兴奋性和抑制性自反馈压制靠近Hopf分岔的神经电活动比较

突触输入刺激神经元产生的电活动,在神经编码中发挥着重要作用.通常认为,兴奋性输入增强电活动,抑制性输入压制电活动.本文选取可调节电流衰减速度的突触模型,研究了兴奋性自突触在亚临界Hopf分岔附近压制神经元电活动的反常作用,与抑制性自突触的压制作用进行了比较,并采用相位响应曲线和相平面分析解释了压制作用的机制.对于单稳的峰放电,快速和中速衰减的兴奋性自突触分别可以诱发频率降低的峰放电和混合振荡(峰放电与阈下振荡的交替),而中速和慢速衰减的抑制性自突触也可以分别诱发频率降低的峰放电和混合振荡.对于与静息共存的峰放电,除上述两种行为外,中速衰减的兴奋性和慢速衰减的抑制性自突触还可以诱发静息.兴奋性和抑制性自突触电流在不同的衰减速度下,分别作用在峰放电的不同相位,才能诱发同类压制行为.结果丰富了兴奋性突触压制电活动反常作用的实例,获得了兴奋性和抑制性自突触压制作用机制的不同,给出了调控神经放电的新手段.     

基于抑制性突触可塑性的神经元放电率自稳态机制

神经元放电率自稳态是指大脑神经网络的放电率维持在相对稳定的状态.大量实验研究发现神经元放电率自稳态是神经电活动的重要特征,并且放电率自稳态是实现神经信息处理及维持正常脑功能的基础,因此放电率自稳态的研究是神经科学领域的重要科学问题.脑神经网络是一个高度复杂的动态系统,存在大量输入扰动信号及由于动态链接导致的参数摄动,因此如何建立并维持神经元放电率自稳态及其鲁棒性仍有待深入研究.反馈神经回路是皮层神经网络的典型连接模式,抑制性突触可塑性对神经元放电率自稳态具有重要的调控作用.本文通过构建包含抑制性突触可塑性的反馈神经回路模型对神经元放电率自稳态及其鲁棒性进行计算研究.结果表明:在抑制性突触可塑性的作用下,神经元放电率可自适应地跟踪目标放电率,从而取得放电率自稳态;在有外部输入干扰和参数摄动的情况下,神经元放电率具有良好的抗扰动性能,表明放电率自稳态具有很强的鲁棒性;理论分析揭示了抑制性突触可塑性学习规则是神经元放电率自稳态的神经机制;仿真分析进一步揭示了学习率及目标放电率对放电率自稳态建立过程具有重要影响.     

利用相位响应曲线解释抑制性反馈增强神经电活动

在众多实验和理论研究中已经发现自突触通过自反馈电流调节神经元电活动和网络时空行为来实现生理功能.本文通过理论研究,发现在一些合适的时滞下,抑制性自反馈电流能引起放电频率增加,这是不同于传统结果—抑制性作用引起频率降低的新发现.进一步,对于没有自反馈的神经元,发现在作用相位合适的抑制性脉冲电流的作用下,放电的相位会提前,导致放电频率增加,这就表现出对应Hopf分岔的II型相位响应曲线的特征.引起放电频率增加的抑制性脉冲刺激的相位与自反馈的时滞相对应,这也就给出了自反馈能够引起放电频率增强的原因.最后,发现抑制性自反馈的时滞较长或耦合强度较大时,噪声诱发的神经元放电峰-峰间期的变异系数较小,也就是放电精确性提高,与实验发现的慢抑制性自突触诱发放电精确性增加的结果相一致.研究结果揭示了负反馈能增强系统响应这一新现象和相应的非线性动力学机制,提供了调控神经电活动的新手段,有助于认识现实神经系统的自突触的潜在功能.     

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.     

快自突触反馈诱发混合簇放电的反常变化及分岔机制

簇放电是神经系统复杂的、多时间尺度的非线性现象,具有多样性,在兴奋性或抑制性作用下实现生理功能.近期较多研究发现了与通常概念(抑制性作用引起电活动降低、兴奋性作用引起放电增强)相反的现象,丰富了非线性科学的内涵.本文关注于抑制性和兴奋性自突触反馈都会诱发的一类复杂的混合簇放电产生的反常现象及其分岔机制.利用快慢变量分离,确认了放电的复杂之处:簇结束于极限环的鞍结分岔之后要先经过去极化阻滞才到休止期.进一步,揭示了该鞍结分岔在反常现象的产生中起到了关键作用.抑制性自反馈引起了该分岔的左移导致簇的参数范围变宽,引起簇内峰个数增多和平均放电频率增加;而兴奋性自突触则引起该分岔右移导致电活动降低.与其他类簇放电只在抑制性自反馈下产生反常现象和慢突触诱发的反常现象不同,该结果给出了簇放电的反常现象的新示例及调控机制,展示了反常现象的多样性,有助于认识脑神经元簇放电和自反馈调控的潜在功能.     

