FEEDBACK CONTROL,DEVELOPMENT STRATEGY,AND MACROECONOMIC PLANNING

The amplitude of a postsynaptic potential attenuates as it spreads from the synapse to the trigger zone, where spike generation is initiated. Thus, especially in neurons with a large dendritic structure, great variability in synaptically evoked changes of the membrane potential at the trigger zone can be expected. Therefore the randomly distributed magnitudes of jumps caused by input processes were assumed in Stein's neuronal model with reversal potentials. Two distributions, normal and exponential, were applied for this purpose. The normal distribution has no influence on statistical characteristics of the interspike interval (1SI). For the exponential distribution the mean ISI, as well as the coefficient of variation, increases nonlinearly with the parameter of the distribution. The coefficient of variation is limited by the value 1. The estimation of the mean ISI by the method of Smith and Smith (1984) is not applicable for the investigated model.     

Information geometry of interspike intervals in spiking neurons

An information geometrical method is developed for characterizing or classifying neurons in cortical areas, whose spike rates fluctuate in time. Under the assumption that the interspike intervals of a spike sequence of a neuron obey a gamma process with a time-variant spike rate and a fixed shape parameter, we formulate the problem of characterization as a semiparametric statistical estimation, where the spike rate is a nuisance parameter. We derive optimal criteria from the information geometrical viewpoint when certain assumptions are added to the formulation, and we show that some existing measures, such as the coefficient of variation and the local variation, are expressed as estimators of certain functions under the same assumptions.     

Optimal signal estimation in neuronal models

We study optimal estimation of a signal in parametric neuronal models on the basis of interspike interval data. Fisher information is the inverse asymptotic variance of the best estimator. Its dependence on the parameter value indicates accuracy of estimation. Our models assume that the input signal is estimated from neuronal output interspike interval data where the frequency transfer function is sigmoidal. If the coefficient of variation of the interspike interval is constant with respect to the signal, the Fisher information is unimodal, and its maximum for the most estimable signal can be found. We obtain a general result and compare the signal producing maximal Fisher information with the inflection point of the sigmoidal transfer function in several basic neuronal models.     

Interspike interval statistics in the stochastic Hodgkin-Huxley model: coexistence of gamma frequency bursts and highly irregular firing

When the classical Hodgkin-Huxley equations are simulated with Na- and K-channel noise and constant applied current, the distribution of interspike intervals is bimodal: one part is an exponential tail, as often assumed, while the other is a narrow gaussian peak centered at a short interspike interval value. The gaussian arises from bursts of spikes in the gamma-frequency range, the tail from the interburst intervals, giving overall an extraordinarily high coefficient of variation--up to 2.5 for 180,000 Na channels when I approximately 7 microA/cm(2). Since neurons with a bimodal ISI distribution are common, it may be a useful model for any neuron with class 2 firing. The underlying mechanism is due to a subcritical Hopf bifurcation, together with a switching region in phase-space where a fixed point is very close to a system limit cycle. This mechanism may be present in many different classes of neurons and may contribute to widely observed highly irregular neural spiking.     

Firing variability is higher than deduced from the empirical coefficient of variation

A convenient and often used summary measure to quantify the firing variability in neurons is the coefficient of variation (CV), defined as the standard deviation divided by the mean. It is therefore important to find an estimator that gives reliable results from experimental data, that is, the estimator should be unbiased and have low estimation variance. When the CV is evaluated in the standard way (empirical standard deviation of interspike intervals divided by their average), then the estimator is biased, underestimating the true CV, especially if the distribution of the interspike intervals is positively skewed. Moreover, the estimator has a large variance for commonly used distributions. The aim of this letter is to quantify the bias and propose alternative estimation methods. If the distribution is assumed known or can be determined from data, parametric estimators are proposed, which not only remove the bias but also decrease the estimation errors. If no distribution is assumed and the data are very positively skewed, we propose to correct the standard estimator. When defining the corrected estimator, we simply use that it is more stable to work on the log scale for positively skewed distributions. The estimators are evaluated through simulations and applied to experimental data from olfactory receptor neurons in rats.     

Classification of stationary neuronal activity according to its information rate

We propose a measure of the information rate of a single stationary neuronal activity with respect to the state of null information. The measure is based on the Kullback-Leibler distance between two interspike interval distributions. The selected activity is compared with the Poisson model with the same mean firing frequency. We show that the approach is related to the notion of specific information and that the method allows us to judge the relative encoding efficiency. Two classes of neuronal activity models are classified according to their information rate: the renewal process models and the first-order Markov chain models. It has been proven that information can be transmitted changing neither the spike rate nor the coefficient of variation and that the increase in serial correlation does not necessarily increase the information gain. We employ the simple, but powerful, Vasicek's estimator of differential entropy to illustrate an application on the experimental data coming from olfactory sensory neurons of rats.     

