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.     

Synchronization transition in gap-junction-coupled leech neurons

Real neurons can exhibit various types of firings including tonic spiking, bursting as well as silent state, which are frequently observed in neuronal electrophysiological experiments. More interestingly, it is found that neurons can demonstrate the co-existing mode of stable tonic spiking and bursting, which depends on initial conditions. In this paper, synchronization in gap-junction-coupled neurons with co-existing attractors of spiking and bursting firings is investigated as the coupling strength gets increased. Synchronization transitions can be identified by means of the bifurcation diagram and the correlation coefficient. It is illustrated that the coupled neurons can exhibit different types of synchronization transitions between spiking and bursting when the coupling strength increases. In the course of synchronization transitions, an intermittent synchronization can be observed. These results may be instructive to understand synchronization transitions in neuronal systems.     

Complex transitions between spike,burst or chaos synchronization states in coupled neurons with coexisting bursting patterns

We investigated the synchronization dynamics of a coupled neuronal system composed of two identical Chay model neurons. The Chay model showed coexisting period-1 and period-2 bursting patterns as a parameter and initial values are varied. We simulated multiple periodic and chaotic bursting patterns with non-(NS), burst phase(BS), spike phase(SS),complete(CS), and lag synchronization states. When the coexisting behavior is near period-2 bursting, the transitions of synchronization states of the coupled system follows very complex transitions that begins with transitions between BS and SS, moves to transitions between CS and SS, and to CS. Most initial values lead to the CS state of period-2 bursting while only a few lead to the CS state of period-1 bursting. When the coexisting behavior is near period-1 bursting, the transitions begin with NS, move to transitions between SS and BS, to transitions between SS and CS, and then to CS. Most initial values lead to the CS state of period-1 bursting but a few lead to the CS state of period-2 bursting. The BS was identified as chaos synchronization. The patterns for NS and transitions between BS and SS are insensitive to initial values. The patterns for transitions between CS and SS and the CS state are sensitive to them. The number of spikes per burst of non-CS bursting increases with increasing coupling strength. These results not only reveal the initial value- and parameterdependent synchronization transitions of coupled systems with coexisting behaviors, but also facilitate interpretation of various bursting patterns and synchronization transitions generated in the nervous system with weak coupling strength.     

Bursting synchronization in clustered neuronal networks

Neuronal networks in the brain exhibit the modular (clustered) property, i.e., they are composed of certain subnetworks with differential internal and external connectivity. We investigate bursting synchronization in a clustered neuronal network. A transition to mutual-phase synchronization takes place on the bursting time scale of coupled neurons, while on the spiking time scale, they behave asynchronously. This synchronization transition can be induced by the variations of inter- and intra- coupling strengths, as well as the probability of random links between different subnetworks. Considering that some pathological conditions are related with the synchronization of bursting neurons in the brain, we analyze the control of bursting synchronization by using a time-periodic external signal in the clustered neuronal network. Simulation results show a frequency locking tongue in the driving parameter plane, where bursting synchronization is maintained, even in the presence of external driving. Hence, effective synchronization suppression can be realized with the driving parameters outside the frequency locking region.     

Chaotic phase synchronization in small-world networks of bursting neurons

We investigate the chaotic phase synchronization in a system of coupled bursting neurons in small-world networks. A transition to mutual phase synchronization takes place on the bursting time scale of coupled oscillators, while on the spiking time scale, they behave asynchronously. It is shown that phase synchronization is largely facilitated by a large fraction of shortcuts, but saturates when it exceeds a critical value. We also study the external chaotic phase synchronization of bursting oscillators in the small-world network by a periodic driving signal applied to a single neuron. It is demonstrated that there exists an optimal small-world topology, resulting in the largest peak value of frequency locking interval in the parameter plane, where bursting synchronization is maintained, even with the external driving. The width of this interval increases with the driving amplitude, but decrease rapidly with the network size. We infer that the externally applied driving parameters outside the frequency locking region can effectively suppress pathologically synchronized rhythms of bursting neurons in the brain.     

Chaotic phase synchronization in a modular neuronal network of small-world subnetworks

We investigate the onset of chaotic phase synchronization of bursting oscillators in a modular neuronal network of small-world subnetworks. A transition to mutual phase synchronization takes place on the bursting time scale of coupled oscillators, while on the spiking time scale, they behave asynchronously. It is shown that this bursting synchronization transition can be induced not only by the variations of inter- and intra-coupling strengths but also by changing the probability of random links between different subnetworks. We also analyze the effect of external chaotic phase synchronization of bursting behavior in this clustered network by an external time-periodic signal applied to a single neuron. Simulation results demonstrate a frequency locking tongue in the driving parameter plane, where bursting synchronization is maintained, even with the external driving. The width of this synchronization region increases with the signal amplitude and the number of driven neurons but decreases rapidly with the network size. Considering that the synchronization of bursting neurons is thought to play a key role in some pathological conditions, the presented results could have important implications for the role of externally applied driving signal in controlling bursting activity in neuronal ensembles.     

