Network mechanism for burst generation

We report on the mechanism of burst generation by populations of intrinsically spiking neurons, when a certain threshold in coupling strength is exceeded. These ensembles synchronize at relatively low coupling strength and lose synchronization at stronger coupling via spatiotemporal intermittency. The latter transition triggers fast repetitive spiking, which results in synchronized bursting. We present evidence that this mechanism is generic for various network topologies from regular to small-world and scale-free ones, different types of coupling and neuronal model.     

Pre-B?tzinger复合体的从簇到峰放电的同步转迁及分岔机制

Pre-Bötzinger复合体是兴奋性耦合的神经元网络,通过产生复杂的放电节律和节律模式的同步转迁参与调控呼吸节律.本文选用复杂簇和峰放电节律的单神经元数学模型构建复合体模型,仿真了与生物学实验相关的多类同步节律模式及其复杂转迁历程,并利用快慢变量分离揭示了相应的分岔机制.当初值相同时,随着兴奋性耦合强度的增加,复合体模型依次表现出完全同步的“fold/homoclinic”,“subHopf/subHopf”簇放电和周期1峰放电.当初值不同时,随耦合强度增加,表现为由“fold/homoclinic”,到“fold/fold limit cycle”、到“subHopf/subHopf”与“fold/fold limit cycle”的混合簇放电、再到“subHopf/subHopf”簇放电的相位同步转迁,最后到反相同步周期1峰放电.完全(同相)同步和反相同步的周期1节律表现出了不同分岔机制.反相峰同步行为给出了与强兴奋性耦合容易诱发同相同步这一传统观念不同的新示例.研究结果给出了preBötzinger复合体的从簇到峰放电节律的同步转迁规律及复杂分岔机制,反常同步行为丰富了非线性动力学的内涵.     

Transition to bursting via deterministic chaos

We study statistical properties of the irregular bursting arising in a class of neuronal models close to the transition from spiking to bursting. Prior to the transition to bursting, the systems in this class develop chaotic attractors, which generate irregular spiking. The chaotic spiking gives rise to irregular bursting. The duration of bursts near the transition can be very long. We describe the statistics of the number of spikes and the interspike interval distributions within one burst as functions of the distance from criticality.     

Intermittent synchronization in a network of bursting neurons

Synchronized oscillations in networks of inhibitory and excitatory coupled bursting neurons are common in a variety of neural systems from central pattern generators to human brain circuits. One example of the latter is the subcortical network of the basal ganglia, formed by excitatory and inhibitory bursters of the subthalamic nucleus and globus pallidus, involved in motor control and affected in Parkinson's disease. Recent experiments have demonstrated the intermittent nature of the phase-locking of neural activity in this network. Here, we explore one potential mechanism to explain the intermittent phase-locking in a network. We simplify the network to obtain a model of two inhibitory coupled elements and explore its dynamics. We used geometric analysis and singular perturbation methods for dynamical systems to reduce the full model to a simpler set of equations. Mathematical analysis was completed using three slow variables with two different time scales. Intermittently, synchronous oscillations are generated by overlapped spiking which crucially depends on the geometry of the slow phase plane and the interplay between slow variables as well as the strength of synapses. Two slow variables are responsible for the generation of activity patterns with overlapped spiking, and the other slower variable enhances the robustness of an irregular and intermittent activity pattern. While the analyzed network and the explored mechanism of intermittent synchrony appear to be quite generic, the results of this analysis can be used to trace particular values of biophysical parameters (synaptic strength and parameters of calcium dynamics), which are known to be impacted in Parkinson's disease.     

On the role of subthreshold currents in the Huber-Braun cold receptor model

We study the role of the strength of subthreshold currents in a four-dimensional Hodgkin-Huxley-type model of mammalian cold receptors. Since a total diminution of subthreshold activity corresponds to a decomposition of the model into a slow, subthreshold, and a fast, spiking subsystem, we first elucidate their respective dynamics separately and draw conclusions about their role for the generation of different spiking patterns. These results motivate a numerical bifurcation analysis of the effect of varying the strength of subthreshold currents, which is done by varying a suitable control parameter. We work out the key mechanisms which can be attributed to subthreshold activity and furthermore elucidate the dynamical backbone of different activity patterns generated by this model.     

