Sampling considerations limit the experimental conditions under which information theoretic analyses of neurophysiological data yield reliable results. We develop a procedure for computing the full temporal entropy and information of ensembles of neural spike trains, which performs reliably for limited samples of data. This approach also yields insight upon the role of correlations between spikes in temporal coding mechanisms. The method, when applied to recordings from complex cells of the monkey primary visual cortex, results in lower RMS error information estimates in comparison to a `brute force' approach.
We propose a numerical method to learn Maximum Entropy (MaxEnt) distributions with spatio-temporal constraints from experimental spike trains. This is an extension of two papers [10] and [4] who proposed the estimation of parameters where only spatial constraints were taken into account. The extension we propose allows to properly handle memory effects in spike statistics, for large sized neural networks.
The major problem in information theoretic analysis of neural responses and other biological data is the reliable estimation of entropy--like quantities from small samples. We apply a recently introduced Bayesian entropy estimator to synthetic data inspired by experiments, and to real experimental spike trains. The estimator performs admirably even very deep in the undersampled regime, where other techniques fail. This opens new possibilities for the information theoretic analysis of experiments, and may be of general interest as an example of learning from limited data.
Large networks of sparsely coupled, excitatory and inhibitory cells occur throughout the brain. A striking feature of these networks is that they are chaotic. How does this chaos manifest in the neural code? Specifically, how variable are the spike patterns that such a network produces in response to an input signal? To answer this, we derive a bound for the entropy of multi-cell spike pattern distributions in large recurrent networks of spiking neurons responding to fluctuating inputs. The analysis is based on results from random dynamical systems theory and is complimented by detailed numeri…
Spike generation in neurons produces a temporal point process, whose statistics is governed by intrinsic phenomena and the external incoming inputs to be coded. In particular, spike-evoked adaptation currents support a slow temporal process that conditions spiking probability at the present time according to past activity. In this work, we study the statistics of interspike interval correlations arising in such non-renewal spike trains, for a neuron model that reproduces different spike modes in a small adaptation scenario. We found that correlations are stronger as the neuron fires at a parti…
Understanding how stimuli and synaptic connectivity in uence the statistics of spike patterns in neural networks is a central question in computational neuroscience. Maximum Entropy approach has been successfully used to characterize the statistical response of simultaneously recorded spiking neurons responding to stimuli. But, in spite of good performance in terms of prediction, the fitting parameters do not explain the underlying mechanistic causes of the observed correlations. On the other hand, mathematical models of spiking neurons (neuro-mimetic models) provide a probabilistic mapping be…
The simple system composed of three neural-like noisy elements is considered. Two of them (sensory neurons or sensors) are stimulated by noise and periodic signals with different ratio of frequencies, and the third one (interneuron) receives the output of these two sensors and noise. We propose the analytical approach to analysis of Interspike Intervals (ISI) statistics of the spike train generated by the interneuron. The ISI distributions of the sensory neurons are considered to be known. The frequencies of the input sinusoidal signals are in ratios, which are usual for music. We show that in…
This paper is based on a lecture given in the LACONEU summer school, Valparaiso, January 2012. We introduce Gibbs distribution in a general setting, including non stationary dynamics, and present then three examples of such Gibbs distributions, in the context of neural networks spike train statistics: (i) Maximum entropy model with spatio-temporal constraints; (ii) Generalized Linear Models; (iii) Conductance based Inte- grate and Fire model with chemical synapses and gap junctions.
Spiking activity from populations of neurons display causal interactions and memory effects. Therefore, they are expected to show some degree of irreversibility in time. Motivated by the spike train statistics, in this paper we build a framework to quantify the degree of irreversibility of any maximum entropy distribution. Our approach is based on the transfer matrix technique, which enables us to find an homogeneous irreducible Markov chain that shares the same maximum entropy measure. We provide relevant examples in the context of spike train statistics
Measures of spike train synchrony have proven a valuable tool in both experimental and computational neuroscience. Particularly useful are time-resolved methods such as the ISI- and the SPIKE-distance, which have already been applied in various bivariate and multivariate contexts. Recently, SPIKE-Synchronization was proposed as another time-resolved synchronization measure. It is based on Event-Synchronization and has a very intuitive interpretation. Here, we present a detailed analysis of the mathematical properties of these three synchronization measures. For example, we were able to obtain …
Understanding how the brain functions is one of the biggest challenges of our time. The analysis of experimentally recorded neural firing patterns (spike trains) plays a crucial role in addressing this problem. Here, the PySpike library is introduced, a Python package for spike train analysis providing parameter-free and time-scale independent measures of spike train synchrony. It allows to compute similarity and dissimilarity profiles, averaged values and distance matrices. Although mainly focusing on neuroscience, PySpike can also be applied in other contexts like climate research or social …
We construct a model that predicts the statistical properties of spike trains generated by a sensory neuron. The model describes the combined effects of the neuron's intrinsic properties, the noise in the surrounding, and the external driving stimulus. We show that the spike trains exhibit universal statistical behavior over short times, modulated by a strongly stimulus-dependent behavior over long times. These predictions are confirmed in experiments on H1, a motion-sensitive neuron in the fly visual system.
