Information encoding and computation with spikes and bursts

Neurons compute and communicate by transforming synaptic input patterns into output spike trains. The nature of this transformation depends crucially on the properties of voltage-gated conductances in neuronal membranes. These intrinsic membrane conductances can enable neurons to generate different spike patterns including brief, high-frequency bursts that are commonly observed in a variety of brain regions. Here we examine how the membrane conductances that generate bursts affect neural computation and encoding. We simulated a bursting neuron model driven by random current input signal and superposed noise. We consider two issues: the timing reliability of different spike patterns and the computation performed by the neuron. Statistical analysis of the simulated spike trains shows that the timing of bursts is much more precise than the timing of single spikes. Furthermore, the number of spikes per burst is highly robust to noise. Next we considered the computation performed by the neuron: how different features of the input current are mapped into specific output spike patterns. Dimensional reduction and statistical classification techniques were used to determine the stimulus features triggering different firing patterns. Our main result is that spikes, and bursts of different durations, code for different stimulus features, which can be quantified without a priori assumptions about those features. These findings lead us to propose that the biophysical mechanisms of spike generation enables individual neurons to encode different stimulus features into distinct spike patterns.     

Influences of synaptic location on the synchronization of rhythmic bursting neurons

We study how the location of synaptic input influences the stablex firing states in coupled model neurons bursting rhythmically at the gamma frequencies (20-70 Hz). The model neuron consists of two compartments and generates one, two, three or four spikes in each burst depending on the intensity of input current and the maximum conductance of M-type potassium current. If the somata are connected by reciprocal excitatory synapses, we find strong correlations between the changes in the bursting mode and those in the stable phase-locked states of the coupled neurons. The stability of the in-phase phase-locked state (synchronously firing state) tends to change when the individual neurons change their bursting patterns. If, however, the synaptic connections are terminated on the dendritic compartments, no such correlated changes occur. In this case, the coupled bursting neurons do not show the in-phase phase-locked state in any bursting mode. These results indicate that synchronization behaviour of bursting neurons significantly depends on the synaptic location, unlike a coupled system of regular spiking neurons.     

皮层锥体神经元模型的动力学分析

簇发放是锥体神经元的一种典型特性,在确定性的信号传递和突触可塑性方面有着很重要的功能作用,本文通过对一类可产生复杂簇发放的皮层锥体神经元房室模型的研究,从非线性动力学角度对模型所产生的复杂簇发放做了详细的分析,讨论了不同电生理参数条件下,模型簇发放中所蕴含着的丰富的动力学性质,如:峰峰间距(InterSpike Intervals,ISIs)的加周期分岔和倍周期分岔等,通过模型分析结果可进一步理解皮层锥体神经元动作电位簇发放中所蕴含的丰富的发放模式和节律编码.     

耦合Pre-Botzinger复合体中神经元的反相簇放电模式及同步研究

本文以pre-B(o)tzinger复合体中两个耦合兴奋性神经元为研究对象,分别研究了耦合神经元反相簇放电类型及其同步转迁.当钠电导参数在一定范围内变化时,耦合神经元分别表现为“sup-Hopf/fold cycle”型和“fold/fold cycle”型反相簇放电.通过计算耦合神经元的簇放电内的峰相位差,进而判定反相簇放电中的峰同步行为,并且研究了反相放电的类型与同步的关系.     

具有时滞的耦合Hindmarsh-Rose神经元系统的放电模式

基于在无时滞的情况下,非全同的HindmarshRose耦合神经元达到几乎完全同步的放电模式,通过数值模拟的方法,研究了时滞对耦合HindmarshRose神经元同步后放电模式的影响.结果表明时滞使得神经元的放电模式发生改变,同时时滞的增加能够诱导簇中的峰逐渐地减小或消失.这一研究将有助于我们更深入地了解时滞对耦合神经元系统行为的影响.     

