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.     

一类新的混沌神经放电的动力学特征的实验和数学模型研究

神经元电活动理论模型Hindmarsh-Rose(HR)模型提示有位于周期1和周期2放电模式之间的一类特殊的混沌放电,但长期以来对其没有获得足够认识.依据回归映射的确定性结构和非线性预报的短期可预报性,确认了在大鼠的实验性神经起步点的实验中发现的位于周期1和周期2放电模式之间的非周期放电是混沌放电模式,还将该混沌放电模式区分为3个不同表观样式.其中1个表观形式与HR模型的仿真结果相类似,验证了HR模型的理论预期;其余2个样式与仿真结果并不相似.进一步揭示了3个表观样式的动力学特征以及相互之间的区别与联系,并与位于周期2和周期3节律之间、周期3和周期4节律之间的混沌比较了异同,也区别了从周期1到混沌再到周期2放电模式的节律转迁历程与其他的从周期1到周期2节律的分岔过程的不同.研究结果确认了该类特殊混沌节律和相应分岔过程的新特征,丰富了混沌放电节律和节律分岔序列的种类.还对仿真该混沌的多样性和非光滑特性,以及揭示该类混沌的产生途径等进行了讨论. 关键词： 混沌 神经放电模式 分岔 节律     

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.     

动态突触、神经耦合与时间延迟对神经元发放的影响

神经元膜电位的受激发放在神经系统的信息传递中起着重要作用.基于一个受动态突触刺激的突触后神经元发放模型,采用数值模拟和傅里叶变换分析的方法研究了动态突触、神经耦合与时间延迟对突触后神经元发放的影响.结果发现:突触前神经元发放频率与Hodgkin-Huxley神经元的固有频率发生共振决定了突触后神经元发放的难易,特定频率范围内的电流刺激有利于神经元激发,动态突触输出的随机突触电流中这些电流刺激所占的比率在很大程度上影响了突触后神经元的发放次数;将突触后神经元换成神经网络后,网络中神经元之间的耦合可以促进神经元的发放,耦合中的时间延迟可以增强这种促进作用,但是不会改变神经耦合对神经元发放的促进模式.     

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.     

一类位于加周期分岔中的貌似混沌的随机神经放电节律的识别

识别非周期神经放电节律是混沌还是随机一直是一个重要的科学问题. 在神经起步点实验中发现了一类介于周期k和周期k+1(k=1,2)节律之间非周期自发放电节律, 其行为是长串的周期k簇和周期k+1簇的交替. 确定性理论模型Chay模型展示出了周期k和周期k+1节律的共存行为. 噪声在共存区诱发出了与实验结果类似的非周期节律, 说明该类节律是噪声引起的两类簇的跃迁. 非线性预报及其回归映射揭示该节律具有确定性机理; 将两类簇分别转换为0和1得到一个二进制序列, 对该序列进行概率分析获得了两类簇跃迁的随机机理. 这不仅说明该节律是具有确定性结构的随机节律而不是混沌, 还为深入识别现实神经系统的混沌和随机节律提供了典型示例和有效方法.     

可兴奋性细胞混沌放电区间的识别机理

在神经起步点记录到加周期分岔过程的生理实验数据，在对此分岔过程中位于周期n爆发 和周期(n+1)爆发之间的混沌的峰峰间期数据检测不稳定的周期轨道时，发现从靠近周期 n爆发的混沌的峰峰间期数据中，可以检测出不稳定的周期n轨道；而从靠近周期(n+1)爆 发的混沌的峰峰间期数据中，不仅可以检测出不稳定的周期(n+1)轨道，还可以检测出不稳 定的周期n轨道.针对该现象，借助于Sherman建议的胰腺β细胞模型，从非线性动力 学角度给出了理论解释.指明了由鞍结分岔和倍周期分岔分别产生第一类阵发和第三类阵发 为出现该 关键词： 峰峰间期 不稳定的周期轨道 鞍结分岔 倍周期分岔     

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.     

“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.     

Modeling and dynamics of double Hindmarsh-Rose neuron with memristor-based magnetic coupling and time delay

The firing of a neuron model is mainly affected by the following factors:the magnetic field, external forcing current, time delay, etc. In this paper, a new time-delayed electromagnetic field coupled dual Hindmarsh-Rose neuron network model is constructed. A magnetically controlled threshold memristor is improved to represent the self-connected and the coupled magnetic fields triggered by the dynamic change of neuronal membrane potential for the adjacent neurons. Numerical simulation confirms that the coupled magnetic field can activate resting neurons to generate rich firing patterns, such as spiking firings, bursting firings, and chaotic firings, and enable neurons to generate larger firing amplitudes. The study also found that the strength of magnetic coupling in the neural network also affects the number of peaks in the discharge of bursting firing. Based on the existing medical treatment background of mental illness, the effects of time lag in the coupling process against neuron firing are studied. The results confirm that the neurons can respond well to external stimuli and coupled magnetic field with appropriate time delay, and keep periodic firing under a wide range of external forcing current.     