具有时滞的抑制性自突触诱发的神经放电的加周期分岔

神经放电节律在神经系统功能实现中起着重要的作用.具有自突触(起始和结束于同一细胞的突触)的神经元普遍存在于神经系统,本文研究了单神经元模型在抑制性自突触作用下的放电节律.结果发现,随着时滞和/或耦合强度的增加,可以诱发Rulkov神经元模型放电节律的加周期分岔.随着放电节律的周期数的增加,平均放电频率增大,当时滞和/或耦合强度大于某一阈值时,频率大于没有自突触时的放电频率.用快慢变量分离方法可以获得没有自突触的神经放电节律的分岔结构,可用于认识外界负向脉冲诱发的新节律.这些新的节律模式与加周期分岔中的节律模式一致.研究结果不仅揭示了抑制性自突触可以诱发典型的非线性现象——加周期分岔,还给出了抑制性自突触可以提高放电频率的新现象,与以前的自突触压制放电的观点不同,进一步丰富了对抑制性自突触诱发的非线性现象的认识.     

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.     

两个非耦合Hindmarsh-Rose神经元同步的非线性特征研究

利用Hindmarsh-Rose(HR)神经元输出的膜电压作为刺激调整两个具有不同初始条件的非耦合HR神经元的电流输入，通过分析神经元放电峰峰间期(ISI)的分布揭示了两个神经元同步过程轨道演化的机理.在周期信号刺激下，两个具有相同参数原处于混沌状态的神经元可以 实现完全同步，且可以同步到不同于刺激信号频率的周期响应上；两个具有不同参数的神经 元可以实现相位同步，参数差别较小的两个神经元可以相位同步到与刺激信号不同频率的周 期响应上，参数差别较大的两个神经元只可能相位同步到与刺激信号相同频率的周期响应上 .混沌信号刺激两个神经元只可能同步到产生混沌信号神经元的放电模式上，可见混沌刺激 更有利于神经元信息编码与解码.分析两个被调整神经元系统的最大条件Lyapunov 指数(Lmc )与刺激强度k的关系表明当k达到某一阈值时两个系统的Lmc均为负值是两个系统实现同 步的必要条件.平均发放率相同的混沌刺激和周期刺激相比较混沌刺激更容易使两个神经元 实现同步，表明混沌刺激产生的效应更强，该结论与实验结果相符合. 关键词： 放电峰峰间期 同步 相位同步 条件Lyapunov 指数     

忆阻Rulkov神经网络同步研究

研究电突触、化学突触以及两者共存对忆阻Rulkov神经模型集体动力学行为的影响。对于两个忆阻Rulkov神经元系统,各种耦合方式都能使系统实现同步。对于不同的耦合强度,神经元呈现不同的放电模式,如方波,三角波,脉冲放电等。当电突触、化学突触同时存在时,系统的同步更依赖于电耦合强度。对全局耦合忆阻Rulkov神经网络同步的研究表明：化学突触单独作用时,同步发生在耦合参数的某个区域范围,当化学耦合强度超过某一阈值时,同步会随着耦合强度的增加而被破坏。电突触单独作用时,系统很快到达同步状态,并且电耦合强度是决定神经元处于静止还是峰放电的关键因素,随着电耦合强度增加,神经元放电频率、振幅增大。当电、化学耦合同时存在时,耦合强度的增加使神经元由静息转变为圆弧放电,并进入同步状态。本文提供了一种通过调整耦合方式和耦合强度,控制神经网络放电模式及其同步的可能方法。     

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。     

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.     

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.     