龙虾心脏神经节模型的动力学分析

通过对龙虾心脏神经节模型的研究,从非线性动力学角度对模型所产生的簇发放做了详细的分析,讨论了不同电生理参数条件下,模型簇发放中所蕴含着的丰富的动力学性质,如：峰峰间距(InterSpike Intervals,ISIs)的加周期分岔和倍周期分岔等.通过模型分析结果可进一步理解龙虾心脏神经节动作电位簇发放中所蕴含的丰富的发放模式和节律编码.     

A-type K(+) current can act as a trigger for bursting in the absence of a slow variable

Models of bursting in single cells typically include two subsystems with different timescales. Variations in one or more slow variables switch the system between a silent and a spiking state. We have developed a model for bursting in the pituitary lactotroph that does not include any slow variable. The model incorporates fast, noninactivating calcium and potassium currents (the spike-generating mechanism), as well as the fast, inactivating A-type potassium current (I(A)). I(A) is active only briefly at the beginning of a burst, but this brief impulse of I(A) acts as a burst trigger, injecting the spike trajectory close to an unstable steady state. The spiraling of the trajectory away from the steady state produces a period of low-amplitude spiking typical of lactotrophs. Increasing the conductance of A-type potassium current brings the trajectory closer to the unstable steady state, increasing burst duration. However, this also increases interburst interval, and for larger conductance values, all activity stops. To our knowledge, this is the first example of a physiologically based, single-compartmental model of bursting with no slow subsystem.     

Patterns of synchrony in neural networks with spike adaptation

We study the emergence of synchronized burst activity in networks of neurons with spike adaptation. We show that networks of tonically firing adapting excitatory neurons can evolve to a state where the neurons burst in a synchronized manner. The mechanism leading to this burst activity is analyzed in a network of integrate-and-fire neurons with spike adaptation. The dependence of this state on the different network parameters is investigated, and it is shown that this mechanism is robust against inhomogeneities, sparseness of the connectivity, and noise. In networks of two populations, one excitatory and one inhibitory, we show that decreasing the inhibitory feedback can cause the network to switch from a tonically active, asynchronous state to the synchronized bursting state. Finally, we show that the same mechanism also causes synchronized burst activity in networks of more realistic conductance-based model neurons.     

A test of metabolically efficient coding in the retina

We tested the hypothesis that aspects of the neural code of retinal ganglion cells are optimized to transmit visual information at minimal metabolic cost. Under a broad ensemble of light patterns, ganglion cell spike trains consisted of sparse, precise bursts of spikes. These bursts were viewed as independent neural symbols. The noise in each burst was measured via repeated presentation of the visual stimulus, and the energy cost was estimated from the total charge flow during ganglion cell spiking. Given these costs and noise, the theory of efficient codes predicts an optimal distribution of symbol usage. Symbols that are either noisy or costly occur less frequently in this optimal code. We found good qualitative and quantitative agreement with the measured distribution of burst sizes for ganglion cells in the tiger salamander retina.     

神经元模型的复杂动力学：分岔与编码

研究了改进的Morris—Lecar（ML）神经元模型的放电节律模式和模式转化的峰峰间期（interspike intervals,ISIs）分岔结构,通过调节模型中的两个重要参数μ和Vk,发现对于固定的μ,改变Vk,神经元呈现出从倍周期级联分岔到加周期分岔的复杂结构,放电模式从静息态转化为周期、混沌簇放电状态;若选取此分岔过程中的某一Vk值,对μ进行调节,呈现出的ISIs分岔结构在很大程度上取决于单个神经元的放电节律模式,且单个神经元处于混沌簇放电时,肛带来的分岔动力学行为较丰富．由于神经元能够通过动作电位对信息进行编码,所以我们推测,研究神经元的放电节律模式和动作电位的ISIs分岔结构能为理解神经信息编码机制提供线索．     

