Spike-timing-dependent plasticity in small-world networks

Biologically plausible excitatory neural networks develop a persistent synchronized pattern of activity depending on spontaneous activity and synaptic refractoriness (short term depression). By fixed synaptic weights synchronous bursts of oscillatory activity are stable and involve the whole network. In our modeling study we investigate the effect of a dynamic Hebbian-like learning mechanism, spike-timing-dependent plasticity (STDP), on the changes of synaptic weights depending on synchronous activity and network connection strategies (small-world topology). We show that STDP modifies the weights of synaptic connections in such a way that synchronization of neuronal activity is considerably weakened. Networks with a higher proportion of long connections can sustain a higher level of synchronization in spite of STDP influence. The resulting distribution of the synaptic weights in single neurons depends both on the global statistics of firing dynamics and on the number of incoming and outgoing connections.     

Dynamics of strongly-coupled spiking neurons

We present a dynamical theory of integrate-and-fire neurons with strong synaptic coupling. We show how phase-locked states that are stable in the weak coupling regime can destabilize as the coupling is increased, leading to states characterized by spatiotemporal variations in the interspike intervals (ISIs). The dynamics is compared with that of a corresponding network of analog neurons in which the outputs of the neurons are taken to be mean firing rates. A fundamental result is that for slow interactions, there is good agreement between the two models (on an appropriately defined timescale). Various examples of desynchronization in the strong coupling regime are presented. First, a globally coupled network of identical neurons with strong inhibitory coupling is shown to exhibit oscillator death in which some of the neurons suppress the activity of others. However, the stability of the synchronous state persists for very large networks and fast synapses. Second, an asymmetric network with a mixture of excitation and inhibition is shown to exhibit periodic bursting patterns. Finally, a one-dimensional network of neurons with long-range interactions is shown to desynchronize to a state with a spatially periodic pattern of mean firing rates across the network. This is modulated by deterministic fluctuations of the instantaneous firing rate whose size is an increasing function of the speed of synaptic response.     

不同耦合类型的三维Hybrid神经元同步动力学

神经元网络的信息传递与多个神经元间的耦合同步密切相关.大量研究表明,神经元耦合系统的同步化问题是研究大脑处理信息的关键.本文基于三维混合神经元模型研究了神经元的复杂放电,该模型由Wilson模型的快子系统和H-R模型的慢子系统组成.该模型能够重现大脑皮层神经元一系列包括规则峰放电、快速峰放电和簇放电等神经动力学行为.本文基于三维混合神经元模型,探讨了三维混合神经元在电突触耦合、化学突触耦合和磁通耦合条件下的同步放电行为,模拟膜电位序列图、相位差图等探讨恒同及非恒同下耦合强度对神经元同步放电的影响.本研究将为人们进一步了解神经系统疾病的发病机制提供指导和帮助,并为神经科学领域提供可能的研究思路.     

Self-organizing dual coding based on spike-time-dependent plasticity

It has been a matter of debate how firing rates or spatiotemporal spike patterns carry information in the brain. Recent experimental and theoretical work in part showed that these codes, especially a population rate code and a synchronous code, can be dually used in a single architecture. However, we are not yet able to relate the role of firing rates and synchrony to the spatiotemporal structure of inputs and the architecture of neural networks. In this article, we examine how feedforward neural networks encode multiple input sources in the firing patterns. We apply spike-time-dependent plasticity as a fundamental mechanism to yield synaptic competition and the associated input filtering. We use the Fokker-Planck formalism to analyze the mechanism for synaptic competition in the case of multiple inputs, which underlies the formation of functional clusters in downstream layers in a self-organizing manner. Depending on the types of feedback coupling and shared connectivity, clusters are independently engaged in population rate coding or synchronous coding, or they interact to serve as input filters. Classes of dual codings and functional roles of spike-time-dependent plasticity are also discussed.     