Control of firing activities in thermosensitive neuron by activating excitatory autapse

Temperature has distinct influence on the activation of ion channels and the excitability of neurons, and careful change in temperature can induce possible mode transition in the neural activities. The formation and development of autapse connection to neuron can enhance its self-adaption to external stimulus, and thus the firing patterns in neuron can be controlled effectively. The autapse is activated to drive a thermosensitive neuron, which is developed from the FitzHugh–Nagumo neural circuit by incorporating a thermistor, and the dynamics in the neural activities is explored to find mode dependence on the temperature and autaptic current. It is found that the firing modes can be controlled by temperature, and the neuron is wakened from resting state to periodic oscillation with the increase of temperature. Furthermore, the intensity and the intrinsic time delay in the autapse are respectively adjusted to control the neural activities, and it is confirmed that appropriate setting for autaptic current can balance and enhance the temperature effect on the neural activities.     

Negative self-feedback induced enhancement and transition of spiking activity for class-3 excitability

Post-inhibitory rebound (PIR) spike, which has been widely observed in diverse nervous systems with different physiological functions and simulated in theoretical models with class-2 excitability, presents a counterintuitive nonlinear phenomenon in that the inhibitory effect can facilitate neural firing behavior. In this study, a PIR spike induced by inhibitory stimulation from the resting state corresponding to class-3 excitability that is not related to bifurcation is simulated in the Morris-Lecar neuron. Additionally, the inhibitory self-feedback mediated by an autapse with time delay can evoke tonic/repetitive spiking from phasic/transient spiking. The dynamical mechanism for the PIR spike and the tonic/repetitive spiking is acquired with the phase plane analysis and the shape of the quasi-separatrix curve. The result extends the counterintuitive phenomenon induced by inhibition to class-3 excitability, which presents a potential function of inhibitory autapse and class-3 neuron in many neuronal systems such as the auditory system.     

Autapse-induced target wave,spiral wave in regular network of neurons

Autapse is a type of synapse that connects axon and dendrites of the same neuron, and the effect is often detected by close-loop feedback in axonal action potentials to the owned dendritic tree. An artificial autapse was introduced into the Hindmarsh-Rose neuron model, and a regular network was designed to detect the regular pattern formation induced by autapse. It was found that target wave emerged in the network even when only a single autapse was considered. By increasing the(autapse density) number of neurons with autapse, for example, a regular area(2×2, 3×3, 4×4, 5×5 neurons) under autapse induced target wave by selecting the feedback gain and time-delay in autapse. Spiral waves were also observed under optimized feedback gain and time delay in autapses because of coherence-like resonance in the network induced by some electric autapses connected to some neurons. This confirmed that the electric autapse has a critical role in exciting and regulating the collective behaviors of neurons by generating stable regular waves(target waves, spiral waves) in the network. The wave length of the induced travelling wave(target wave, spiral wave), because of local effect of autapse, was also calculated to understand the waveprofile in the network of neurons.     

Autaptic modulation-induced neuronal electrical activities and wave propagation on network under electromagnetic induction

Based on an improved HR neuron model, the effects of electrical and chemical autapses on the firing activities of single neurons are studied, and the wave propagation in forward feedback neural network is also discussed by considering autapstic regulation under different intensities of electromagnetic induction. It is found that the electrical activities of single neuron can be changed by exerting excitatory or inhibitory of electrical and chemical autapses. With different feedback gains of electromagnetic induction current, membrane potential shows the oscillatory solutions and steady states. Under the condition of different autapse or electromagnetic induction, the propagation of electrical activities caused by the central neuron is transformed in the forward feedback network. Moreover, the spatial synchronization of the network will be changed by choosing different coupling intensities and feedback gains. It is proved that the electrical and chemical autapses play a significant role in firing modes of single neuron and the wave propagation of the forward feedback networks under the electromagnetic induction.     

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.     

Dynamical origin of independent spiking and bursting activity in neural microcircuits

The relationship between spiking and bursting dynamics is a key question in neuroscience, particularly in understanding the origins of different neural coding strategies and the mechanisms of motor command generation and neural circuit coordination. Experiments indicate that spiking and bursting dynamics can be independent. We hypothesize that different mechanisms for spike and burst generation, intrinsic neuron dynamics for spiking and a modulational network instability for bursting, are the origin of this independence. We tested the hypothesis in a detailed dynamical analysis of a minimal inhibitory neural microcircuit (motif) of three reciprocally connected Hodgkin-Huxley neurons. We reduced this high-dimensional dynamical system to a rate model and showed that both systems have identical bifurcations from tonic spiking to burst generation, which, therefore, does not depend on the details of spiking activity.