Phase synchronization in ensembles of bursting oscillators

We study the effects of mutual and external chaotic phase synchronization in ensembles of bursting oscillators. These oscillators (used for modeling neuronal dynamics) are essentially multiple time scale systems. We show that a transition to mutual phase synchronization takes place on the bursting time scale of globally coupled oscillators, while on the spiking time scale, they behave asynchronously. We also demonstrate the effect of the onset of external chaotic phase synchronization of the bursting behavior in the studied ensemble by a periodic driving applied to one arbitrarily taken neuron. We also propose an explanation of the mechanism behind this effect. We infer that the demonstrated phenomenon can be used efficiently for controlling bursting activity in neural ensembles.     

New dynamics in cerebellar Purkinje cells: torus canards

We describe a transition from bursting to rapid spiking in a reduced mathematical model of a cerebellar Purkinje cell. We perform a slow-fast analysis of the system and find that-after a saddle node bifurcation of limit cycles-the full model dynamics temporarily follow a repelling branch of limit cycles. We propose that the system exhibits a dynamical phenomenon new to realistic, biophysical applications: torus canards.     

兴奋性作用诱发神经簇放电个数不增反降的分岔机制

非线性动力学在识别神经放电的复杂现象、机制和功能方面发挥了重要作用.不同于传统观念,本文提出了兴奋性作用可以降低而不是增加簇内放电个数的新观点.在簇放电模式休止期的适合相位施加强度合适的脉冲或自突触电流,能诱发簇内放电个数降低;电流的施加相位越早,所需的强度阈值越大,簇内放电个数越少.进一步,利用快慢变量分离获得的簇放电的动力学性质进行了理论解释.簇放电模式表现出低电位的休止期和高电位的放电的交替,存在于快子系统的鞍结分岔点和同宿轨分岔点之间;放电起始于鞍结分岔、结束于同宿轨分岔;越靠近同宿轨分岔从休止期跨越到放电所需的电流强度越大.因此,电流在休止期上的作用相位越早,就越靠近同宿轨分岔,因而从休止期跨越到放电需要的电流强度阈值越大,放电起始相位到同宿轨分岔之间的区间变小导致放电个数变少.研究结果丰富了非线性现象及机制,对兴奋性作用提出了新看法,给出了调控簇放电模式的新途径.     

Parameter-dependent synchronization transition of coupled neurons with co-existing spiking and bursting

A firing pattern transition is simulated in the Leech neuron model, firstly from bursting to co-existence of spiking and bursting and then to spiking. The attraction domain of spiking and bursting for three different parameter values are calculated. Synchronization transition processes of two coupled Leech neurons, one is bursting and the other the co-existing spiking, are simulated for the three parameters. The three synchronization processes appear similar as the coupling strength increases, beginning from non-synchronization to complete synchronization through a complex dynamical procedure, but their detailed processes are different depending on the parameter values. The transition procedure is complex and the complete synchronization is in bursting for larger parameter values, while the process is simple with complete synchronization of spiking for smaller values. The potential relationship between complete synchronization and the attraction domain is also discussed. The results are instructive to understanding the synchronization behaviors of the coupled neuronal system with co-existing attractors.     

Phase synchronization of bursting neural networks with electrical and delayed dynamic chemical couplings

Diffusive electrical connections in neuronal networks are instantaneous, while excitatoryor inhibitory couplings through chemical synapses contain a transmission time-delay.Moreover, chemical synapses are nonlinear dynamical systems whose behavior can bedescribed by nonlinear differential equations. In this work, neuronal networks withdiffusive electrical couplings and time-delayed dynamic chemical couplings are considered.We investigate the effects of distributed time delays on phase synchronization of burstingneurons. We observe that in both excitatory and Inhibitory chemical connections, the phasesynchronization might be enhanced when time-delay is taken into account. This distributedtime delay can induce a variety of phase-coherent dynamical behaviors. We also study thecollective dynamics of network of bursting neurons. The network model presents theso-called Small-World property, encompassing neurons whose dynamics have two time scales(fast and slow time scales). The neuron parameters in such Small-World network, aresupposed to be slightly different such that, there may be synchronization of the bursting(slow) activity if the coupling strengths are large enough. Bounds for the criticalcoupling strengths to obtain burst synchronization in terms of the network structure aregiven. Our studies show that the network synchronizability is improved, as itsheterogeneity is reduced. The roles of synaptic parameters, more precisely those of thecoupling strengths and the network size are also investigated.     