Mechanisms that modulate the transfer of spiking correlations

Correlations between neuronal spike trains affect network dynamics and population coding. Overlapping afferent populations and correlations between presynaptic spike trains introduce correlations between the inputs to downstream cells. To understand network activity and population coding, it is therefore important to understand how these input correlations are transferred to output correlations.Recent studies have addressed this question in the limit of many inputs with infinitesimal postsynaptic response amplitudes, where the total input can be approximated by gaussian noise. In contrast, we address the problem of correlation transfer by representing input spike trains as point processes, with each input spike eliciting a finite postsynaptic response. This approach allows us to naturally model synaptic noise and recurrent coupling and to treat excitatory and inhibitory inputs separately.We derive several new results that provide intuitive insights into the fundamental mechanisms that modulate the transfer of spiking correlations.     

Relating macroscopic measures of brain activity to fast, dynamic neuronal interactions

In this article we used biologically plausible simulations of coupled neuronal populations to address the relationship between phasic and fast coherent neuronal interactions and macroscopic measures of activity that are integrated over time, such as the BOLD response in functional magnetic resonance imaging. Event-related, dynamic correlations were assessed using joint peristimulus time histograms and, in particular, the mutual information between stimulus-induced transients in two populations. This mutual information can be considered as an index of functional connectivity. Our simulations showed that functional connectivity or dynamic integration between two populations increases with mean background activity and stimulus-related rate modulation. Furthermore, as the background activity increases, the populations become increasingly sensitive to the intensity of the stimulus in terms of a predisposition to transient phase locking. This reflects an interaction between background activity and stimulus intensity in producing dynamic correlations, in that background activity augments stimulus-induced coherence modulation. This is interesting from a computational perspective because background activity establishes a context that may have a profound effect on event-related interactions or functional connectivity between neuronal populations. Finally, total firing rates, which subsume both background activity and stimulus-related rate modulation, were almost linearly related to the expression of dynamic correlations over large ranges of activities. These observations show that under the assumptions implicit in our model, rate-specific metrics based on rate or coherence modulation may be different perspectives on the same underlying dynamics. This suggests that activity (averaged over all peristimulus times), as measured in neuroimaging, may be tightly coupled to the expression of dynamic correlations.     

前馈型神经元网络中的放电频率传递分析

本文建立了一个基于Hodgkin-Huxley神经元的前馈神经元网络模型,研究了平均放电频率在前馈神经元网络中的传递情况。研究结果显示,适当的层间连接概率与输人噪声强度能够提髙前馈神经元网络的同步效率,进而增强网络稳定传递放电频率的性能。此外,通过引入并调节突触时滞,发现适当的时滞对神经元耦合系统的完全同步和前馈神经元网络内信息传输有明显的促进作用。     

呼吸中间神经元及呼吸网络模型的动力学分析

呼吸节律的产生起源于Pre-Btzinger复合体,这其中包含了Pre-Btzinger神经元在内的许多种呼吸神经元的参与,这些呼吸神经元和肺通过突触联系构成了脑桥-髓质的动态呼吸网络.由于目前对于呼吸节律产生和变化的网络机制尙不完全清楚,因此,本文从非线性动力学角度入手,通过构造与实际结构比较接近的呼吸网络模型,分别考察了网络中单个Pre-Btzinger中间神经元多样性的发放模式以及网络中群体神经元周期性和同步性的放电变化.数值结果为进一步揭示呼吸节律的产生和调控机制提供了一定的帮助.     

