Distortion of neural signals by spike coding

Analog neural signals must be converted into spike trains for transmission over electrically leaky axons. This spike encoding and subsequent decoding leads to distortion. We quantify this distortion by deriving approximate expressions for the mean square error between the inputs and outputs of a spiking link. We use integrate-and-fire and Poisson encoders to convert naturalistic stimuli into spike trains and spike count and inter-spike interval decoders to generate reconstructions of the stimulus. The distortion expressions enable us to compare these spike coding schemes over a large parameter space. We verify that the integrate-and-fire encoder is more effective than the Poisson encoder. The disparity between the two encoders diminishes as the stimulus coefficient of variation (CV) increases, at which point, the variability attributed to the stimulus overwhelms the variability attributed to Poisson statistics. When the stimulus CV is small, the interspike interval decoder is superior, as the distortion resulting from spike count decoding is dominated by a term that is attributed to the discrete nature of the spike count. In this regime, additive noise has a greater impact on the interspike interval decoder than the spike count decoder. When the stimulus CV is large, the average signal excursion is much larger than the quantization step size, and spike count decoding is superior.     

脉冲神经元的信息处理

介绍累积放电脉冲神经元的数学描述;讨论脉冲神经元如何将激励信号转化为脉冲序列;讨论脉冲神经元如何将输入脉冲序列转化为输出脉冲序列。实验结果表明脉冲神经元具有很好的信息表示能力、信号鉴别能力和图像信号重构能力。给出利用脉冲神经网络进行图像信号处理的方法。     

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.     

A computational model as neurodecoder based on synchronous oscillation in the visual cortex

Based on synchronized responses of neuronal populations in the visual cortex to external stimuli, we proposed a computational model consisting primarily of a neuronal phase-locked loop (NPLL) and multiscaled operator. The former reveals the function of synchronous oscillations in the visual cortex. Regardless of which of these patterns of the spike trains may be an average firing-rate code, a spike-timing code, or a rate-time code, the NPLL can decode original visual information from neuronal spike trains modulated with patterns of external stimuli, because a voltage-controlled oscillator (VCO), which is included in the NPLL, can precisely track neuronal spike trains and instantaneous variations, that is, VCO can make a copy of an external stimulus pattern. The latter, however, describes multi-scaled properties of visual information processing, but not merely edge and contour detection. In this study, in which we combined NPLL with a multiscaled operator and maximum likelihood estimation, we proved that the model, as a neurodecoder, implements optimum algorithm decoding visual information from neuronal spike trains at the system level. At the same time, the model also obtains increasingly important supports, which come from a series of experimental results of neurobiology on stimulus-specific neuronal oscillations or synchronized responses of the neuronal population in the visual cortex. In addition, the problem of how to describe visual acuity and multiresolution of vision by wavelet transform is also discussed. The results indicate that the model provides a deeper understanding of the role of synchronized responses in decoding visual information.     

Dynamic analyses of information encoding in neural ensembles

Neural spike train decoding algorithms and techniques to compute Shannon mutual information are important methods for analyzing how neural systems represent biological signals. Decoding algorithms are also one of several strategies being used to design controls for brain-machine interfaces. Developing optimal strategies to design decoding algorithms and compute mutual information are therefore important problems in computational neuroscience. We present a general recursive filter decoding algorithm based on a point process model of individual neuron spiking activity and a linear stochastic state-space model of the biological signal. We derive from the algorithm new instantaneous estimates of the entropy, entropy rate, and the mutual information between the signal and the ensemble spiking activity. We assess the accuracy of the algorithm by computing, along with the decoding error, the true coverage probability of the approximate 0.95 confidence regions for the individual signal estimates. We illustrate the new algorithm by reanalyzing the position and ensemble neural spiking activity of CA1 hippocampal neurons from two rats foraging in an open circular environment. We compare the performance of this algorithm with a linear filter constructed by the widely used reverse correlation method. The median decoding error for Animal 1 (2) during 10 minutes of open foraging was 5.9 (5.5) cm, the median entropy was 6.9 (7.0) bits, the median information was 9.4 (9.4) bits, and the true coverage probability for 0.95 confidence regions was 0.67 (0.75) using 34 (32) neurons. These findings improve significantly on our previous results and suggest an integrated approach to dynamically reading neural codes, measuring their properties, and quantifying the accuracy with which encoded information is extracted.     