Rate limitations of unitary event analysis

Unitary event analysis is a new method for detecting episodes of synchronized neural activity (Riehle, Grün, Diesmann, & Aertsen, 1997). It detects time intervals that contain coincident firing at higher rates than would be expected if the neurons fired as independent inhomogeneous Poisson processes; all coincidences in such intervals are called unitary events (UEs). Changes in the frequency of UEs that are correlated with behavioral states may indicate synchronization of neural firing that mediates or represents the behavioral state. We show that UE analysis is subject to severe limitations due to the underlying discrete statistics of the number of coincident events. These limitations are particularly stringent for low (0-10 spikes/s) firing rates. Under these conditions, the frequency of UEs is a random variable with a large variation relative to its mean. The relative variation decreases with increasing firing rate, and we compute the lowest firing rate, at which the 95% confidence interval around the mean frequency of UEs excludes zero. This random variation in UE frequency makes interpretation of changes in UEs problematic for neurons with low firing rates. As a typical example, when analyzing 150 trials of an experiment using an averaging window 100 ms wide and a 5 ms coincidence window, firing rates should be greater than 7 spikes per second.     

Analysis of spike statistics in neuronal systems with continuous attractors or multiple, discrete attractor States

Attractor networks are likely to underlie working memory and integrator circuits in the brain. It is unknown whether continuous quantities are stored in an analog manner or discretized and stored in a set of discrete attractors. In order to investigate the important issue of how to differentiate the two systems, here we compare the neuronal spiking activity that arises from a continuous (line) attractor with that from a series of discrete attractors. Stochastic fluctuations cause the position of the system along its continuous attractor to vary as a random walk, whereas in a discrete attractor, noise causes spontaneous transitions to occur between discrete states at random intervals. We calculate the statistics of spike trains of neurons firing as a Poisson process with rates that vary according to the underlying attractor network. Since individual neurons fire spikes probabilistically and since the state of the network as a whole drifts randomly, the spike trains of individual neurons follow a doubly stochastic (Poisson) point process. We compare the series of spike trains from the two systems using the autocorrelation function, Fano factor, and interspike interval (ISI) distribution. Although the variation in rate can be dramatically different, especially for short time intervals, surprisingly both the autocorrelation functions and Fano factors are identical, given appropriate scaling of the noise terms. Since the range of firing rates is limited in neurons, we also investigate systems for which the variation in rate is bounded by either rigid limits or because of leak to a single attractor state, such as the Ornstein-Uhlenbeck process. In these cases, the time dependence of the variance in rate can be different between discrete and continuous systems, so that in principle, these processes can be distinguished using second-order spike statistics.     

Mean-driven and fluctuation-driven persistent activity in recurrent networks

Spike trains from cortical neurons show a high degree of irregularity, with coefficients of variation (CV) of their interspike interval (ISI) distribution close to or higher than one. It has been suggested that this irregularity might be a reflection of a particular dynamical state of the local cortical circuit in which excitation and inhibition balance each other. In this "balanced" state, the mean current to the neurons is below threshold, and firing is driven by current fluctuations, resulting in irregular Poisson-like spike trains. Recent data show that the degree of irregularity in neuronal spike trains recorded during the delay period of working memory experiments is the same for both low-activity states of a few Hz and for elevated, persistent activity states of a few tens of Hz. Since the difference between these persistent activity states cannot be due to external factors coming from sensory inputs, this suggests that the underlying network dynamics might support coexisting balanced states at different firing rates. We use mean field techniques to study the possible existence of multiple balanced steady states in recurrent networks of current-based leaky integrate-and-fire (LIF) neurons. To assess the degree of balance of a steady state, we extend existing mean-field theories so that not only the firing rate, but also the coefficient of variation of the interspike interval distribution of the neurons, are determined self-consistently. Depending on the connectivity parameters of the network, we find bistable solutions of different types. If the local recurrent connectivity is mainly excitatory, the two stable steady states differ mainly in the mean current to the neurons. In this case, the mean drive in the elevated persistent activity state is suprathreshold and typically characterized by low spiking irregularity. If the local recurrent excitatory and inhibitory drives are both large and nearly balanced, or even dominated by inhibition, two stable states coexist, both with subthreshold current drive. In this case, the spiking variability in both the resting state and the mnemonic persistent state is large, but the balance condition implies parameter fine-tuning. Since the degree of required fine-tuning increases with network size and, on the other hand, the size of the fluctuations in the afferent current to the cells increases for small networks, overall we find that fluctuation-driven persistent activity in the very simplified type of models we analyze is not a robust phenomenon. Possible implications of considering more realistic models are discussed.     