Asynchronous states and the emergence of synchrony in large networks of interacting excitatory and inhibitory neurons

We investigate theoretically the conditions for the emergence of synchronous activity in large networks, consisting of two populations of extensively connected neurons, one excitatory and one inhibitory. The neurons are modeled with quadratic integrate-and-fire dynamics, which provide a very good approximation for the subthreshold behavior of a large class of neurons. In addition to their synaptic recurrent inputs, the neurons receive a tonic external input that varies from neuron to neuron. Because of its relative simplicity, this model can be studied analytically. We investigate the stability of the asynchronous state (AS) of the network with given average firing rates of the two populations. First, we show that the AS can remain stable even if the synaptic couplings are strong. Then we investigate the conditions under which this state can be destabilized. We show that this can happen in four generic ways. The first is a saddle-node bifurcation, which leads to another state with different average firing rates. This bifurcation, which occurs for strong enough recurrent excitation, does not correspond to the emergence of synchrony. In contrast, in the three other instability mechanisms, Hopf bifurcations, which correspond to the emergence of oscillatory synchronous activity, occur. We show that these mechanisms can be differentiated by the firing patterns they generate and their dependence on the mutual interactions of the inhibitory neurons and cross talk between the two populations. We also show that besides these codimension 1 bifurcations, the system can display several codimension 2 bifurcations: Takens-Bogdanov, Gavrielov-Guckenheimer, and double Hopf bifurcations.     

A network of coincidence detector neurons with periodic and chaotic dynamics

We propose a simple neural network model to understand the dynamics of temporal pulse coding. The model is composed of coincidence detector neurons with uniform synaptic efficacies and random pulse propagation delays. We also assume a global negative feedback mechanism which controls the network activity, leading to a fixed number of neurons firing within a certain time window. Due to this constraint, the network state becomes well defined and the dynamics equivalent to a piecewise nonlinear map. Numerical simulations of the model indicate that the latency of neuronal firing is crucial to the global network dynamics; when the timing of postsynaptic firing is less sensitive to perturbations in timing of presynaptic spikes, the network dynamics become stable and periodic, whereas increased sensitivity leads to instability and chaotic dynamics. Furthermore, we introduce a learning rule which decreases the Lyapunov exponent of an attractor and enlarges the basin of attraction.     

局部耦合的双层忆阻神经元网络的嵌合态和同步

嵌合态被发现存在于神经系统并且可能在神经元节律、大脑的睡眠和记忆等诸多神经过程中发挥重要作用.本文考虑神经元交互中的电磁感应现象,建立了以Hindmarsh Rose神经元为节点的局部耦合的双层忆阻神经元网络,研究其嵌合态时空动力学模式及产生机理.结果发现,改变层内、层间突触耦合强度会使网络产生移动和不完美移动嵌合态等多种类型的嵌合模式,其中不完美移动嵌合态中不相干的区域会扩展到网络的相干域.特别地,在特定耦合强度下,存在一种新的嵌合态活动模式,即一部分神经元处于嵌合态,另一部分神经元处于移动嵌合态.考虑神经元突触的忆阻特性,发现忆阻参数的增加能够使处于嵌合态的神经元网络转变为同步态,且耦合强度越大,达到同步态所需要的忆阻参数值越小.进一步探究双层网络的同步性,发现层间耦合强度和忆阻参数的增大有助于网络达到更好的同步.研究结果表明神经元之间的相互作用可以激发双层神经元网络产生多种嵌合态模式,电磁感应可以促进网络由嵌合态向同步态转迁,这些结果有助于理解人脑中复杂的神经放电过程和信息处理机制,并为可能的类脑装置应用提供参考.     

耦合Hindmarsh-Rose神经元的同步放电模式及转迁

通过数值模拟和分岔分析,探究了具有不同放电模式的两电耦合Hindmarsh-Rose神经元的几乎完全同步,并研究了同步放电模式对单个个体的放电模式的依赖性.研究结果有助于我们更好地理解神经元放电模式转迁的动力学机理和生物学意义.     