Spatiotemporal spike encoding of a continuous external signal

Interspike intervals of spikes emitted from an integrator neuron model of sensory neurons can encode input information represented as a continuous signal from a deterministic system. If a real brain uses spike timing as a means of information processing, other neurons receiving spatiotemporal spikes from such sensory neurons must also be capable of treating information included in deterministic interspike intervals. In this article, we examine functions of neurons modeling cortical neurons receiving spatiotemporal spikes from many sensory neurons. We show that such neuron models can encode stimulus information passed from the sensory model neurons in the form of interspike intervals. Each sensory neuron connected to the cortical neuron contributes equally to the information collection by the cortical neuron. Although the incident spike train to the cortical neuron is a superimposition of spike trains from many sensory neurons, it need not be decomposed into spike trains according to the input neurons. These results are also preserved for generalizations of sensory neurons such as a small amount of leak, noise, inhomogeneity in firing rates, or biases introduced in the phase distributions.     

Analysis and modelling of variability and covariability of population spike trains across multiple time scales

As multi-electrode and imaging technology begin to provide us with simultaneous recordings of large neuronal populations, new methods for modelling such data must also be developed. We present a model of responses to repeated trials of a sensory stimulus based on thresholded Gaussian processes that allows for analysis and modelling of variability and covariability of population spike trains across multiple time scales. The model framework can be used to specify the values of many different variability measures including spike timing precision across trials, coefficient of variation of the interspike interval distribution, and Fano factor of spike counts for individual neurons, as well as signal and noise correlations and correlations of spike counts across multiple neurons. Using both simulated data and data from different stages of the mammalian auditory pathway, we demonstrate the range of possible independent manipulations of different variability measures, and explore how this range depends on the sensory stimulus. The model provides a powerful framework for the study of experimental and surrogate data and for analyzing dependencies between different statistical properties of neuronal populations.     

Spike timing precision and neural error correction: local behavior

The effects of spike timing precision and dynamical behavior on error correction in spiking neurons were investigated. Stationary discharges-phase locked, quasiperiodic, or chaotic-were induced in a simulated neuron by presenting pacemaker presynaptic spike trains across a model of a prototypical inhibitory synapse. Reduced timing precision was modeled by jittering presynaptic spike times. Aftereffects of errors-in this communication, missed presynaptic spikes-were determined by comparing postsynaptic spike times between simulations identical except for the presence or absence of errors. Results show that the effects of an error vary greatly depending on the ongoing dynamical behavior. In the case of phase lockings, a high degree of presynaptic spike timing precision can provide significantly faster error recovery. For nonlocked behaviors, isolated missed spikes can have little or no discernible aftereffects (or even serve to paradoxically reduce uncertainty in postsynaptic spike timing), regardless of presynaptic imprecision. This suggests two possible categories of error correction: high-precision locking with rapid recovery and low-precision nonlocked with error immunity.     

Finding the event structure of neuronal spike trains

Neurons in sensory systems convey information about physical stimuli in their spike trains. In vitro, single neurons respond precisely and reliably to the repeated injection of the same fluctuating current, producing regions of elevated firing rate, termed events. Analysis of these spike trains reveals that multiple distinct spike patterns can be identified as trial-to-trial correlations between spike times (Fellous, Tiesinga, Thomas, & Sejnowski, 2004 ). Finding events in data with realistic spiking statistics is challenging because events belonging to different spike patterns may overlap. We propose a method for finding spiking events that uses contextual information to disambiguate which pattern a trial belongs to. The procedure can be applied to spike trains of the same neuron across multiple trials to detect and separate responses obtained during different brain states. The procedure can also be applied to spike trains from multiple simultaneously recorded neurons in order to identify volleys of near-synchronous activity or to distinguish between excitatory and inhibitory neurons. The procedure was tested using artificial data as well as recordings in vitro in response to fluctuating current waveforms.     

Spike train probability models for stimulus-driven leaky integrate-and-fire neurons