Discrimination with Spike Times and ISI Distributions

We study the discrimination capability of spike time sequences using the Chernoff distance as a metric. We assume that spike sequences are generated by renewal processes and study how the Chernoff distance depends on the shape of interspike interval (ISI) distribution. First, we consider a lower bound to the Chernoff distance because it has a simple closed form. Then we consider specific models of ISI distributions such as the gamma, inverse gaussian (IG), exponential with refractory period (ER), and that of the leaky integrate-and-fire (LIF) neuron. We found that the discrimination capability of spike times strongly depends on high-order moments of ISI and that it is higher when the spike time sequence has a larger skewness and a smaller kurtosis. High variability in terms of coefficient of variation (CV) does not necessarily mean that the spike times have less discrimination capability. Spike sequences generated by the gamma distribution have the minimum discrimination capability for a given mean and variance of ISI. We used series expansions to calculate the mean and variance of ISIs for LIF neurons as a function of the mean input level and the input noise variance. Spike sequences from an LIF neuron are more capable of discrimination than those of IG and gamma distributions when the stationary voltage level is close to the neuron's threshold value of the neuron.     

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.     

The Ornstein-Uhlenbeck process does not reproduce spiking statistics of neurons in prefrontal cortex

Cortical neurons of behaving animals generate irregular spike sequences. Recently, there has been a heated discussion about the origin of this irregularity. Softky and Koch (1993) pointed out the inability of standard single-neuron models to reproduce the irregularity of the observed spike sequences when the model parameters are chosen within a certain range that they consider to be plausible. Shadlen and Newsome (1994), on the other hand, demonstrated that a standard leaky integrate-and-fire model can reproduce the irregularity if the inhibition is balanced with the excitation. Motivated by this discussion, we attempted to determine whether the Ornstein-Uhlenbeck process, which is naturally derived from the leaky integration assumption, can in fact reproduce higher-order statistics of biological data. For this purpose, we consider actual neuronal spike sequences recorded from the monkey prefrontal cortex to calculate the higher-order statistics of the interspike intervals. Consistency of the data with the model is examined on the basis of the coefficient of variation and the skewness coefficient, which are, respectively, a measure of the spiking irregularity and a measure of the asymmetry of the interval distribution. It is found that the biological data are not consistent with the model if the model time constant assumes a value within a certain range believed to cover all reasonable values. This fact suggests that the leaky integrate-and-fire model with the assumption of uncorrelated inputs is not adequate to account for the spiking in at least some cortical neurons.     

Calculation of interspike intervals for integrate-and-fire neurons with poisson distribution of synaptic inputs

We present a new technique for calculating the interspike intervals of integrate-and-fire neurons. There are two new components to this technique. First, the probability density of the summed potential is calculated by integrating over the distribution of arrival times of the afferent post-synaptic potentials (PSPs), rather than using conventional stochastic differential equation techniques. A general formulation of this technique is given in terms of the probability distribution of the inputs and the time course of the postsynaptic response. The expressions are evaluated in the gaussian approximation, which gives results that become more accurate for large numbers of small-amplitude PSPs. Second, the probability density of output spikes, which are generated when the potential reaches threshold, is given in terms of an integral involving a conditional probability density. This expression is a generalization of the renewal equation, but it holds for both leaky neurons and situations in which there is no time-translational invariance. The conditional probability density of the potential is calculated using the same technique of integrating over the distribution of arrival times of the afferent PSPs. For inputs with a Poisson distribution, the known analytic solutions for both the perfect integrator model and the Stein model (which incorporates membrane potential leakage) in the diffusion limit are obtained. The interspike interval distribution may also be calculated numerically for models that incorporate both membrane potential leakage and a finite rise time of the postsynaptic response. Plots of the relationship between input and output firing rates, as well as the coefficient of variation, are given, and inputs with varying rates and amplitudes, including inhibitory inputs, are analyzed. The results indicate that neurons functioning near their critical threshold, where the inputs are just sufficient to cause firing, display a large variability in their spike timings.     

一种分布式数据流相关性分析的有效方法

在分布式数据流中,数据流之间相关性分析可以揭示被监测对象之间存在的内在联系。提出了一个基于基窗口的相关系数的计算方法,该方法先将计算相关系数的公式变形为由适合基窗口聚集的因子组成,然后用基于基窗口的方法聚集每个因子。基于基窗口的聚集方法是将窗口中的数据项划分成一系列基窗口并分别对基窗口进行计算。当窗口随机滑动后,新窗口中数据项的聚集可以部分地利用上一次窗口聚集的结果。模拟实验表明,与每次对窗口中所有数据进行聚集相比,基于基窗口的方法可以有效地降低数据流相关系数的计算时间。     

