Response variability in balanced cortical networks

We study the spike statistics of neurons in a network with dynamically balanced excitation and inhibition. Our model, intended to represent a generic cortical column, comprises randomly connected excitatory and inhibitory leaky integrate-and-fire neurons, driven by excitatory input from an external population. The high connectivity permits a mean-field description in which synaptic currents can be treated as Gaussian noise, the mean and autocorrelation function of which are calculated self-consistently from the firing statistics of single model neurons. Within this description, we find that the irregularity of spike trains is controlled mainly by the strength of the synapses relative to the difference between the firing threshold and the post-firing reset level of the membrane potential. For moderately strong synapses we find spike statistics very similar to those observed in primary visual cortex.Comment: 22 pages, 7 figures, submitted to Neural Computatio

Tracking dynamic interactions between structural and functional connectivity : a TMS/EEG-dMRI study

Transcranial magnetic stimulation (TMS) in combination with neuroimaging techniques allows to measure the effects of a direct perturbation of the brain. When coupled with high-density electroencephalography (TMS/hd-EEG), TMS pulses revealed electrophysiological signatures of different cortical modules in health and disease. However, the neural underpinnings of these signatures remain unclear. Here, by applying multimodal analyses of cortical response to TMS recordings and diffusion magnetic resonance imaging (dMRI) tractography, we investigated the relationship between functional and structural features of different cortical modules in a cohort of awake healthy volunteers. For each subject, we computed directed functional connectivity interactions between cortical areas from the source-reconstructed TMS/hd-EEG recordings and correlated them with the correspondent structural connectivity matrix extracted from dMRI tractography, in three different frequency bands (alpha, beta, gamma) and two sites of stimulation (left precuneus and left premotor). Each stimulated area appeared to mainly respond to TMS by being functionally elicited in specific frequency bands, that is, beta for precuneus and gamma for premotor. We also observed a temporary decrease in the whole-brain correlation between directed functional connectivity and structural connectivity after TMS in all frequency bands. Notably, when focusing on the stimulated areas only, we found that the structure-function correlation significantly increases over time in the premotor area controlateral to TMS. Our study points out the importance of taking into account the major role played by different cortical oscillations when investigating the mechanisms for integration and segregation of information in the human brain

Ising Models for Inferring Network Structure From Spike Data

Now that spike trains from many neurons can be recorded simultaneously, there is a need for methods to decode these data to learn about the networks that these neurons are part of. One approach to this problem is to adjust the parameters of a simple model network to make its spike trains resemble the data as much as possible. The connections in the model network can then give us an idea of how the real neurons that generated the data are connected and how they influence each other. In this chapter we describe how to do this for the simplest kind of model: an Ising network. We derive algorithms for finding the best model connection strengths for fitting a given data set, as well as faster approximate algorithms based on mean field theory. We test the performance of these algorithms on data from model networks and experiments.Comment: To appear in "Principles of Neural Coding", edited by Stefano Panzeri and Rodrigo Quian Quirog

Locking of correlated neural activity to ongoing oscillations

Population-wide oscillations are ubiquitously observed in mesoscopic signals of cortical activity. In these network states a global oscillatory cycle modulates the propensity of neurons to fire. Synchronous activation of neurons has been hypothesized to be a separate channel of signal processing information in the brain. A salient question is therefore if and how oscillations interact with spike synchrony and in how far these channels can be considered separate. Experiments indeed showed that correlated spiking co-modulates with the static firing rate and is also tightly locked to the phase of beta-oscillations. While the dependence of correlations on the mean rate is well understood in feed-forward networks, it remains unclear why and by which mechanisms correlations tightly lock to an oscillatory cycle. We here demonstrate that such correlated activation of pairs of neurons is qualitatively explained by periodically-driven random networks. We identify the mechanisms by which covariances depend on a driving periodic stimulus. Mean-field theory combined with linear response theory yields closed-form expressions for the cyclostationary mean activities and pairwise zero-time-lag covariances of binary recurrent random networks. Two distinct mechanisms cause time-dependent covariances: the modulation of the susceptibility of single neurons (via the external input and network feedback) and the time-varying variances of single unit activities. For some parameters, the effectively inhibitory recurrent feedback leads to resonant covariances even if mean activities show non-resonant behavior. Our analytical results open the question of time-modulated synchronous activity to a quantitative analysis.Comment: 57 pages, 12 figures, published versio

VIOLA - A multi-purpose and web-based visualization tool for neuronal-network simulation output