Mathematical models of neurons are widely used to improve understanding of neuronal spiking behavior. These models can produce artificial spike trains that resemble actual spike train data in important ways, but they are not very easy to apply to the analysis of spike train data. Instead, statistical methods based on point process models of spike trains provide a wide range of data-analytical techniques. Two simplified point process models have been introduced in the literature: the time-rescaled renewal process (TRRP) and the multiplicative inhomogeneous Markov interval (m-IMI) model. In this letter we investigate the extent to which the TRRP and m-IMI models are able to fit spike trains produced by stimulus-driven leaky integrate-and-fire (LIF) neurons. With a constant stimulus, the LIF spike train is a renewal process, and the m-IMI and TRRP models will describe accurately the LIF spike train variability. With a time-varying stimulus, the probability of spiking under all three of these models depends on both the experimental clock time relative to the stimulus and the time since the previous spike, but it does so differently for the LIF, m-IMI, and TRRP models. We assessed the distance between the LIF model and each of the two empirical models in the presence of a time-varying stimulus. We found that while lack of fit of a Poisson model to LIF spike train data can be evident even in small samples, the m-IMI and TRRP models tend to fit well, and much larger samples are required before there is statistical evidence of lack of fit of the m-IMI or TRRP models. We also found that when the mean of the stimulus varies across time, the m-IMI model provides a better fit to the LIF data than the TRRP, and when the variance of the stimulus varies across time, the TRRP provides the better fit.     

Model-based decoding, information estimation, and change-point detection techniques for multineuron spike trains

One of the central problems in systems neuroscience is to understand how neural spike trains convey sensory information. Decoding methods, which provide an explicit means for reading out the information contained in neural spike responses, offer a powerful set of tools for studying the neural coding problem. Here we develop several decoding methods based on point-process neural encoding models, or forward models that predict spike responses to stimuli. These models have concave log-likelihood functions, which allow efficient maximum-likelihood model fitting and stimulus decoding. We present several applications of the encoding model framework to the problem of decoding stimulus information from population spike responses: (1) a tractable algorithm for computing the maximum a posteriori (MAP) estimate of the stimulus, the most probable stimulus to have generated an observed single- or multiple-neuron spike train response, given some prior distribution over the stimulus; (2) a gaussian approximation to the posterior stimulus distribution that can be used to quantify the fidelity with which various stimulus features are encoded; (3) an efficient method for estimating the mutual information between the stimulus and the spike trains emitted by a neural population; and (4) a framework for the detection of change-point times (the time at which the stimulus undergoes a change in mean or variance) by marginalizing over the posterior stimulus distribution. We provide several examples illustrating the performance of these estimators with simulated and real neural data.     

Modeling synaptic plasticity in conjuction with the timing of pre- and postsynaptic action potentials

We present a spiking neuron model that allows for an analytic calculation of the correlations between pre- and postsynaptic spikes. The neuron model is a generalization of the integrate-and-fire model and equipped with a probabilistic spike-triggering mechanism. We show that under certain biologically plausible conditions, pre- and postsynaptic spike trains can be described simultaneously as an inhomogeneous Poisson process. Inspired by experimental findings, we develop a model for synaptic long-term plasticity that relies on the relative timing of pre- and post-synaptic action potentials. Being given an input statistics, we compute the stationary synaptic weights that result from the temporal correlations between the pre- and postsynaptic spikes. By means of both analytic calculations and computer simulations, we show that such a mechanism of synaptic plasticity is able to strengthen those input synapses that convey precisely timed spikes at the expense of synapses that deliver spikes with a broad temporal distribution. This may be of vital importance for any kind of information processing based on spiking neurons and temporal coding.     

Detecting and estimating signals in noisy cable structure, I: neuronal noise sources

In recent theoretical approaches addressing the problem of neural coding, tools from statistical estimation and information theory have been applied to quantify the ability of neurons to transmit information through their spike outputs. These techniques, though fairly general, ignore the specific nature of neuronal processing in terms of its known biophysical properties. However, a systematic study of processing at various stages in a biophysically faithful model of a single neuron can identify the role of each stage in information transfer. Toward this end, we carry out a theoretical analysis of the information loss of a synaptic signal propagating along a linear, one-dimensional, weakly active cable due to neuronal noise sources along the way, using both a signal reconstruction and a signal detection paradigm. Here we begin such an analysis by quantitatively characterizing three sources of membrane noise: (1) thermal noise due to the passive membrane resistance, (2) noise due to stochastic openings and closings of voltage-gated membrane channels (NA+ and K+), and (3) noise due to random, background synaptic activity. Using analytical expressions for the power spectral densities of these noise sources, we compare their magnitudes in the case of a patch of membrane from a cortical pyramidal cell and explore their dependence on different biophysical parameters.     