Phase precession through synaptic facilitation

Phase precession is a relational code that is thought to be important for episodic-like memory, for instance, the learning of a sequence of places. In the hippocampus, places are encoded through bursting activity of so-called place cells. The spikes in such a burst exhibit a precession of their firing phases relative to field potential theta oscillations (4-12 Hz); the theta phase of action potentials in successive theta cycles progressively decreases toward earlier phases. The mechanisms underlying the generation of phase precession are, however, unknown. In this letter, we show through mathematical analysis and numerical simulations that synaptic facilitation in combination with membrane potential oscillations of a neuron gives rise to phase precession. This biologically plausible model reproduces experimentally observed features of phase precession, such as (1) the progressive decrease of spike phases, (2) the nonlinear and often also bimodal relation between spike phases and the animal's place, (3) the range of phase precession being smaller than one theta cycle, and (4) the dependence of phase jitter on the animal's location within the place field. The model suggests that the peculiar features of the hippocampal mossy fiber synapse, such as its large efficacy, long-lasting and strong facilitation, and its phase-locked activation, are essential for phase precession in the CA3 region of the hippocampus.     

不同时间尺度的慢变量引起的簇放电研究

本文基于两个重要的慢负反馈机制给出了一个组合型的胰腺β-细胞模型.在这个模型中,不同簇放电模式对快、中、慢的振荡周期具有鲁棒性,这样可以通过快振荡周期簇放电模式的快慢动力学分析得到所有簇放电的动力学机理和拓扑类型.对于快振荡周期的簇放电,较慢的慢变量α几乎为常数,较快的慢变量Cer对快子系统没有影响,因此只要考虑慢变量...     

Dynamical analysis of bursting oscillations in the Chay-Keizer model with three time scales

Based on the Chay-Keizer model with three time scales, we investigate the role of the slowest variable in generating bursting oscillations in pancreaticcells. It is shown that both of the two slow processes can interact to drive fast, medium and slow bursting oscillations typically observed in pancreaticcells. Moreover, diverse patterns of electrical bursting are presented, including the fold/fold bursting, fold/homoclinic bursting, fold/Hopf bursting via fold/fold hysteresis loop, and the fold/fold burstin...     

Parallel Mining of Neuronal Spike Streams on Graphics Processing Units

Multi-electrode arrays (MEAs) provide dynamic and spatial perspectives into brain function by capturing the temporal behavior of spikes recorded from cultures and living tissue. Understanding the firing patterns of neurons implicit in these spike trains is crucial to gaining insight into cellular activity. We present a solution involving a massively parallel graphics processing unit (GPU) to mine spike train datasets. We focus on mining frequent episodes of firing patterns that capture coordinated events even in the presence of intervening background events. We present two algorithmic strategies—hybrid mining and two-pass elimination—to map the finite state machine-based counting algorithms onto GPUs. These strategies explore different computation-to-core mapping schemes and illustrate innovative parallel algorithm design patterns for temporal data mining. We also provide a multi-GPU mining framework, which exhibits additional performance enhancement. Together, these contributions move us towards a real-time solution to neuronal data mining.     

Predicting single-neuron activity in locally connected networks

The characterization of coordinated activity in neuronal populations has received renewed interest in the light of advancing experimental techniques that allow recordings from multiple units simultaneously. Across both in vitro and in vivo preparations, nearby neurons show coordinated responses when spontaneously active and when subject to external stimuli. Recent work (Truccolo, Hochberg, & Donoghue, 2010 ) has connected these coordinated responses to behavior, showing that small ensembles of neurons in arm-related areas of sensorimotor cortex can reliably predict single-neuron spikes in behaving monkeys and humans. We investigate this phenomenon using an analogous point process model, showing that in the case of a computational model of cortex responding to random background inputs, one is similarly able to predict the future state of a single neuron by considering its own spiking history, together with the spiking histories of randomly sampled ensembles of nearby neurons. This model exhibits realistic cortical architecture and displays bursting episodes in the two distinct connectivity schemes studied. We conjecture that the baseline predictability we find in these instances is characteristic of locally connected networks more broadly considered.     