The number of synaptic inputs and the synchrony of large, sparse neuronal networks

The prevalence of coherent oscillations in various frequency ranges in the central nervous system raises the question of the mechanisms that synchronize large populations of neurons. We study synchronization in models of large networks of spiking neurons with random sparse connectivity. Synchrony occurs only when the average number of synapses, M, that a cell receives is larger than a critical value, Mc. Below Mc, the system is in an asynchronous state. In the limit of weak coupling, assuming identical neurons, we reduce the model to a system of phase oscillators that are coupled via an effective interaction, gamma. In this framework, we develop an approximate theory for sparse networks of identical neurons to estimate Mc analytically from the Fourier coefficients of gamma. Our approach relies on the assumption that the dynamics of a neuron depend mainly on the number of cells that are presynaptic to it. We apply this theory to compute Mc for a model of inhibitory networks of integrate-and-fire (I&F) neurons as a function of the intrinsic neuronal properties (e.g., the refractory period Tr), the synaptic time constants, and the strength of the external stimulus, Iext. The number Mc is found to be nonmonotonous with the strength of Iext. For Tr = 0, we estimate the minimum value of Mc over all the parameters of the model to be 363.8. Above Mc, the neurons tend to fire in smeared one-cluster states at high firing rates and smeared two-or-more-cluster states at low firing rates. Refractoriness decreases Mc at intermediate and high firing rates. These results are compared to numerical simulations. We show numerically that systems with different sizes, N, behave in the same way provided the connectivity, M, is such that 1/Meff = 1/M - 1/N remains constant when N varies. This allows extrapolating the large N behavior of a network from numerical simulations of networks of relatively small sizes (N = 800 in our case). We find that our theory predicts with remarkable accuracy the value of Mc and the patterns of synchrony above Mc, provided the synaptic coupling is not too large. We also study the strong coupling regime of inhibitory sparse networks. All of our simulations demonstrate that increasing the coupling strength reduces the level of synchrony of the neuronal activity. Above a critical coupling strength, the network activity is asynchronous. We point out a fundamental limitation for the mechanisms of synchrony relying on inhibition alone, if heterogeneities in the intrinsic properties of the neurons and spatial fluctuations in the external input are also taken into account.     

网络噪声和振子数量对同步化行为的影响

选取了三个反映同步化程度的指标平均向量场、同步因子和放电概率,数值模拟研究了网络噪声和振子数量对同步化行为的影响.随着噪声强度的增大,三个指标都出现了先增加再降低的现象,即发生了相干共振.在不同的耦合强度和噪声强度下,三个同步化指标随着振子数量的增加都呈现出了降低的趋势,表明了网络同步化行为的减弱.研究结果对如何利用噪...     

An exact method to quantify the information transmitted by different mechanisms of correlational coding

We derive a new method to quantify the impact of correlated firing on the information transmitted by neuronal populations. This new method considers, in an exact way, the effects of high order spike train statistics, with no approximation involved, and it generalizes our previous work that was valid for short time windows and small populations. The new technique permits one to quantify the information transmitted if each cell were to convey fully independent information separately from the information available in the presence of synergy-redundancy effects. Synergy-redundancy effects are shown to arise from three possible contributions: a redundant contribution due to similarities in the mean response profiles of different cells; a synergistic stimulus-dependent correlational contribution quantifying the information content of changes of correlations with stimulus, and a stimulus-independent correlational contribution term that reflects interactions between the distribution of rates of individual cells and the average level of cross-correlation. We apply the new method to simultaneously recorded data from somatosensory and visual cortices. We demonstrate that it constitutes a reliable tool to determine the role of cross-correlated activity in stimulus coding even when high firing rate data (such as multi-unit recordings) are considered.     

Dual information representation with stable firing rates and chaotic spatiotemporal spike patterns in a neural network model.

