No Arabic abstract
A major goal in neuroscience is to understand how populations of neurons code for stimuli or actions. While the number of neurons that can be recorded simultaneously is increasing at a fast pace, in most cases these recordings cannot access a complete population. In particular, it is hard to simultaneously record all the neurons of the same type in a given area. Recent progress have made possible to profile each recorded neuron in a given area thanks to genetic and physiological tools, and to pool together recordings from neurons of the same type across different experimental sessions. However, it is unclear how to infer the activity of a full population of neurons of the same type from these sequential recordings. Neural networks exhibit collective behaviour, e.g. noise correlations and synchronous activity, that are not directly captured by a conditionally-independent model that would just put together the spike trains from sequential recordings. Here we show that we can infer the activity of a full population of retina ganglion cells from sequential recordings, using a novel method based on copula distributions and maximum entropy modeling. From just the spiking response of each ganglion cell to a repeated stimulus, and a few pairwise recordings, we could predict the noise correlations using copulas, and then the full activity of a large population of ganglion cells of the same type using maximum entropy modeling. Remarkably, we could generalize to predict the population responses to different stimuli and even to different experiments. We could therefore use our method to construct a very large population merging cells responses from different experiments. We predicted synchronous activity accurately and showed it grew substantially with the number of neurons. This approach is a promising way to infer population activity from sequential recordings in sensory areas.
Electrical stimulation of neural systems is a key tool for understanding neural dynamics and ultimately for developing clinical treatments. Many applications of electrical stimulation affect large populations of neurons. However, computational models of large networks of spiking neurons are inherently hard to simulate and analyze. We evaluate a reduced mean-field model of excitatory and inhibitory adaptive exponential integrate-and-fire (AdEx) neurons which can be used to efficiently study the effects of electrical stimulation on large neural populations. The rich dynamical properties of this basic cortical model are described in detail and validated using large network simulations. Bifurcation diagrams reflecting the networks state reveal asynchronous up- and down-states, bistable regimes, and oscillatory regions corresponding to fast excitation-inhibition and slow excitation-adaptation feedback loops. The biophysical parameters of the AdEx neuron can be coupled to an electric field with realistic field strengths which then can be propagated up to the population description.We show how on the edge of bifurcation, direct electrical inputs cause network state transitions, such as turning on and off oscillations of the population rate. Oscillatory input can frequency-entrain and phase-lock endogenous oscillations. Relatively weak electric field strengths on the order of 1 V/m are able to produce these effects, indicating that field effects are strongly amplified in the network. The effects of time-varying external stimulation are well-predicted by the mean-field model, further underpinning the utility of low-dimensional neural mass models.
During wakefulness and deep sleep brain states, cortical neural networks show a different behavior, with the second characterized by transients of high network activity. To investigate their impact on neuronal behavior, we apply a pairwise Ising model analysis by inferring the maximum entropy model that reproduces single and pairwise moments of the neurons spiking activity. In this work we first review the inference algorithm introduced in Ferrari,Phys. Rev. E (2016). We then succeed in applying the algorithm to infer the model from a large ensemble of neurons recorded by multi-electrode array in human temporal cortex. We compare the Ising model performance in capturing the statistical properties of the network activity during wakefulness and deep sleep. For the latter, the pairwise model misses relevant transients of high network activity, suggesting that additional constraints are necessary to accurately model the data.
The models in statistical physics such as an Ising model offer a convenient way to characterize stationary activity of neural populations. Such stationary activity of neurons may be expected for recordings from in vitro slices or anesthetized animals. However, modeling activity of cortical circuitries of awake animals has been more challenging because both spike-rates and interactions can change according to sensory stimulation, behavior, or an internal state of the brain. Previous approaches modeling the dynamics of neural interactions suffer from computational cost; therefore, its application was limited to only a dozen neurons. Here by introducing multiple analytic approximation methods to a state-space model of neural population activity, we make it possible to estimate dynamic pairwise interactions of up to 60 neurons. More specifically, we applied the pseudolikelihood approximation to the state-space model, and combined it with the Bethe or TAP mean-field approximation to make the sequential Bayesian estimation of the model parameters possible. The large-scale analysis allows us to investigate dynamics of macroscopic properties of neural circuitries underlying stimulus processing and behavior. We show that the model accurately estimates dynamics of network properties such as sparseness, entropy, and heat capacity by simulated data, and demonstrate utilities of these measures by analyzing activity of monkey V4 neurons as well as a simulated balanced network of spiking neurons.
Neurons in the intact brain receive a continuous and irregular synaptic bombardment from excitatory and inhibitory pre-synaptic neurons, which determines the firing activity of the stimulated neuron. In order to investigate the influence of inhibitory stimulation on the firing time statistics, we consider Leaky Integrate-and-Fire neurons subject to inhibitory instantaneous post-synaptic potentials. In particular, we report exact results for the firing rate, the coefficient of variation and the spike train spectrum for various synaptic weight distributions. Our results are not limited to stimulations of infinitesimal amplitude, but they apply as well to finite amplitude post-synaptic potentials, thus being able to capture the effect of rare and large spikes. The developed methods are able to reproduce also the average firing properties of heterogeneous neuronal populations.
Much of neuroscience aims at reverse engineering the brain, but we only record a small number of neurons at a time. We do not currently know if reverse engineering the brain requires us to simultaneously record most neurons or if multiple recordings from smaller subsets suffice. This is made even more important by the development of novel techniques that allow recording from selected subsets of neurons, e.g. using optical techniques. To get at this question, we analyze a neural network, trained on the MNIST dataset, using only partial recordings and characterize the dependency of the quality of our reverse engineering on the number of simultaneously recorded neurons. We find that reverse engineering of the nonlinear neural network is meaningfully possible if a sufficiently large number of neurons is simultaneously recorded but that this number can be considerably smaller than the number of neurons. Moreover, recording many times from small random subsets of neurons yields surprisingly good performance. Application in neuroscience suggests to approximate the I/O function of an actual neural system, we need to record from a much larger number of neurons. The kind of scaling analysis we perform here can, and arguably should be used to calibrate approaches that can dramatically scale up the size of recorded data sets in neuroscience.