Hindmarsh-Rose神经元的随机自共振和噪声影响下的同步

研究了噪声对Hindmarsh-Rose(HR)神经元随机自共振和同步的影响。将高斯白噪声加入HR神经元模型的膜电位上，把外界直流电作为分岔参数，分别考虑参数处于Hopf分岔前、Hopf分岔附近和Hopf分岔后时，噪声影响下的随机自共振现象。两个未经耦合的全同HR神经元，如果接受相同的噪声激励，只要噪声强度高于某临界值，就能达到完全同步。进一步，噪声能够增强弱耦合神经元的完全同步。数值结果表明簇放电的神经元比峰放电的神经元更容易被噪声诱导而达到完全同步，耦合也增强了神经元对噪声激励的灵敏度。     

Coherence resonance and synchronization of Hindmarsh-Rose neurons with noise

Noise effects on coherence resonance and synchronization of Hindmarsh-Rose (HR) neuron model are studied. The coherence resonance of a single HR neuron with Gaussian white noise added to the membrane potential is investigated in situations before, near and after the Hopf bifurcation, separately, with the external direct current as a bifurcation parameter. It is shown that even though there is no coupling between neurons, uncoupled identical HR neurons driven by a common noise can achieve complete synchronization when the noise intensity is higher than a critical value. Furthermore, noise also enhances complete synchronization of weakly coupled neurons. It is concluded that synchronization in bursting neurons is easier to be induced than in spiking ones, and coupling enhances the sensitivity of synchronization of neurons to noise stimulus.     

Ordered bursting synchronization and complex wave propagation in a ring neuronal network

Ordered bursting synchronization and complex propagation are investigated for a ring neuronal network in which each neuron exhibits chaotic bursting behaviour. The neurons become more and more synchronous in chaotic bursting as the synaptic strength is increased. It is shown that excitatory chemical synapses can effectively tame the chaos, and ordered bursting synchronization can be observed as the synaptic strength is further increased. However, synchronization among neurons is weakened as the number of neurons is increased. More importantly, it is shown that ordered bursting synchronization can be turned into spiking synchronization at certain noise intensity. Complex spatio-temporal patterns propagating towards both sides of pacemaker are found in this network before the emergence of spiking synchronization.     

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.     

耦合Hindmarsh-Rose神经元的放电模式和完全同步

通过数值模拟和分岔分析的方法研究了Hindmarsh-Rose（HR）神经元的放电模式。当外加直流激励变化时，单个的神经元表现为静息态、周期性峰放电、周期性簇放电以及混沌的放电模式。利用快慢动力学分析的方法研究了HR神经元的动力学行为。当每个神经元表现为静息态、周期性放电和混沌时，两个耦合的神经元在一定的耦合强度下均会达到完全同步。神经元的耦合方式模拟神经元之间缝隙连接的电耦合。理论分析了完全同步的判断准则，并给出相应的数值模拟结果。电耦合HR神经元耦合系统的峰峰间期的分岔结构在耦合的作用下仍然能保持未耦合时的分岔结构。     

耦合不连续系统同步转换过程中的多吸引子共存

本文研究了耦合不连续系统的同步转换过程中的动力学行为, 发现由混沌非同步到混沌同步的转换过程中特殊的多吸引子共存现象. 通过计算耦合不连续系统的同步序参量和最大李雅普诺夫指数随耦合强度的变化, 发现了较复杂的同步转换过程: 临界耦合强度之后出现周期非同步态(周期性窗口); 分析了系统周期态的迭代轨道,发现其具有两类不同的迭代轨道: 对称周期轨道和非对称周期轨道, 这两类周期吸引子和同步吸引子同时存在, 系统表现出对初值敏感的多吸引子共存现象. 分析表明, 耦合不连续系统中的周期轨道是由于局部动力学的不连续特性和耦合动力学相互作用的结果. 最后, 对耦合不连续系统的同步转换过程进行了详细的分析, 结果表明其同步呈现出较复杂的转换过程.     