Dynamical phases of the Hindmarsh-Rose neuronal model: studies of the transition from bursting to spiking chaos

The dynamical phases of the Hindmarsh-Rose neuronal model are analyzed in detail by varying the external current I. For increasing current values, the model exhibits a peculiar cascade of nonchaotic and chaotic period-adding bifurcations leading the system from the silent regime to a chaotic state dominated by bursting events. At higher I-values, this phase is substituted by a regime of continuous chaotic spiking and finally via an inverse period doubling cascade the system returns to silence. The analysis is focused on the transition between the two chaotic phases displayed by the model: one dominated by spiking dynamics and the other by bursts. At the transition an abrupt shrinking of the attractor size associated with a sharp peak in the maximal Lyapunov exponent is observable. However, the transition appears to be continuous and smoothed out over a finite current interval, where bursts and spikes coexist. The beginning of the transition (from the bursting side) is signaled from a structural modification in the interspike interval return map. This change in the map shape is associated with the disappearance of the family of solutions responsible for the onset of the bursting chaos. The successive passage from bursting to spiking chaos is associated with a progressive pruning of unstable long-lasting bursts.     

Breakdown of order preservation in symmetric oscillator networks with pulse-coupling

Symmetric networks of coupled dynamical units exhibit invariant subspaces with two or more units synchronized. In time-continuously coupled systems, these invariant sets constitute barriers for the dynamics. For networks of units with local dynamics defined on the real line, this implies that the units' ordering is preserved and that their winding number is identical. Here, we show that in permutation-symmetric networks with pulse-coupling, the order is often no longer preserved. We analytically study a class of pulse-coupled oscillators (characterizing for instance the dynamics of spiking neural networks) and derive quantitative conditions for the breakdown of order preservation. We find that in general pulse-coupling yields additional dimensions to the state space such that units may change their order by avoiding the invariant sets. We identify a system of two symmetrically pulse-coupled identical oscillators where, contrary to intuition, the oscillators' average frequencies and thus their winding numbers are different.     

Routes to bursting in a periodically driven oscillator

The dynamical behaviors of a periodic excited oscillator with multiple time scales in the form that order gap exists between the frequency of the excitation and the natural frequency, are investigated in this Letter. By regarding the whole excitation term as a parameter, bifurcation sets are derived, which divide the generalized parameter space into several regions corresponding to different kinds of dynamics. Different types of bursting phenomena, such as fold/Hopf bursting, fold/Hopf/homoclinic bursting and Hopf/homoclinic bursting, are presented, the mechanism of which is obtained based on the bifurcations of the generalized autonomous system as well as the introduction of the so-called transformed phase portraits. Furthermore, the evolution of the bursting is discussed in details, in which one may find that when the two limit cycles caused by the Hopf bifurcations of the two related equilibrium points interact with each other, homoclinic bifurcation may occur, leading to the merge of the two cycles to form a large amplitude cycle. The homoclinic bifurcation may cause the two asymmetric bursters to merge into a symmetric enlarged burster, in which the large amplitude of the spiking state agrees well with the amplitude of the cycle caused by the homoclinic bifurcation.     

计算相位响应曲线的方波扰动直接算法

通过相位响应曲线可对具有极限环周期运动的动力系统的性质有更为深入的理解.神经元是一个典型的动力系统,因此相位响应曲线提供了一种研究神经元重复周期放电行为的新思路.本文提出一种求解相位响应曲线的方法,即方波扰动的直接算法,通过Hodgkin-Huxley,Fitz Hugh-Nagumo,Morris-Lecar和Hindmarsh-Rose神经元模型验证该算法可计算周期峰放电、周期簇放电的相位响应曲线.该算法克服了其他算法在运用过程中的局限性.利用该算法计算结果表明:周期峰放电的相位响应曲线类型是由其分岔类型所决定;在Morris-Lecar模型中发现一种开始于Hopf分岔终止于鞍点同宿轨道分岔的阈上周期振荡,其相位响应曲线属于第二类型.通过大量的相位响应曲线的计算发现相位响应的相对大小及正负性仅取决于扰动所施加的时间,而且周期簇放电的相位响应曲线比周期峰放电的相位响应曲线更为复杂.     

Bursting phenomena as well as the bifurcation mechanism in a coupled BVP oscillator with periodic excitation

We explore the complicated bursting oscillations as well as the mechanism in a high-dimensional dynamical system.By introducing a periodically changed electrical power source in a coupled BVP oscillator, a fifth-order vector field with two scales in frequency domain is established when an order gap exists between the natural frequency and the exciting frequency.Upon the analysis of the generalized autonomous system, bifurcation sets are derived, which divide the parameter space into several regions associated with different types of dynamical behaviors. Two typical cases are focused on as examples,in which different types of bursting oscillations such as sub Hopf/sub Hopf burster, sub Hopf/fold-cycle burster, and doublefold/fold burster can be observed. By employing the transformed phase portraits, the bifurcation mechanism of the bursting oscillations is presented, which reveals that different bifurcations occurring at the transition between the quiescent states(QSs) and the repetitive spiking states(SPs) may result in different forms of bursting oscillations. Furthermore, because of the inertia of the movement, delay may exist between the locations of the bifurcation points on the trajectory and the bifurcation points obtained theoretically.     