Including long-range dependence in integrate-and-fire models of the high interspike-interval variability of cortical neurons

Many different types of integrate-and-fire models have been designed in order to explain how it is possible for a cortical neuron to integrate over many independent inputs while still producing highly variable spike trains. Within this context, the variability of spike trains has been almost exclusively measured using the coefficient of variation of interspike intervals. However, another important statistical property that has been found in cortical spike trains and is closely associated with their high firing variability is long-range dependence. We investigate the conditions, if any, under which such models produce output spike trains with both interspike-interval variability and long-range dependence similar to those that have previously been measured from actual cortical neurons. We first show analytically that a large class of high-variability integrate-and-fire models is incapable of producing such outputs based on the fact that their output spike trains are always mathematically equivalent to renewal processes. This class of models subsumes a majority of previously published models, including those that use excitation-inhibition balance, correlated inputs, partial reset, or nonlinear leakage to produce outputs with high variability. Next, we study integrate-and-fire models that have (nonPoissonian) renewal point process inputs instead of the Poisson point process inputs used in the preceding class of models. The confluence of our analytical and simulation results implies that the renewal-input model is capable of producing high variability and long-range dependence comparable to that seen in spike trains recorded from cortical neurons, but only if the interspike intervals of the inputs have infinite variance, a physiologically unrealistic condition. Finally, we suggest a new integrate-and-fire model that does not suffer any of the previously mentioned shortcomings. By analyzing simulation results for this model, we show that it is capable of producing output spike trains with interspike-interval variability and long-range dependence that match empirical data from cortical spike trains. This model is similar to the other models in this study, except that its inputs are fractional-gaussian-noise-driven Poisson processes rather than renewal point processes. In addition to this model's success in producing realistic output spike trains, its inputs have long-range dependence similar to that found in most subcortical neurons in sensory pathways, including the inputs to cortex. Analysis of output spike trains from simulations of this model also shows that a tight balance between the amounts of excitation and inhibition at the inputs to cortical neurons is not necessary for high interspike-interval variability at their outputs. Furthermore, in our analysis of this model, we show that the superposition of many fractional-gaussian-noise-driven Poisson processes does not approximate a Poisson process, which challenges the common assumption that the total effect of a large number of inputs on a neuron is well represented by a Poisson process.     

Dynamic analysis of neural encoding by point process adaptive filtering

Neural receptive fields are dynamic in that with experience, neurons change their spiking responses to relevant stimuli. To understand how neural systems adapt their representations of biological information, analyses of receptive field plasticity from experimental measurements are crucial. Adaptive signal processing, the well-established engineering discipline for characterizing the temporal evolution of system parameters, suggests a framework for studying the plasticity of receptive fields. We use the Bayes' rule Chapman-Kolmogorov paradigm with a linear state equation and point process observation models to derive adaptive filters appropriate for estimation from neural spike trains. We derive point process filter analogues of the Kalman filter, recursive least squares, and steepest-descent algorithms and describe the properties of these new filters. We illustrate our algorithms in two simulated data examples. The first is a study of slow and rapid evolution of spatial receptive fields in hippocampal neurons. The second is an adaptive decoding study in which a signal is decoded from ensemble neural spiking activity as the receptive fields of the neurons in the ensemble evolve. Our results provide a paradigm for adaptive estimation for point process observations and suggest a practical approach for constructing filtering algorithms to track neural receptive field dynamics on a millisecond timescale.     

Temporal decoding by phase-locked loops: unique features of circuit-level implementations and their significance for vibrissal information processing

Rhythmic active touch, such as whisking, evokes a periodic reference spike train along which the timing of a novel stimulus, induced, for example, when the whiskers hit an external object, can be interpreted. Previous work supports the hypothesis that the whisking-induced spike train entrains a neural implementation of a phase-locked loop (NPLL) in the vibrissal system. Here we extend this work and explore how the entrained NPLL decodes the delay of the novel, contact-induced stimulus and facilitates object localization. We consider two implementations of NPLLs, which are based on a single neuron or a neural circuit, respectively, and evaluate the resulting temporal decoding capabilities. Depending on the structure of the NPLL, it can lock in either a phase- or co-phase-sensitive mode, which is sensitive to the timing of the input with respect to the beginning of either the current or the next cycle, respectively. The co-phase-sensitive mode is shown to be unique to circuit-based NPLLs. Concentrating on temporal decoding in the vibrissal system of rats, we conclude that both the nature of the information processing task and the response characteristics suggest that the computation is sensitive to the co-phase. Consequently, we suggest that the underlying thalamocortical loop should implement a circuit-based NPLL.     

