一类自突触作用下神经元电路的设计和模拟

神经元在自突触作用下可以诱发各类放电活动的迁移, 神经元动作电位对电自突触的响应比较敏感. 通常用包含延迟因子和增益的反馈回路电流来刻画自突触对神经元动作电位的影响. 基于Pspice软件, 设计了包含自突触效应的神经元电路, 用以延迟反馈电路来模拟电自突触对电位的调制作用. 研究结果发现: 1)在外界刺激和电自突触回路协同作用下, 神经元电路输出信号可以呈现静息态, 尖峰放电, 簇放电状态. 2)在时变增大的外界刺激下和自突触回路驱动下, 神经元电路的输出电位序列在多种电活动模式之间(静息, 尖峰放电, 簇放电)交替出现, 其机理在于自突触回路具有记忆特性, 神经元对于不同的外界刺激可以做出不同模式的响应. 3)在给定比较大外界刺激下, 改变反馈回路的增益, 发现电路输出的序列也可以呈现不同模式交替, 即神经元对于相同的刺激可以通过自我调节自突触增益来产生不同模式的响应, 其机理可能在于回路的有效反馈, 这有助于理解突触的可塑性.     

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.     

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.     

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.     

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.     

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.     

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.     

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.