No Arabic abstract
Understanding information processing in the brain requires the ability to determine the functional connectivity between the different regions of the brain. We present a method using transfer entropy to extract this flow of information between brain regions from spike-train data commonly obtained in neurological experiments. Transfer entropy is a statistical measure based in information theory that attempts to quantify the information flow from one process to another, and has been applied to find connectivity in simulated spike-train data. Due to statistical error in the estimator, inferring functional connectivity requires a method for determining significance in the transfer entropy values. We discuss the issues with numerical estimation of transfer entropy and resulting challenges in determining significance before presenting the trial-shuffle method as a viable option. The trial-shuffle method, for spike-train data that is split into multiple trials, determines significant transfer entropy values independently for each individual pair of neurons by comparing to a created baseline distribution using a rigorous statistical test. This is in contrast to either globally comparing all neuron transfer entropy values or comparing pairwise values to a single baseline value. In establishing the viability of this method by comparison to several alternative approaches in the literature, we find evidence that preserving the inter-spike-interval timing is important. We then use the trial-shuffle method to investigate information flow within a model network as we vary model parameters. This includes investigating the global flow of information within a connectivity network divided into two well-connected subnetworks, going beyond local transfer of information between pairs of neurons.
Background: Spike trains of multiple neurons can be analyzed following the summed population (SP) or the labeled line (LL) hypothesis. Responses to external stimuli are generated by a neuronal population as a whole or the individual neurons have encoding capacities of their own. The SPIKE-distance estimated either for a single, pooled spike train over a population or for each neuron separately can serve to quantify these responses. New Method: For the SP case we compare three algorithms that search for the most discriminative subpopulation over all stimulus pairs. For the LL case we introduce a new algorithm that combines neurons that individually separate different pairs of stimuli best. Results: The best approach for SP is a brute force search over all possible subpopulations. However, it is only feasible for small populations. For more realistic settings, simulated annealing clearly outperforms gradient algorithms with only a limited increase in computational load. Our novel LL approach can handle very involved coding scenarios despite its computational ease. Comparison with Existing Methods: Spike train distances have been extended to the analysis of neural populations interpolating between SP and LL coding. This includes parametrizing the importance of distinguishing spikes being fired in different neurons. Yet, these approaches only consider the population as a whole. The explicit focus on subpopulations render our algorithms complimentary. Conclusions: The spectrum of encoding possibilities in neural populations is broad. The SP and LL cases are two extremes for which our algorithms provide correct identification results.
We propose a statistical method for modeling the non-Poisson variability of spike trains observed in a wide range of brain regions. Central to our approach is the assumption that the variance and the mean of interspike intervals are related by a power function characterized by two parameters: the scale factor and exponent. It is shown that this single assumption allows the variability of spike trains to have an arbitrary scale and various dependencies on the firing rate in the spike count statistics, as well as in the interval statistics, depending on the two parameters of the power function. We also propose a statistical model for spike trains that exhibits the variance-to-mean power relationship, and based on this a maximum likelihood method is developed for inferring the parameters from rate-modulated spike trains. The proposed method is illustrated on simulated and experimental spike trains.
Rhythmic activity has been associated with a wide range of cognitive processes. Previous studies have shown that spike-timing-dependent plasticity can facilitate the transfer of rhythmic activity downstream the information processing pathway. However, STDP has also been known to generate strong winner-take-all like competitions between subgroups of correlated synaptic inputs. Consequently, one might expect that STDP would induce strong competition between different rhythmicity channels thus preventing the multiplexing of information across different frequency channels. This study explored whether STDP facilitates the multiplexing of information across multiple frequency channels, and if so, under what conditions. We investigated the STDP dynamics in the framework of a model consisting of two competing subpopulations of neurons that synapse in a feedforward manner onto a single postsynaptic neuron. Each sub-population was assumed to oscillate in an independent manner and in a different frequency band. To investigate the STDP dynamics, a mean field Fokker-Planck theory was developed in the limit of the slow learning rate. Surprisingly, our theory predicted limited interactions between the different sub-groups. Our analysis further revealed that the interaction between these channels was mainly mediated by the shared component of the mean activity. Next, we generalized these results beyond the simplistic model using numerical simulations. We found that for a wide range of parameters, the system converged to a solution in which the post-synaptic neuron responded to both rhythms. Nevertheless, all the synaptic weights remained dynamic and did not converge to a fixed point. These findings imply that STDP can support the multiplexing of rhythmic information and demonstrate how functionality can be retained in the face of continuous remodeling of all the synaptic weights.
Entropy is a classical measure to quantify the amount of information or complexity of a system. Various entropy-based measures such as functional and spectral entropies have been proposed in brain network analysis. However, they are less widely used than traditional graph theoretic measures such as global and local efficiencies because either they are not well-defined on a graph or difficult to interpret its biological meaning. In this paper, we propose a new entropy-based graph invariant, called volume entropy. It measures the exponential growth rate of the number of paths in a graph, which is a relevant measure if information flows through the graph forever. We model the information propagation on a graph by the generalized Markov system associated to the weighted edge-transition matrix. We estimate the volume entropy using the stationary equation of the generalized Markov system. A prominent advantage of using the stationary equation is that it assigns certain distribution of weights on the edges of the brain graph, which we call the stationary distribution. The stationary distribution shows the information capacity of edges and the direction of information flow on a brain graph. The simulation results show that the volume entropy distinguishes the underlying graph topology and geometry better than the existing graph measures. In brain imaging data application, the volume entropy of brain graphs was significantly related to healthy normal aging from 20s to 60s. In addition, the stationary distribution of information propagation gives a new insight into the information flow of functional brain graph.
We propose a method, based on persistent homology, to uncover topological properties of a priori unknown covariates of neuron activity. Our input data consist of spike train measurements of a set of neurons of interest, a candidate list of the known stimuli that govern neuron activity, and the corresponding state of the animal throughout the experiment performed. Using a generalized linear model for neuron activity and simple assumptions on the effects of the external stimuli, we infer away any contribution to the observed spike trains by the candidate stimuli. Persistent homology then reveals useful information about any further, unknown, covariates.