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Publication# Fitting summary statistics of neural data with a differentiable spiking network simulator

Abstract

Fitting network models to neural activity is an important tool in neuroscience. A popular approach is to model a brain area with a probabilistic recurrent spiking network whose parameters maximize the likelihood of the recorded activity. Although this is widely used, we show that the resulting model does not produce realistic neural activity. To correct for this, we suggest to augment the log-likelihood with terms that measure the dissimilarity between simulated and recorded activity. This dissimilarity is defined via summary statistics commonly used in neuroscience and the optimization is efficient because it relies on back-propagation through the stochastically simulated spike trains. We analyze this method theoretically and show empirically that it generates more realistic activity statistics. We find that it improves upon other fitting algorithms for spiking network models like GLMs (Generalized Linear Models) which do not usually rely on back-propagation. This new fitting algorithm also enables the consideration of hidden neurons which is otherwise notoriously hard, and we show that it can be crucial when trying to infer the network connectivity from spike recordings.

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Neural network

A neural network can refer to a neural circuit of biological neurons (sometimes also called a biological neural network), a network of artificial neurons or nodes in the case of an artificial neur

Brain

A brain is an organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It is located in the head, usually close to the sensory organs for senses such as

Simulation

A simulation is the imitation of the operation of a real-world process or system over time. Simulations require the use of models; the model represents the key characteristics or behaviors of the se

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Claudia Clopath, Wulfram Gerstner, Eleni Vasilaki

Electrophysiological connectivity patterns in cortex often have a few strong connections, which are sometimes bidirectional, among a lot of weak connections. To explain these connectivity patterns, we created a model of spike timing–dependent plasticity (STDP) in which synaptic changes depend on presynaptic spike arrival and the postsynaptic membrane potential, filtered with two different time constants. Our model describes several nonlinear effects that are observed in STDP experiments, as well as the voltage dependence of plasticity. We found that, in a simulated recurrent network of spiking neurons, our plasticity rule led not only to development of localized receptive fields but also to connectivity patterns that reflect the neural code. For temporal coding procedures with spatio-temporal input correlations, strong connections were predominantly unidirectional, whereas they were bidirectional under rate-coded input with spatial correlations only. Thus, variable connectivity patterns in the brain could reflect different coding principles across brain areas; moreover, our simulations suggested that plasticity is fast.

Fast oscillations and in particular gamma-band oscillation (20-80 Hz) are commonly observed during brain function and are at the center of several neural processing theories. In many cases, mathematical analysis of fast oscillations in neural networks has been focused on the transition between irregular and oscillatory firing viewed as an instability of the asynchronous activity. But in fact, brain slice experiments as well as detailed simulations of biological neural networks have produced a large corpus of results concerning the properties of fully developed oscillations that are far from this transition point. We propose here a mathematical approach to deal with nonlinear oscillations in a network of heterogeneous or noisy integrate-and-fire neurons connected by strong inhibition. This approach involves limited mathematical complexity and gives a good sense of the oscillation mechanism, making it an interesting tool to understand fast rhythmic activity in simulated or biological neural networks. A surprising result of our approach is that under some conditions, a change of the strength of inhibition only weakly influences the period of the oscillation. This is in contrast to standard theoretical and experimental models of interneuron network gamma oscillations (ING), where frequency tightly depends on inhibition strength, but it is similar to observations made in some in vitro preparations in the hippocampus and the olfactory bulb and in some detailed network models. This result is explained by the phenomenon of suppression that is known to occur in strongly coupled oscillating inhibitory networks but had not yet been related to the behavior of oscillation frequency.

Moritz Deger, Joao Emanuel Felipe Gerhard

Point process generalized linear models (PP-GLMs) provide an important statistical framework for modeling spiking activity in single-neurons and neuronal networks. Stochastic stability is essential when sampling from these models, as done in computational neuroscience to analyze statistical properties of neuronal dynamics and in neuro-engineering to implement closed-loop applications. Here we show, however, that despite passing common goodness-of-fit tests, PP-GLMs estimated from data are often unstable, leading to divergent firing rates. The inclusion of absolute refractory periods is not a satisfactory solution since the activity then typically settles into unphysiological rates. To address these issues, we derive a framework for determining the existence and stability of fixed points of the expected conditional intensity function (CIF) for general PP-GLMs. Specifically, in nonlinear Hawkes PP-GLMs, the CIF is expressed as a function of the previous spike history and exogenous inputs. We use a mean-field quasi-renewal (QR) approximation that decomposes spike history effects into the contribution of the last spike and an average of the CIF over all spike histories prior to the last spike. Fixed points for stationary rates are derived as self-consistent solutions of integral equations. Bifurcation analysis and the number of fixed points predict that the original models can show stable, divergent, and metastable (fragile) dynamics. For fragile models, fluctuations of the single-neuron dynamics predict expected divergence times after which rates approach unphysiologically high values. This metric can be used to estimate the probability of rates to remain physiological for given time periods, e.g., for simulation purposes. We demonstrate the use of the stability framework using simulated single-neuron examples and neurophysiological recordings. Finally, we show how to adapt PPGLM estimation procedures to guarantee model stability. Overall, our results provide a stability framework for data-driven PP-GLMs and shed new light on the stochastic dynamics of state-of-the-art statistical models of neuronal spiking activity.