A unified approach to the study of temporal, correlational, and rate coding

We demonstrate that the information contained in the spike occurrence times of a population of neurons can be broken up into a series of terms, each reflecting something about potential coding mechanisms. This is possible in the coding regime in which few spikes are emitted in the relevant time window. This approach allows us to study the additional information contributed by spike timing beyond that present in the spike counts and to examine the contributions to the whole information of different statistical properties of spike trains, such as firing rates and correlation functions. It thus forms the basis for a new quantitative procedure for analyzing simultaneous multiple neuron recordings and provides theoretical constraints on neural coding strategies. We find a transition between two coding regimes, depending on the size of the relevant observation timescale. For time windows shorter than the timescale of the stimulus-induced response fluctuations, there exists a spike count coding phase, in which the purely temporal information is of third order in time. For time windows much longer than the characteristic timescale, there can be additional timing information of first order, leading to a temporal coding phase in which timing information may affect the instantaneous information rate. In this new framework, we study the relative contributions of the dynamic firing rate and correlation variables to the full temporal information, the interaction of signal and noise correlations in temporal coding, synergy between spikes and between cells, and the effect of refractoriness. We illustrate the utility of the technique by analyzing a few cells from the rat barrel cortex.     

Short-term synaptic plasticity can enhance weak signal detectability in nonrenewal spike trains

We study the encoding of weak signals in spike trains with interspike interval (ISI) correlations and the signals' subsequent detection in sensory neurons. Motivated by the observation of negative ISI correlations in auditory and electrosensory afferents, we assess the theoretical performance limits of an individual detector neuron receiving a weak signal distributed across multiple afferent inputs. We assess the functional role of ISI correlations in the detection process using statistical detection theory and derive two sequential likelihood ratio detector models: one for afferents with renewal statistics; the other for afferents with negatively correlated ISIs. We suggest a mechanism that might enable sensory neurons to implicitly compute conditional probabilities of presynaptic spikes by means of short-term synaptic plasticity. We demonstrate how this mechanism can enhance a postsynaptic neuron's sensitivity to weak signals by exploiting the correlation structure of the input spike trains. Our model not only captures fundamental aspects of early electrosensory signal processing in weakly electric fish, but may also bear relevance to the mammalian auditory system and other sensory modalities.     

面向图像识别的多层脉冲神经网络学习算法综述

相较于第1代和第2代神经网络,第3代神经网络的脉冲神经网络是一种更加接近于生物神经网络的模型,因此更具有生物可解释性和低功耗性。基于脉冲神经元模型,脉冲神经网络可以通过脉冲信号的形式模拟生物信号在神经网络中的传播,通过脉冲神经元的膜电位变化来发放脉冲序列,脉冲序列通过时空联合表达不仅传递了空间信息还传递了时间信息。当前面向模式识别任务的脉冲神经网络模型性能还不及深度学习,其中一个重要原因在于脉冲神经网络的学习方法不成熟,深度学习中神经网络的人工神经元是基于实数形式的输出,这使得其可以使用全局性的反向传播算法对深度神经网络的参数进行训练,脉冲序列是二值性的离散输出,这直接导致对脉冲神经网络的训练存在一定困难,如何对脉冲神经网络进行高效训练是一个具有挑战的研究问题。本文首先总结了脉冲神经网络研究领域中的相关学习算法,然后对其中主要的方法:直接监督学习、无监督学习的算法以及ANN2SNN的转换算法进行分析介绍,并对其中代表性的工作进行对比分析,最后基于对当前主流方法的总结,对未来更高效、更仿生的脉冲神经网络参数学习方法进行展望。     

Analyzing functional connectivity using a network likelihood model of ensemble neural spiking activity

Analyzing the dependencies between spike trains is an important step in understanding how neurons work in concert to represent biological signals. Usually this is done for pairs of neurons at a time using correlation-based techniques. Chornoboy, Schramm, and Karr (1988) proposed maximum likelihood methods for the simultaneous analysis of multiple pair-wise interactions among an ensemble of neurons. One of these methods is an iterative, continuous-time estimation algorithm for a network likelihood model formulated in terms of multiplicative conditional intensity functions. We devised a discrete-time version of this algorithm that includes a new, efficient computational strategy, a principled method to compute starting values, and a principled stopping criterion. In an analysis of simulated neural spike trains from ensembles of interacting neurons, the algorithm recovered the correct connectivity matrices and interaction parameters. In the analysis of spike trains from an ensemble of rat hippocampal place cells, the algorithm identified a connectivity matrix and interaction parameters consistent with the pattern of conjoined firing predicted by the overlap of the neurons' spatial receptive fields. These results suggest that the network likelihood model can be an efficient tool for the analysis of ensemble spiking activity.