Bursting synchronization in non-locally coupled maps

Neuron activity presents two timescales, a fast one related to action-potential spiking, and a slow timescale in which bursting takes place. Bursting activity in neuron ensembles can be synchronized, meaning the adjustment of the bursting phases due to coupling. We investigated bursting synchronization in a non-locally coupled lattice using a two-dimensional map to describe neuron activity. The coupling involves all sites in a lattice, the corresponding strength decreasing with the lattice distance in a power-law fashion. We observed bursting synchronization for wide intervals of the coupling parameters. We also investigated the bursting synchronization of the ensemble with an external time-periodic signal applied to one or more selected neurons.     

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.     

Forced bursting and transition mechanism in CO oxidation with three time scales

The mathematical model of CO oxidation with three time scales on platinum group metals is investigated, in which order gaps between the time scales related to external perturbation and the rates associated with different chemical reaction steps exist. Forced bursters, such as point-point type forced bursting and point-cycle type forced bursting, are presented. The bifurcation mechanism of forced bursting is novel, and the phenomenon where two different kinds of spiking states coexist in point-cycle type forced bursting has not been reported in previous work. A double-parameter bifurcation set of the fast subsystem is explored to reveal the transition mechanisms of different forced bursters with parameter variation.     

Taming desynchronized bursting with delays in the Macaque cortical network

Inhibitory coupled bursting Hindmarsh-Rose neurons are considered as constitutive units of the Macaque cortical network. In the absence of information transmission delay the bursting activity is desynchronized,giving rise to spatiotemporally disordered dynamics. This paper shows that the introduction of finite delays can lead to the synchronization of bursting and thus to the emergence of coherent propagating fronts of excitation in the space-time domain. Moreover,it shows that the type of synchronous bursting is uniquely determined by the delay length,with the transitions from one type to the other occurring in a step-like manner depending on the delay. Interestingly,as the delay is tuned close to the transition points,the synchronization deteriorates,which implies the coexistence of different bursting attractors. These phenomena can be observed by different but fixed coupling strengths,thus indicating a new role for information transmission delays in realistic neuronal networks.     

Transition to complete synchronization via near-synchronization in two coupled chaotic neurons

The synchronization transition in two coupled chaotic Morris-Lecar （ML） neurons with gap junction is studied with the coupling strength increasing. The conditional Lyapunov exponents, along with the synchronization errors are calculated to diagnose synchronization of two coupled chaotic ML neurons. As a result, it is shown that the increase in the coupling strength leads to incoherence, then induces a transition process consisting of three different synchronization states in succession, namely, burst synchronization, near-synchronization and embedded burst synchronization, and achieves complete synchronization of two coupled neurons finally. These sequential transitions to synchronization reveal a new transition route from incoherence to complete synchronization in coupled systems with multi-time scales.     

“Hopf/homoclinic”簇放电和“SubHopf/homoclinic"簇放电之间的同步

以修正过的Morris-Lecar神经元模型为例,讨论了“Hopf/homoclinic”簇放电和“SubHopf/homoclinic"簇放电之间的同步行为.首先,分别考察了同一拓扑类型的两个耦合簇放电神经元的同步行为,发现“Hopf/homoclinic”簇放电比“SubHopf/homoclinic”簇放电达到膜电位完全同步所需要的耦合强度小,即前者比后者更容易达到膜电位完全同步.其次,对这两个不同拓扑类型的簇放电神经元的耦合同步行为进行了讨论.通过数值分析发现随着耦合强度的增加,两种不同类型的簇放电首先达到簇放电同步,然后当耦合强度足够大时甚至可以达到膜电位完全同步,并且同步后的放电类型更接近容易同步的簇放电类型,即“Hopf/homoclinic”簇放电.然而令人奇怪的是此时慢变量并没有达到完全同步,而是相位同步;慢变量之间呈现为一种线性关系.这一点和现有文献的结果截然不同.     

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.     

Spatiotemporal patterns and chaotic burst synchronization in a small-world neuronal network

The spatiotemporal patterns and chaotic burst synchronization of a small-world neuronal network are studied in this paper. The synchronization parameter, similarity parameter and order parameter are introduced to investigate the dynamics behaviour of the neurons. Chaotic burst synchronization and nearly complete synchronization can be observed if the link probability and the coupling strength are large enough. It is found that with increasing link probability and the coupling strength chaotic bursts become appreciably synchronous in space and coherent in time, and the maximal spatiotemporal order appears at some particular values of the probability and the coupling strength, respectively. The larger the size of the network, the smaller the probability and the coupling strength are needed for the network to achieve burst synchronization. Moreover, the bursting activity and the spatiotemporal patterns are robust to small noise.