Firing activities in a fractional-order Hindmarsh-Rose neuron with multistable memristor as autapse

Considering the fact that memristors have the characteristics similar to biological synapses, a fractional-order multistable memristor is proposed in this paper. It is verified that the fractional-order memristor has multiple local active regions and multiple stable hysteresis loops, and the influence of fractional-order on its nonvolatility is also revealed. Then by considering the fractional-order memristor as an autapse of Hindmarsh-Rose (HR) neuron model, a fractional-order memristive neuron model is developed. The effects of the initial value, external excitation current, coupling strength and fractional-order on the firing behavior are discussed by time series, phase diagram, Lyapunov exponent and inter spike interval (ISI) bifurcation diagram. Three coexisting firing patterns, including irregular asymptotically periodic (A-periodic) bursting, A-periodic bursting and chaotic bursting, dependent on the memristor initial values, are observed. It is also revealed that the fractional-order can not only induce the transition of firing patterns, but also change the firing frequency of the neuron. Finally, a neuron circuit with variable fractional-order is designed to verify the numerical simulations.     

神经元电活动不同节律模式的几种变化过程

利用神经元Chay模型,对实验中观察到的三种放电节律模式序列进行数值模拟,并应用余维1极限环分岔分析研究了其产生机理.首先考虑的是周期性放电模式的变化过程;其次,具有不同表象的一种超临界和一种亚临界倍周期簇放电序列产生并导致混沌现象的出现,然后以不同的方式转迁到逆超临界倍周期峰放电序列;最后研究无混沌的加周期簇放电序列,得出加周期分岔仅是一种与倍周期分岔密切相关的分岔现象.     

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。     

电磁辐射诱发神经元放电节律转迁的动力学行为研究

本文首先根据能量转换理论建立了电磁辐射影响下神经元电流变量模型, 然后结合Hodgkin-Huxley(HH)神经元模型研究了电磁辐射对单个神经元以及耦合神经元放电行为的影响. 结果表明, 随着电磁辐射强度的增大, 神经元放电率逐渐减小, 最后达到一个比较稳定的值. 神经元原有的周期型放电由于辐射强度的增大而逐步过渡到簇放电状态, 并借助动态分岔理论解释了这种放电模式的转换. 同时证明了磁辐射对单个神经元放电的影响可以通过神经元间的耦合传递到临近其他神经元中.     

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.     

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.     

Delayed excitatory self-feedback-induced negative responses of complex neuronal bursting patterns

In traditional viewpoint, excitatory modulation always promotes neural firing activities. On contrary, the negative responses of complex bursting behaviors to excitatory self-feedback mediated by autapse with time delay are acquired in the present paper. Two representative bursting patterns which are identified respectively to be "Fold/Big Homoclinic"bursting and "Circle/Fold cycle" bursting with bifurcations are studied. For both burstings, excitatory modulation can induce less spikes per burst for suitable time delay and strength of the self-feedback/autapse, because the modulation can change the initial or termination phases of the burst. For the former bursting composed of quiescent state and burst, the mean firing frequency exhibits increase, due to that the quiescent state becomes much shorter than the burst. However, for the latter bursting pattern with more complex behavior which is depolarization block lying between burst and quiescent state, the firing frequency manifests decrease in a wide range of time delay and strength, because the duration of both depolarization block and quiescent state becomes long. Therefore, the decrease degree of spike number per burst is larger than that of the bursting period, which is the cause for the decrease of firing frequency. Such reduced bursting activity is explained with the relations between the bifurcation points of the fast subsystem and the bursting trajectory. The present paper provides novel examples of paradoxical phenomenon that the excitatory effect induces negative responses, which presents possible novel modulation measures and potential functions of excitatory self-feedback/autapse to reduce bursting activities.     

慢变控制下Chen系统的复杂行为及其机理

由于Chen系统的控制分析大都是基于同一时间尺度,而两时间尺度耦合问题的相关研究基本上局限于单维慢变量情形.本文探讨了基于慢时间尺度上的Duffing振子,即含有两维慢子系统控制下Chen系统的动力学演化过程.给出了诸如对称式fold/fold、对称式fold/Hopf、对称式homoclinic/homoclinic等不同形式的簇发振荡行为,并揭示了其相应的产生机制,指出慢子系统中两维慢变量的相互影响导致系统产生了类似于周期激励下的簇发行为.     

Different types of bursting in Chay neuronal model

Based on actual neuronal firing activities, bursting in the Chay neuronal model is considered, in which V _K, reversal potentials for K⁺, V _C, reversal potentials for Ca²⁺, time kinetic constant λ _n and an additional depolarized current I are considered as dynamical parameters. According to the number of the Hopf bifurcation points on the upper branch of the bifurcation curve of fast subsystem, which is associated with the stable limit cycle corresponding to spiking states, different types of bursting and their respective dynamical behavior are surveyed by means of fast-slow dynamical bifurcation analysis. Supported by the National Natural Science Foundation of China (Grant Nos. 10432010, 10526002 and 10702002)     

Firing Patterns and Transitions in Coupled Neurons Controlled by a Pacemaker

To reveal the dynamics of neuronal networks with pacemakers, the firing patterns and their transitions are investigated in a ring HR neuronal network with gap junctions under the control of a pacemaker. Compared with the situation without pacemaker, the neurons in the network can exhibit wrious firing patterns as the externed current is applied or the coupling strength of pacemaker varies. The results are beneficial for understanding the complex cooperative behaviour of large neural assemblies with pacemaker control.