采用密度比估计的多窗口变点检测方法

针对基于密度比估计的时间序列变点检测方法受时间窗窗宽限制,识别变点类型单一的问题,利用和发展动态多重过滤算法MFA(multiple filtering algorithm),提出一种多窗口变点检测方法 mDRCPD(multiple window density-ratio change point detection)。将处理后的时间序列按多个时间窗进行适当划分,通过比较相邻时间窗数据的分布差异来识别变点,采用基于密度比估计的相对皮尔逊散度来度量不同时间窗数据分布的差异性;固定窗宽下寻找变点集,并按照MFA方法集成各变点集。模拟实验和实证分析表明,与基于密度比的单窗口变点检测方法相比,mDRCPD方法在多变点时间序列变点检测中绝对误差、召回率、F1得分等指标均有改善。将mDRCPD方法应用到COVID-19的传播进程分析中,通过对传播率的分段建模来刻画疫情的阶段性特点,评估国家政策在降低疫情传播速度上的效果。     

一种新的信号控制干道行程时间实时估计模型

给出了一种新的信号控制干道行程时间实时估计模型. 其建模思路是: 将分析时段分为若干个较短的时间窗, 然后进一步把一个时间窗离散为多个时间间隔. 将干道各交叉口停车线前的车辆是否处于排队定义为干道系统的状态. 在一个时间窗内, 确定每个时间间隔上的干道系统状态, 由此构造出一个无记忆特性的随机过程, 根据离散马尔可夫决策过程理论, 实现了单个时间窗的干道行程时间估计. 在每个时间窗上应用该过程, 实现了干道行程时间的实时估计. 与现有模型相比较, 文中模型的优势体现在: 模型输入是通用的流量和信号配时数据, 模型参数少且容易标定, 模型应用方便、成本低和可移植性强. 最后, 该模型在广州市的某条实际干道上进行了检验.     

Detecting localized repeats in genomic sequences: a new strategy and its application to Bacillus subtilis and Arabidopsis thaliana sequences

A new method for the search of local repeats in long DNA sequences, such as complete genomes, is presented. It detects a large variety of repeats varying in length from one to several hundred bases, which may contain many mutations. By mutations we mean substitutions, insertions or deletions of one or more bases. The method is based on counting occurrences of short words (3-12 bases) in sequence fragments called windows. A score is computed for each window, based on calculating exact word occurrence probabilities for all the words of a given length in the window. The probabilities are defined using a Bernoulli model (independent letters) for the sequence, using the actual letter frequencies from each window. A plot of the probabilities along the sequence for high-scoring windows facilitates the identification of the repeated patterns. We applied the method to the 1.87 Mb sequence of chromosome 4 of Arabidopsis thaliana and to the complete genome of Bacillus subtilis (4.2 Mb). The repeats that we found were classified according to their size, number of occurrences, distance between occurrences, and location with respect to genes. The method proves particularly useful in detecting long, inexact repeats that are local, but not necessarily tandem. The method is implemented as a C program called EXCEP, which is available on request from the authors.     

Robust window operators

It is a common practice in computer vision and image processing to convolve rectangular constant coefficient windows with digital images to perform local smoothing and derivative estimation for edge detection and other purposes. If all data points in each image window belong to the same statistical population, this practice is reasonable and fast. But, as is well known, constant coefficient window operators produce incorrect results if more than one statistical population is present within a window, for example, if a gray-level or gradient discontinuity is present. This paper shows one way to apply the theory of robust statistics to the data smoothing and derivative estimation problem. A robust window operator is demonstrated that preserves gray-level and gradient discontinuities in digital images as it smooths and estimates derivatives.     

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.     

Multiple-window parallel adaptive boundary finding in computer vision

The problem considered in this paper is the estimation of highly variable object boundaries in noisy images. Boundaries may be those of a tank in an IR image, a spinal canal in a CAT scan, a cloud in a visible light image, etc. Or they may be internal to an object such as the boundary between a spherical surface and a cylindrical surface in a manufactured object. The focus of the paper is on parallel multiple-window boundary estimation algorithms. Here the image field is parti-tioned into an array of rectangular windows, and boundary finders are run simultaneously within the windows. The boundary segments found within the windows are then seamed together to obtain meaningful global boundaries. The entire procedure is treated within a maximum likelihood estimation framework that we have developed for boundary finding. Although our multiple-window estimation approach can be used with a number of local boundary finding algorithms, we concen-trate on one which is based on dynamic programming and will produce the true maximum likelihood boundary. Some theoretical considera-tions for boundary model design and boundary-finding runtime are covered. Included is the use of a low computational cost F-test for test-ing whether a window contains a boundary, and an analytical treatment which shows that use of coarse pixels with a chi-square test or an F-test improves the probability of correctly recognizing whether a boundary is present in a window.