/TASC.2023.3242901
![]() |
[12] | L. Blomer, On the voltage gated ion channels involved in action potential generation and back propagation in layer 5 pyramidal neurons, 2022. Available from: https://theses.hal.science/tel-04077615/file/BLOMER_2022_archivage.pdf. |
[13] |
M. V. Roshchin, V. N. Ierusalimsky, P. M. Balaban, E. S. Nikitin, Ca2+-activated KCa3.1 potassium channels contribute to the slow afterhyperpolarization in L5 neocortical pyramidal neurons, Sci. Rep., 10 (2020), 14484. https://doi.org/10.1038/s41598-020-71415-x doi: 10.1038/s41598-020-71415-x
![]() |
[14] |
R. Orfali, N. Albanyan, Ca2+-Sensitive potassium channels, Molecules, 28 (2023), 885. https://doi.org/10.3390/molecules28020885 doi: 10.3390/molecules28020885
![]() |
[15] |
L. Duan, Q. Lu, Q. Wang, Two-parameter bifurcation analysis of firing activities in the Chay neuronal model, Neurocomputing, 72 (2008), 341–351. https://doi.org/10.1016/j.neucom.2008.01.019 doi: 10.1016/j.neucom.2008.01.019
![]() |
[16] |
L. Duan, Q. Lu, Codimension-two bifurcation analysis on firing activities in Chay neuron model, Chaos, Solitons Fractals, 30 (2006), 1172–1179. https://doi.org/10.1016/j.chaos.2005.08.179 doi: 10.1016/j.chaos.2005.08.179
![]() |
[17] |
L. Duan, Q. Lu, Bursting oscillations near codimension-two bifurcations in the Chay Neuron model, Int. J. Nonlinear Sci. Numer. Simul., 7 (2006), 59–64. https://doi.org/10.1515/IJNSNS.2006.7.1.59 doi: 10.1515/IJNSNS.2006.7.1.59
![]() |
[18] |
Q. Xu, X. Tan, D. Zhu, H. Bao, Y. Hu, B. Bao, Bifurcations to bursting and spiking in the Chay neuron and their validation in a digital circuit, Chaos, Solitons Fractals, 141 (2020), 110353. https://doi.org/10.1016/j.chaos.2020.110353 doi: 10.1016/j.chaos.2020.110353
![]() |
[19] |
Q. Xu, X. Tan, D. Zhu, M. Chen, J. Zhou, H. Wu, Synchronous behavior for memristive synapse-connected Chay twin-neuron network and hardware implementation, Math. Probl. Eng., 2020 (2020), 8218740. https://doi.org/10.1155/2020/8218740 doi: 10.1155/2020/8218740
![]() |
[20] |
L. Lu, M. Yi, Z. Gao, Y. Wu, X. Zhao, Critical state of energy-efficient firing patterns with different bursting kinetics in temperature-sensitive Chay neuron, Nonlinear Dyn., 111 (2023), 16557–16567. https://doi.org/10.1007/s11071-023-08700-7 doi: 10.1007/s11071-023-08700-7
![]() |
[21] |
L. L. Lu, M. Yi, X. Q. Liu, Energy-efficient firing modes of chay neuron model in different bursting kinetics, Sci. China Technol. Sci., 65 (2022), 1661–1674. https://doi.org/10.1007/s11431-021-2066-7 doi: 10.1007/s11431-021-2066-7
![]() |
[22] |
F. Zhu, R. Wang, K. Aihara, X. Pan, Energy-efficient firing patterns with sparse bursts in the Chay neuron model, Nonlinear Dyn., 100 (2020), 2657–2672. https://doi.org/10.1007/s11071-020-05593-8 doi: 10.1007/s11071-020-05593-8
![]() |
[23] |
T. R. Chay, Chaos in a three-variable model of an excitable cell, Physica D, 16 (1985), 233–242. https://doi.org/10.1016/0167-2789(85)90060-0 doi: 10.1016/0167-2789(85)90060-0
![]() |