Estimating the degree of synchrony or reliability between two or more spike trains is a frequent task in both experimental and computational neuroscience. In recent years, many different methods have been proposed that typically compare the timing of spikes on a certain time scale to be fixed beforehand. Here, we propose the ISI-distance, a simple complementary approach that extracts information from the interspike intervals by evaluating the ratio of the instantaneous frequencies. The method is parameter free, time scale independent and easy to visualize as illustrated by an application to re…
Recently, the SPIKE-distance has been proposed as a parameter-free and time-scale independent measure of spike train synchrony. This measure is time-resolved since it relies on instantaneous estimates of spike train dissimilarity. However, its original definition led to spuriously high instantaneous values for event-like firing patterns. Here we present a substantial improvement of this measure which eliminates this shortcoming. The reliability gained allows us to track changes in instantaneous clustering, i.e., time-localized patterns of (dis)similarity among multiple spike trains. Additional…
We investigate the effect of electric synapses (gap junctions) on collective neuronal dynamics and spike statistics in a conductance-based Integrate-and-Fire neural network, driven by a Brownian noise, where conductances depend upon spike history. We compute explicitly the time evolution operator and show that, given the spike-history of the network and the membrane potentials at a given time, the further dynamical evolution can be written in a closed form. We show that spike train statistics is described by a Gibbs distribution whose potential can be approximated with an explicit formula, whe…
We consider a conductance based neural network inspired by the generalized Integrate and Fire model introduced by Rudolph and Destexhe. We show the existence and uniqueness of a unique Gibbs distribution characterizing spike train statistics. The corresponding Gibbs potential is explicitly computed. These results hold in presence of a time-dependent stimulus and apply therefore to non-stationary dynamics.
A wide variety of approaches to estimate the degree of synchrony between two or more spike trains have been proposed. One of the most recent methods is the ISI-distance which extracts information from the interspike intervals (ISIs) by evaluating the ratio of the instantaneous firing rates. In contrast to most previously proposed measures it is parameter free and time-scale independent. However, it is not well suited to track changes in synchrony that are based on spike coincidences. Here we propose the SPIKE-distance, a complementary measure which is sensitive to spike coincidences but still …
Techniques for recording large-scale neuronal spiking activity are developing very fast. This leads to an increasing demand for algorithms capable of analyzing large amounts of experimental spike train data. One of the most crucial and demanding tasks is the identification of similarity patterns with a very high temporal resolution and across different spatial scales. To address this task, in recent years three time-resolved measures of spike train synchrony have been proposed, the ISI-distance, the SPIKE-distance, and event synchronization. The Matlab source codes for calculating and visualiz…
We initiate a mathematical analysis of hidden effects induced by binning spike trains of neurons. Assuming that the original spike train has been generated by a discrete Markov process, we show that binning generates a stochastic process which is not Markov any more, but is instead a Variable Length Markov Chain (VLMC) with unbounded memory. We also show that the law of the binned raster is a Gibbs measure in the DLR (Dobrushin-Lanford-Ruelle) sense coined in mathematical statistical mechanics. This allows the derivation of several important consequences on statistical properties of binned spi…
This paper uses a simple optogenetic model to compare the timing distortion between a randomly-generated target spike sequence and an externally-stimulated neuron spike sequence. Optogenetics is an emerging field of neuroscience where neurons are genetically modified to express light-sensitive receptors that enable external control over when the neurons fire. Given the prominence of neuronal signaling within the brain and throughout the body, optogenetics has significant potential to improve the understanding of the nervous system and to develop treatments for neurological diseases. This paper…
Optogenetics is an emerging field of neuroscience where neurons are genetically modified to express light-sensitive receptors that enable external control over when the neurons fire. Given the prominence of neuronal signaling within the brain and throughout the body, optogenetics has significant potential to improve the understanding of the nervous system and to develop treatments for neurological diseases. This paper uses a simple optogenetic model to compare the timing distortion between a randomly-generated target spike sequence and an externally-stimulated neuron spike sequence. The distor…
Here is proposed a review of the different choices to structure spike trains, using deterministic metrics. Temporal constraints observed in biological or computational spike trains are first taken into account. The relation with existing neural codes (rate coding, rank coding, phase coding, ..) is then discussed. To which extend the "neural code" contained in spike trains is related to a metric appears to be a key point, a generalization of the Victor-Purpura metric family being proposed for temporal constrained causal spike trains
Spiking activity in cortical networks is nonlinear in nature. The linear-nonlinear cascade model, some versions of which are also known as point-process generalized linear model, can efficiently capture the nonlinear dynamics exhibited by such networks. Of particular interest in such models are theoretical predictions of spike train statistics. However, due to the moment-closure problem, approximations are inevitable. We suggest here a series expansion that explains how higher-order moments couple to lower-order ones. Our approach makes predictions in terms of certain integrals, the so-called …
Statistical properties of spike trains measured from a sensory neuron in-vivo are studied experimentally and theoretically. Experiments are performed on an identified neuron in the visual system of the blowfly. It is shown that the spike trains exhibit universal behavior over short time, modulated by a stimulus-dependent envelope over long time. A model of the neuron as a nonlinear oscillator driven by noise and an external stimulus, is suggested to account for these results. The model enables a theoretic distinction of the effects of internal neuronal properties from effects of external stimu…
Background: In neurophysiological data, latency refers to a global shift of spikes from one spike train to the next, either caused by response onset fluctuations or by finite propagation speed. Such systematic shifts in spike timing lead to a spurious decrease in synchrony which needs to be corrected. New Method: We propose a new algorithm of multivariate latency correction suitable for sparse data for which the relevant information is not primarily in the rate but in the timing of each individual spike. The algorithm is designed to correct systematic delays while maintaining all other kinds o…