Neuronal network models and corresponding computer simulations are invaluable tools to aid the interpretation of the relationship between neuron properties, connectivity and measured activity in cortical tissue. Spatiotemporal patterns of activity propagating across the cortical surface as observed experimentally can for example be described by neuronal network models with layered geometry and distance-dependent connectivity. The interpretation of the resulting stream of multi-modal and multi-dimensional simulation data calls for integrating interactive visualization steps into existing simulation-analysis workflows. Here, we present a set of interactive visualization concepts called views for the visual analysis of activity data in topological network models, and a corresponding reference implementation VIOLA (VIsualization Of Layer Activity). The software is a lightweight, open-source, web-based and platform-independent application combining and adapting modern interactive visualization paradigms, such as coordinated multiple views, for massively parallel neurophysiological data. For a use-case demonstration we consider spiking activity data of a two-population, layered point-neuron network model subject to a spatially confined excitation originating from an external population. With the multiple coordinated views, an explorative and qualitative assessment of the spatiotemporal features of neuronal activity can be performed upfront of a detailed quantitative data analysis of specific aspects of the data. Furthermore, ongoing efforts including the European Human Brain Project aim at providing online user portals for integrated model development, simulation, analysis and provenance tracking, wherein interactive visual analysis tools are one component. Browser-compatible, web-technology based solutions are therefore required. Within this scope, with VIOLA we provide a first prototype.Comment: 38 pages, 10 figures, 3 table

The Effect of Nonstationarity on Models Inferred from Neural Data

Neurons subject to a common non-stationary input may exhibit a correlated firing behavior. Correlations in the statistics of neural spike trains also arise as the effect of interaction between neurons. Here we show that these two situations can be distinguished, with machine learning techniques, provided the data are rich enough. In order to do this, we study the problem of inferring a kinetic Ising model, stationary or nonstationary, from the available data. We apply the inference procedure to two data sets: one from salamander retinal ganglion cells and the other from a realistic computational cortical network model. We show that many aspects of the concerted activity of the salamander retinal neurons can be traced simply to the external input. A model of non-interacting neurons subject to a non-stationary external field outperforms a model with stationary input with couplings between neurons, even accounting for the differences in the number of model parameters. When couplings are added to the non-stationary model, for the retinal data, little is gained: the inferred couplings are generally not significant. Likewise, the distribution of the sizes of sets of neurons that spike simultaneously and the frequency of spike patterns as function of their rank (Zipf plots) are well-explained by an independent-neuron model with time-dependent external input, and adding connections to such a model does not offer significant improvement. For the cortical model data, robust couplings, well correlated with the real connections, can be inferred using the non-stationary model. Adding connections to this model slightly improves the agreement with the data for the probability of synchronous spikes but hardly affects the Zipf plot.Comment: version in press in J Stat Mec

State Dependence of Stimulus-Induced Variability Tuning in Macaque MT

Behavioral states marked by varying levels of arousal and attention modulate some properties of cortical responses (e.g. average firing rates or pairwise correlations), yet it is not fully understood what drives these response changes and how they might affect downstream stimulus decoding. Here we show that changes in state modulate the tuning of response variance-to-mean ratios (Fano factors) in a fashion that is neither predicted by a Poisson spiking model nor changes in the mean firing rate, with a substantial effect on stimulus discriminability. We recorded motion-sensitive neurons in middle temporal cortex (MT) in two states: alert fixation and light, opioid anesthesia. Anesthesia tended to lower average spike counts, without decreasing trial-to-trial variability compared to the alert state. Under anesthesia, within-trial fluctuations in excitability were correlated over longer time scales compared to the alert state, creating supra-Poisson Fano factors. In contrast, alert-state MT neurons have higher mean firing rates and largely sub-Poisson variability that is stimulus-dependent and cannot be explained by firing rate differences alone. The absence of such stimulus-induced variability tuning in the anesthetized state suggests different sources of variability between states. A simple model explains state-dependent shifts in the distribution of observed Fano factors via a suppression in the variance of gain fluctuations in the alert state. A population model with stimulus-induced variability tuning and behaviorally constrained information-limiting correlations explores the potential enhancement in stimulus discriminability by the cortical population in the alert state.Comment: 36 pages, 18 figure

Self-organization of network dynamics into local quantized states

Self-organization and pattern formation in network-organized systems emerges from the collective activation and interaction of many interconnected units. A striking feature of these non-equilibrium structures is that they are often localized and robust: only a small subset of the nodes, or cell assembly, is activated. Understanding the role of cell assemblies as basic functional units in neural networks and socio-technical systems emerges as a fundamental challenge in network theory. A key open question is how these elementary building blocks emerge, and how they operate, linking structure and function in complex networks. Here we show that a network analogue of the Swift-Hohenberg continuum model---a minimal-ingredients model of nodal activation and interaction within a complex network---is able to produce a complex suite of localized patterns. Hence, the spontaneous formation of robust operational cell assemblies in complex networks can be explained as the result of self-organization, even in the absence of synaptic reinforcements. Our results show that these self-organized, local structures can provide robust functional units to understand natural and socio-technical network-organized processes.Comment: 11 pages, 4 figure