Homoclinic bifurcations leading to the emergence of bursting oscillations in cell models

We present a qualitative analysis of a generic model structure that can simulate the bursting and spiking dynamics of many biological cells. Four different scenarios for the emergence of bursting are described. In this connection a number of theorems are stated concerning the relation between the phase portraits of the fast subsystem and the global behavior of the full model. It is emphasized that the onset of bursting involves the formation of a homoclinic orbit that travels along the route of the bursting oscillations and, hence, cannot be explained in terms of bifurcations in the fast subsystem. In one of the scenarios, the bursting oscillations arise in a homoclinic bifurcation in which the one-dimensional (1D) stable manifold of a saddle point becomes attracting to its whole 2D unstable manifold. This type of homoclinic bifurcation, and the complex behavior that it can produce, have not previously been examined in detail. We derive a 2D flow-defined map for this situation and show how the map transforms a disk-shaped cross-section of the flow into an annulus. Preliminary investigations of the stable dynamics of this map show that it produces an interesting cascade of alternating pitchfork and boundary collision bifurcations. Received 24 June 1999 and Received in final form 17 February 2000     

兴奋性自突触引起神经簇放电频率降低或增加的非线性机制

兴奋和抑制性作用分别会增强和压制神经电活动,这是神经调控的通常观念,在神经信息处理中起重要作用.本文选取了放电簇和阈下振荡相交替、放电簇谷值小于阈下振荡谷值的Homoclinic/Homoclinic型簇放电,研究发现时滞和强度合适的兴奋性自突触电流作用在放电簇的谷值附近时,能引起簇内放电个数降低,并进而导致平均放电频率降低,这是不同于通常观念的新现象.进一步,用快慢变量分离获得的分岔和相轨迹,揭示了阈下振荡和放电簇分别对应快子系统的阈下和阈上极限环,兴奋性自突触电流引起阈上极限环向阈下极限环的转迁导致放电提前结束是频率降低原因.并与近期在Fold/Homoclinic簇放电报道的兴奋性自突触诱发的簇内放电个数降低但放电频率增加的现象和机制进行了比较.研究结果丰富了神经电活动的反常现象并揭示了背后的非线性机制,给出了调控簇放电的新手段,揭示了兴奋性自突触的潜在功能.     

Burst synchronization of electrically and chemically coupled map-based neurons

Burst synchronization and burst dynamics of a system consisting of two map-based neurons coupled through electrical or chemical synapses are discussed. Some basic characteristic quantities are introduced to describe burst synchronization and burst dynamics of neurons. It is observed that excitatory coupling leads to in-phase burst synchronization but inhibitory coupling results in anti-phase one. By using the basic characteristics of burst dynamics, the effects of the intrinsic bursting properties and the coupling schemes on complex bursting behaviors are also presented for both inhibitory and excitatory couplings. The results are instructive to identify bursting behaviors through experimental data.     

Transition between tonic spiking and bursting in a neuron model via the blue-sky catastrophe

We study a continuous and reversible transition between periodic tonic spiking and bursting activities in a neuron model. It is described as the blue-sky catastrophe, which is a homoclinic bifurcation of a saddle-node periodic orbit of codimension one. This transition constitutes a biophysically plausible mechanism for the regulation of burst duration that increases with no bound like 1/square root alpha-alpha0 as the transition value alpha0 is approached.     

多平衡态下簇发振荡产生机理及吸引子结构分析

以周期激励下受控Lorenz模型为例, 考察了多平衡态共存下激励频率与系统固有频率之间存在量级差距也即存在频域上的不同尺度时的耦合效应. 由于激励频率远小于系统的固有频率, 因此将整个激励项视为慢变参数, 分析随慢变参数变化下的各种分岔模式及其相应的分岔行为, 指出在一定条件下, 不同平衡点会产生Hopf分岔和fold分岔. 根据分岔条件的不同, 给出了两种典型情况下的簇发振荡, 并通过引入转换相图, 揭示了不同簇发的产生机理, 指出多平衡态和多种分岔共存不仅会导致沉寂态和激发态的多样性, 而且会使得不同沉寂态和激发态之间存在着不同的转换形式.