Information measure for analyzing specific spiking patterns and applications to LGN bursts

Neural spiking responses can include a variety of spiking patterns. However, neither the mere presence of the patterns nor the pattern's frequency indicates that the pattern conveys distinct stimulus information. Here, we present an in-depth analysis of a Pattern Information measure, which quantifies how informative it is to distinguish a particular pattern of spikes from either a single spike or an another pattern. (1) We show how a shuffle-controlled estimation method minimizes the impact of sampling bias. (2) We describe how the Pattern Information could arise from time-varying firing rates, and we demonstrate an analysis to determine whether Pattern Information associated with a particular pattern captures structure not contained in the time-varying firing rate. (3) Because patterns may contain several spikes or inter-spike intervals, we extend the Pattern Information measure to determine whether the complete pattern carries information distinct from sub-patterns containing only a fraction of these spikes or intervals. (4) The Pattern Information is applied to determine whether a plurality of patterns carry distinct stimulus information from one another. In particular, we demonstrate these concepts using data from cells of the lateral geniculate nucleus (LGN), thereby extending previous analysis demonstrating that distinguishes between bursts of spikes and single spikes providing visual information.     

Fuzzy characterization of spike synchrony in parallel spike trains

We present a framework for characterizing spike (and spike-train) synchrony in parallel neuronal spike trains that is based on the identification of spikes with what we call influence maps: real-valued functions that describe an influence region around the corresponding spike times within which possibly graded (i.e., fuzzy) synchrony with other spikes is defined. We formalize two models of synchrony in this framework: the bin-based model (the almost exclusively applied model in the field) and a novel, alternative model based on a continuous, graded notion of synchrony, aimed at overcoming the drawbacks of the bin-based model. We study the task of identifying frequent (and synchronous) neuronal patterns from parallel spike trains in our framework, formalized as an instance of what we call the fuzzy frequent pattern mining problem (a generalization of standard frequent pattern mining) and briefly evaluate our synchrony models on this task.     

Synergy in a neural code

We show that the information carried by compound events in neural spike trains-patterns of spikes across time or across a population of cells-can be measured, independent of assumptions about what these patterns might represent. By comparing the information carried by a compound pattern with the information carried independently by its parts, we directly measure the synergy among these parts. We illustrate the use of these methods by applying them to experiments on the motion-sensitive neuron H1 of the fly's visual system, where we confirm that two spikes close together in time carry far more than twice the information carried by a single spike. We analyze the sources of this synergy and provide evidence that pairs of spikes close together in time may be especially important patterns in the code of H1.     

Invariant Recognition of Spatio-Temporal Patterns in The Olfactory System Model

This paper presents a model of a network of integrate-and-fire neurons with time delay weights, capable of invariant spatio-temporal pattern recognition. Spatio-temporal patterns are formed by spikes according to the encoding principle that the phase shifts of the spikes encode the input stimulus intensity which corresponds to the concentration of constituent molecules of an odor. We applied the Hopfield's phase shift encoding principle at the output level for spatio-temporal pattern recognition: Firing of an output neuron indicates that corresponding odor is recognized and phase shift of its firing encodes the concentration of the recognized odor. The temporal structure of the model provides the base for the modeling of higher level tasks, where temporal correlation is involved, such as feature binding and segmentation, object recognition, etc.     

Robust sound onset detection using leaky integrate-and-fire neurons with depressing synapses

A biologically inspired technique for detecting onsets in sound is presented. Outputs from a cochlea-like filter are spike coded, in a way similar to the auditory nerve (AN). These AN-like spikes are presented to a leaky integrate-and-fire neuron through a depressing synapse. Onsets are detected with essentially zero latency relative to these AN spikes. Onset detection results for a tone burst, musical sounds and the DARPA/NIST TIMIT speech corpus are presented.