Although various means of information representation in the cortex have been considered, the fundamental mechanism for such representation is not well understood. The relation between neural network dynamics and properties of information representation needs to be examined. We examined spatial pattern properties of mean firing rates and spatiotemporal spikes in an interconnected spiking neural network model. We found that whereas the spatiotemporal spike patterns are chaotic and unstable, the spatial patterns of mean firing rates (SPMFR) are steady and stable, reflecting the internal structure of synaptic weights. Interestingly, the chaotic instability contributes to fast stabilization of the SPMFR. Findings suggest that there are two types of network dynamics behind neuronal spiking: internally-driven dynamics and externally driven dynamics. When the internally driven dynamics dominate, spikes are relatively more chaotic and independent of external inputs; the SPMFR are steady and stable. When the externally driven dynamics dominate, the spiking patterns are relatively more dependent on the spatiotemporal structure of external inputs. These emergent properties of information representation imply that the brain may adopt a dual coding system. Recent experimental data suggest that internally driven and externally driven dynamics coexist and work together in the cortex.     

模块神经元网络中耦合时滞诱导的簇同步转迁

利用Courbage-Nekorkin-Vdovin神经元构建含有耦合时滞的模块神经元网络模型,通过数值模拟研究了耦合强度及耦合时滞对模块神经元网络簇同步放电特性的影响.研究结果表明,适当大的耦合强度可以诱导模块神经元网络达到簇同步.同时,研究发现耦合时滞可以诱导模块神经元网络出现簇同步转迁,且当时滞大小约为网络中所有神经元平均振荡周期的整数倍数时,模块神经元网络的簇同步现象能够间歇性出现.此外,研究结果表明时滞诱导的簇同步转迁对子网络内的耦合强度、子网络间的连接概率具有鲁棒性.     

考虑树突整合效应的神经元网络的放电和同步特性

生物神经网络的同步被认为在大脑神经信息的处理过程中发挥了重要作用.本文在Hodgkin-Huxley(HH)神经元网络模型中考虑树突整合效应,得到修正后的DHH(Dendritic-integration-rule-based HH)神经元网络模型,研究了网络的放电和同步特性.首先以三个抑制性神经元构成的耦合系统为例,发现树突整合效应的加入提高了神经元的放电阈值;然后分别建立全局耦合的抑制性和兴奋性神经元网络,发现大的耦合强度能够诱导抑制性和兴奋性神经元网络达到几乎完全同步的状态,并且对神经元的放电幅值有较大的影响;更有趣的是,当树突整合系数为某一值时,抑制性神经元网络的同步达到最高,而兴奋性神经网络的同步达到最低.     

Synchronous firing and higher-order interactions in neuron pool

The stochastic mechanism of synchronous firing in a population of neurons is studied from the point of view of information geometry. Higher-order interactions of neurons, which cannot be reduced to pairwise correlations, are proved to exist in synchronous firing. In a neuron pool where each neuron fires stochastically, the probability distribution q(r) of the activity r, which is the fraction of firing neurons in the pool, is studied. When q(r) has a widespread distribution, in particular, when q(r) has two peaks, the neurons fire synchronously at one time and are quiescent at other times. The mechanism of generating such a probability distribution is interesting because the activity r is concentrated on its mean value when each neuron fires independently, because of the law of large numbers. Even when pairwise interactions, or third-order interactions, exist, the concentration is not resolved. This shows that higher-order interactions are necessary to generate widespread activity distributions. We analyze a simple model in which neurons receive common overlapping inputs and prove that such a model can have a widespread distribution of activity, generating higher-order stochastic interactions.     

Difficulty of singularity in population coding

Fisher information has been used to analyze the accuracy of neural population coding. This works well when the Fisher information does not degenerate, but when two stimuli are presented to a population of neurons, a singular structure emerges by their mutual interactions. In this case, the Fisher information matrix degenerates, and the regularity condition ensuring the Cramér-Rao paradigm of statistics is violated. An animal shows pathological behavior in such a situation. We present a novel method of statistical analysis to understand information in population coding in which algebraic singularity plays a major role. The method elucidates the nature of the pathological case by calculating the Fisher information. We then suggest that synchronous firing can resolve singularity and show a method of analyzing the binding problem in terms of the Fisher information. Our method integrates a variety of disciplines in population coding, such as nonregular statistics, Bayesian statistics, singularity in algebraic geometry, and synchronous firing, under the theme of Fisher information.