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Sue Ann Campbell

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Published work

10 published item(s)

preprint2026arXiv

Approximate Macroscopic Dynamics of Spiking Neural Networks Based on Solutions to the Transport Equation

Firing rate fluctuations in neural populations are observed experimentally over multiple time scales, in single neurons, across trials when elicited by stimuli, and across populations. In this work, we examine how firing rate fluctuations emerge in networks of coupled integrate-and-fire neurons as a function of the initial distribution of voltages in networks with time-varying inputs. We analytically derive an approximation for the evolution of the instantaneous population rate or flux as a function of the initial voltage distribution through a Fokker-Planck system. Unlike earlier mean field approaches based on asynchronous or constant flux steady state solutions to the Fokker-Planck system, the approach considered here is based on the transport solution to the advection equation and assumes that the time-varying inputs are slow, and the neurons are in the excitation-driven regime. The transport mean field system predicts how firing rate fluctuations emerge from a dynamic interaction between time-varying inputs, initial densities, and coupling in populations of neurons.

preprint2022arXiv

Exact mean-field models for spiking neural networks with adaptation

Networks of spiking neurons with adaption have been shown to be able to reproduce a wide range of neural activities, including the emergent population bursting and spike synchrony that underpin brain disorders and normal function. Exact mean-field models derived from spiking neural networks are extremely valuable, as such models can be used to determine how individual neuron and network parameters interact to produce macroscopic network behaviour. In the paper, we derive and analyze a set of exact mean-field equations for the neural network with spike frequency adaptation. Specifically, our model is a network of Izhikevich neurons, where each neuron is modeled by a two dimensional system consisting of a quadratic integrate and fire equation plus an equation which implements spike frequency adaptation. Previous work deriving a mean-field model for this type of network, relied on the assumption of sufficiently slow dynamics of the adaptation variable. However, this approximation did not succeeded in establishing an exact correspondence between the macroscopic description and the realistic neural network, especially when the adaptation time constant was not large. The challenge lies in how to achieve a closed set of mean-field equations with the inclusion of the mean-field expression of the adaptation variable. We address this challenge by using a Lorentzian ansatz combined with the moment closure approach to arrive at the mean-field system in the thermodynamic limit. The resulting macroscopic description is capable of qualitatively and quantitatively describing the collective dynamics of the neural network, including transition between tonic firing and bursting.

preprint2021arXiv

Hysteresis bifurcation and application to delayed Fitzhugh-Nagumo neural systems

Hysteresis dynamics has been described in a vast number of biological experimental studies. Many such studies are phenomenological and a mathematical appreciation has not attracted enough attention. In the paper, we explore the nature of hysteresis and study it from the dynamical system point of view by using the bifurcation and perturbation theories. We firstly make a classification of hysteresis according to the system behaviours transiting between different types of attractors. Then, we focus on a mathematically amenable situation where hysteretic movements between the equilibrium point and the limit cycle are initiated by a subcritical Hopf bifurcation and a saddle-node bifurcation of limit cycles. We present a analytical framework by using the method of multiple scales to obtain the normal form up to the fifth order. Theoretical results are compared with time domain simulations and numerical continuation, showing good agreement. Although we consider the time-delayed FitzHugh-Nagumo neural system in the paper, the generalization should be clear to other systems or parameters. The general framework we present in the paper can be naturally extended to the notion of bursting activity in neuroscience where hysteresis is a dominant mechanism to generate bursting oscillations.

preprint2021arXiv

M-current Induced Bogdanov-Takens Bifurcation and Switching of Neuron Excitability Class

In this work, we consider a general conductance-based neuron model with the inclusion of the acetycholine sensitive, M-current. We study bifurcations in the parameter space consisting of the applied current, $I_{app}$ the maximal conductance of the M-current, $g_M$, and the conductance of the leak current, $g_L$. We give precise conditions for the model that ensure the existence of a Bogdanov-Takens (BT) point and show such a point can occur by varying $I_{app}$ and $g_{M}$. We discuss the case when the BT point becomes a Bogdanov-Takens-Cusp (BTC) point and show that such a point can occur in the three dimensional parameter space. The results of the bifurcation analysis are applied to different neuronal models and are verified and supplemented by numerical bifurcation diagrams generated using the package MATCONT. We conclude that there is a transition in the neuronal excitability type organized by the BT point and the neuron switches from Class-I to Class-II as conductance of the M-current increases.

preprint2020arXiv

A Phase Model with Large Time Delayed Coupling

We consider two identical oscillators with weak, time delayed coupling. We start with a general system of delay differential equations then reduce it to a phase model. With the assumption of large time delay, the resulting phase model has an explicit delay and phase shift in the argument of the phases and connection function, respectively. Using the phase model, we prove that for any type of oscillators and any coupling, the in-phase and anti-phase phase-locked solutions always exist and give conditions for their stability. We show that for small delay these solutions are unique, but with large enough delay multiple solutions of each type with different frequencies may occur. We give conditions for the existence and stability of other types of phase-locked solutions. We discuss the various bifurcations that can occur in the phase model as the time delay is varied. The results of the phase model analysis are applied to Morris-Lecar oscillators with diffusive coupling and compared with numerical studies of the full system of delay differential equations. We also consider the case of small time delay and compare the results with the existing ones in the literature.

preprint2020arXiv

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 investigated 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 considered 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 old age.

preprint2016arXiv

Phase models and clustering in networks of oscillators with delayed coupling

We consider a general model for a network of oscillators with time delayed, circulant coupling. We use the theory of weakly coupled oscillators to reduce the system of delay differential equations to a phase model where the time delay enters as a phase shift. We use the phase model to study the existence and stability of cluster solutions. Cluster solutions are phase locked solutions where the oscillators separate into groups. Oscillators within a group are synchronized while those in different groups are phase-locked. We give model independent existence and stability results for symmetric cluster solutions. We show that the presence of the time delay can lead to the coexistence of multiple stable clustering solutions. We apply our analytical results to a network of Morris Lecar neurons and compare these results with numerical continuation and simulation studies.

preprint2014arXiv

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

Mean-field systems have been recently derived that adequately predict the behaviors of large networks of coupled integrate-and-fire neurons [14]. 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 mean synaptic gating variable of the network. These differential equations are non-smooth, and in particular are piecewise smooth continuous (PWSC). Here, we analyze the smooth and non-smooth 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 2 bifurcations can coalesce into a co-dimension 3 non-smooth bifurcation. As the mean-field system we study is a non-generic piecewise smooth continuous system, we discuss possible regularizations of this system and how the bifurcations which occur are related to non-smooth bifurcations displayed by generic PWSC systems.

preprint2014arXiv

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.

preprint2013arXiv

Mean-Field Models for Heterogeneous Networks of Two-Dimensional Integrate and Fire Models

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.