One-Dimensional Population Density Approaches to Recurrently Coupled Networks of Neurons with Noise

Mean-field systems have been previously derived for networks of coupled, two-dimensional, integrate-and-fire neurons such as the Izhikevich, adapting exponential (AdEx) and quartic integrate and fire (QIF), among others. Unfortunately, the mean-field systems have a degree of frequency error and the networks analyzed often do not include noise when there is adaptation. Here, we derive a one-dimensional partial differential equation (PDE) approximation for the marginal voltage density under a first order moment closure for coupled networks of integrate-and-fire neurons with white noise inputs. The PDE has substantially less frequency error than the mean-field system, and provides a great deal more information, at the cost of analytical tractability. The convergence properties of the mean-field system in the low noise limit are elucidated. A novel method for the analysis of the stability of the asynchronous tonic firing solution is also presented and implemented. Unlike previous attempts at stability analysis with these network types, information about the marginal densities of the adaptation variables is used. This method can in principle be applied to other systems with nonlinear partial differential equations.Comment: 26 Pages, 6 Figure

Bifurcation Analysis of Large Networks of Neurons

The human brain contains on the order of a hundred billion neurons, each with several thousand synaptic connections. Computational neuroscience has successfully modeled both the individual neurons as various types of oscillators, in addition to the synaptic coupling between the neurons. However, employing the individual neuronal models as a large coupled network on the scale of the human brain would require massive computational and financial resources, and yet is the current undertaking of several research groups. Even if one were to successfully model such a complicated system of coupled differential equations, aside from brute force numerical simulations, little insight may be gained into how the human brain solves problems or performs tasks. Here, we introduce a tool that reduces large networks of coupled neurons to a much smaller set of differential equations that governs key statistics for the network as a whole, as opposed to tracking the individual dynamics of neurons and their connections. This approach is typically referred to as a mean-field system. As the mean-field system is derived from the original network of neurons, it is predictive for the behavior of the network as a whole and the parameters or distributions of parameters that appear in the mean-field system are identical to those of the original network. As such, bifurcation analysis is predictive for the behavior of the original network and predicts where in the parameter space the network transitions from one behavior to another. Additionally, here we show how networks of neurons can be constructed with a mean-field or macroscopic behavior that is prescribed. This occurs through an analytic extension of the Neural Engineering Framework (NEF). This can be thought of as an inverse mean-field approach, where the networks are constructed to obey prescribed dynamics as opposed to deriving the macroscopic dynamics from an underlying network. Thus, the work done here analyzes neuronal networks through both top-down and bottom-up approaches

Mean-field models for heterogeneous networks of two-dimensional integrate and fire neurons

We analytically derive mean-field models for all-to-all coupled networks of heterogeneous, adapting, two-dimensional integrate and fire neurons. The class of models we consider includes the Izhikevich, adaptive exponential and quartic integrate and fire models. The heterogeneity in the parameters leads to different moment closure assumptions that can be made in the derivation of the mean-field model from the population density equation for the large network. Three different moment closure assumptions lead to three different mean-field systems. These systems can be used for distinct purposes such as bifurcation analysis of the large networks, prediction of steady state firing rate distributions, parameter estimation for actual neurons and faster exploration of the parameter space. We use the mean-field systems to analyze adaptation induced bursting under realistic sources of heterogeneity in multiple parameters. Our analysis demonstrates that the presence of heterogeneity causes the Hopf bifurcation associated with the emergence of bursting to change from sub-critical to super-critical. This is confirmed with numerical simulations of the full network for biologically reasonable parameter values. This change decreases the plausibility of adaptation being the cause of bursting in hippocampal area CA3, an area with a sizable population of heavily coupled, strongly adapting neurons.Natural Sciences and Engineering Research Council of CanadaOntario Graduate Scholarship progra

Nonsmooth Bifurcations of Mean Field Systems of Two-Dimensional Integrate and Fire Neurons

First Published in SIAM Journal on Applied Dynamical Systems in 15[1], 2016 published by the Society for Industrial and Applied Mathematics (SIAM) Copyright © by SIAM. Unauthorized reproduction of this article is prohibited.Mean field systems have recently been derived that adequately predict the behaviors of large networks of coupled integrate-and-fire neurons [W. Nicola and S.A. Campbell, J. Comput. Neurosci., 35 (2013), pp. 87-108]. The mean field system for a network of neurons with spike frequency adaptation is typically a pair of differential equations for the mean adaptation and synaptic gating variable of the network. These differential equations are nonsmooth, and, in particular, are piecewise smooth continuous (PWSC). Here, we analyze the smooth and nonsmooth bifurcation structure of these equations and show that the system is organized around a pair of co-dimension-two bifurcations that involve, respectively, the collision between a Hopf equilibrium point and a switching manifold, and a saddle-node equilibrium point and a switching manifold. These two co-dimension-two bifurcations can coalesce into a co-dimension-three nonsmooth bifurcation. As the mean field system we study is a nongeneric piecewise smooth continuous system, we discuss possible regularizations of this system and how the bifurcations which occur are related to nonsmooth bifurcations displayed by generic PWSC systems

Normalized Connectomes Show Increased Synchronizability with Age through Their Second Largest Eigenvalue

The synchronization of different brain regions is widely observed under both normal and pathological conditions, such as epilepsy. However, the relationship between the dynamics of these brain regions, the connectivity between them, and the ability to synchronize remains an open question. We investigate the problem of inter-region synchronization in networks of Wilson--Cowan/neural field equations with homeostatic plasticity, each of which acts as a model for an isolated brain region. We consider arbitrary connection profiles with only one constraint: the rows of the connection matrices are all identically normalized. We found that these systems often synchronize to the solution obtained from a single, self-coupled neural region. We analyze the stability of this solution through a straightforward modification of the master stability function (MSF) approach and found that synchronized solutions lose stability for connectivity matrices when the second largest positive eigenvalue is sufficiently large for values of the global coupling parameter that are not too large. This result was numerically confirmed for ring systems and lattices and was also robust to small amounts of heterogeneity in the homeostatic set points in each node. Finally, we tested this result on connectomes obtained from 196 subjects over a broad age range (4--85 years) from the Human Connectome Project. We found that the second largest eigenvalue tended to decrease with age, indicating an increase in synchronizability that may be related to the increased prevalence of epilepsy with advancing age.Natural Sciences and Engineering Research Council, Discovery grant

The Impact of Small Time Delays on the Onset of Oscillations and Synchrony in Brain Networks

The human brain constitutes one of the most advanced networks produced by nature, consisting of billions of neurons communicating with each other. However, this communication is not in real-time, with different communication or time-delays occurring between neurons in different brain areas. Here, we investigate the impacts of these delays by modeling large interacting neural circuits as neural-field systems which model the bulk activity of populations of neurons. By using a Master Stability Function analysis combined with numerical simulations, we find that delays (1) may actually stabilize brain dynamics by temporarily preventing the onset to oscillatory and pathologically synchronized dynamics and (2) may enhance or diminish synchronization depending on the underlying eigenvalue spectrum of the connectivity matrix. Real eigenvalues with large magnitudes result in increased synchronizability while complex eigenvalues with large magnitudes and positive real parts yield a decrease in synchronizability in the delay vs. instantaneously coupled case. This result applies to networks with fixed, constant delays, and was robust to networks with heterogeneous delays. In the case of real brain networks, where the eigenvalues are predominantly real, owing to the nearly symmetric nature of these weight matrices, biologically plausible, small delays, are likely to increase synchronization, rather than decreasing it.Natural Sciences and Engineering Research Council of Canada