Recurrence-Based Synchronization Analysis of Weakly Coupled Bursting Neurons under External ELF Fields

We investigate the response characteristics of a two-dimensional neuron model exposed to an externally applied extremely low frequency (ELF) sinusoidal electric field and the synchronization of neurons weakly coupled with gap junction. We find, by numerical simulations, that neurons can exhibit different spiking patterns, which are well observed in the structure of the recurrence plot (RP). We further study the synchronization between weakly coupled neurons in chaotic regimes under the influence of a weak ELF electric field. In general, detecting the phases of chaotic spiky signals is not easy by using standard methods. Recurrence analysis provides a reliable tool for defining phases even for noncoherent regimes or spiky signals. Recurrence-based synchronization analysis reveals that, even in the range of weak coupling, phase synchronization of the coupled neurons occurs and, by adding an ELF electric field, this synchronization increases depending on the amplitude of the externally applied ELF electric field. We further suggest a novel measure for RP-based phase synchronization analysis, which better takes into account the probabilities of recurrences.Peer Reviewe

Predicting and identifying signs of criticality near neuronal phase transition

This thesis examines the critical transitions between distinct neural states associated with the transition to neuron spiking and with the induction of anaesthesia. First, mathematical and electronic models of a single spiking neuron are investigated, focusing on stochastic subthreshold dynamics on close approach to spiking and to depolarisation-blocked quiescence (spiking death) transition points. Theoretical analysis of subthreshold neural behaviour then shifts to the anaesthetic-induced phase transition into unconsciousness using a mean-field model for interacting populations of excitatory and inhibitory neurons. The anaesthetic-induced changes are validated experimentally using published electrophysiological data recorded in anaesthetised rats. The criticality hypothesis associated with brain state change is examined using neuronal avalanches for experimentally recorded rat local field potential (LFP) data and mean-field pseudoLFP simulation data. We compare three different implementations of the FitzHugh--Nagumo single spiking neuron model: a mathematical model by H. R. Wilson, an alternative due to Keener and Sneyd, and an op-amp based nonlinear oscillator circuit. Although all three models can produce nonlinear ``spiking" oscillations, our focus is on the altering characteristics of noise-induced fluctuations near spiking onset and death via Hopf bifurcation. We introduce small-amplitude white noise to enable a linearised stochastic analyses using Ornstein--Uhlenbeck theory to predict variance, power spectrum and correlation of voltage fluctuations during close approach to the critical point, identified as the point at which the real part of the dominant eigenvalue becomes zero. We validate the theoretical predictions with numerical simulations and show that the fluctuations exhibit critical slowing down divergences when approaching the critical point: power-law increases in the variance of the fluctuations simultaneous with prolongation of the system response. We expand the study of stochastic behaviour to two spatial dimensions using the Waikato mean-field model operating near phase transition points controlled by the infusion or elimination of anaesthetic inhibition. Specifically, we investigate close approach to the critical point (CP), and to the points of loss of consciousness (LOC) and recovery of consciousness (ROC). We select the equilibrium states using $\lambda$ anaesthetic inhibition and $\Delta V^{\text{rest}}_e$ cortical excitation as control parameters, then analyse the voltage fluctuations evoked by small-amplitude spatiotemporal white noise. We predict the variance and power spectrum of voltage fluctuations near the marginally stable LOC and ROC transition points, then validate via numerical simulation. The results demonstrate a marked increase in voltage fluctuations and spectral power near transition points. This increased susceptibility to low-intensity white noise stimulation provides an early warning of impending phase transition. Effects of anaesthetic agents on cortical activity are reflected in local field potentials (LFPs) by the variation of amplitude and frequency in voltage fluctuations. To explore these changes, we investigate LFPs acquired from published electrophysiological experiments of anaesthetised rats to extract amplitude distribution, variance and time-correlation statistics. The analysis is broadened by applying detrended fluctuation analysis (DFA) to detect long-range dependencies in the time-series, and we compare DFA results with power spectral density (PSD). We find that the DFA exponent increases with anaesthetic concentration, but is always close to 1. The penultimate chapter investigates the evidence of criticality in anaesthetic induced phase-transitions using avalanche analysis. Rat LFP data reveal an avalanche power-law exponent close to $\alpha = 1.5$, but this value depends on both the time-bin width chosen to separate the events and the \textit{z}-score threshold used to detect these events. Power-law behaviour is only evident at lower anaesthetic concentrations; at higher concentrations the avalanche size distribution fails to align with a power-law nature. Criticality behaviour is also indicated in the Waikato mean-field model for anaesthetic-induced phase-transition using avalanches detected from the pseudoLFP time-series, but only at the critical point (CP) and at the secondary phase-transition points of LOC and ROC. In summary, this thesis unveils evidence of characteristic changes near phase transition points using computer-based mathematical modelling and electrophysiological data analysis. We find that noise-driven fluctuations become larger and persist for longer as the critical point is closely approached, with similar properties being seen not only in single-neuron and neural population models, but also in biological LFP signals. These results consistent with an increase of susceptibility to noise perturbations near phase transition point. Identification of neuronal avalanches in rat LFP data for low anaesthetic concentrations provides further support for the criticality hypothesis

Complex oscillations with multiple timescales - Application to neuronal dynamics

The results gathered in this thesis deal with multiple time scale dynamical systems near non-hyperbolic points, giving rise to canard-type solutions, in systems of dimension 2, 3 and 4. Bifurcation theory and numerical continuation methods adapted for such systems are used to analyse canard cycles as well as canard-induced complex oscillations in three-dimensional systems. Two families of such complex oscillations are considered: mixed-mode oscillations (MMOs) in systems with two slow variables, and bursting oscillations in systems with two fast variables. In the last chapter, we present recent results on systems with two slow and two fast variables, where both MMO-type dynamics and bursting-type dynamics can arise and where complex oscillations are also organised by canard solutions. The main application area that we consider here is that of neuroscience, more precisely low-dimensional point models of neurons displaying both sub-threshold and spiking behaviour. We focus on analysing how canard objects allow to control the oscillatory patterns observed in these neuron models, in particular the crossings of excitability thresholds

Pattern formation in electrically coupled pacemaker cells : a thesis presented in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Mathematics at Massey University, Manawatū, New Zealand

Figures are re-used with permission.In this thesis we study electrical activity in smooth muscle cells in the absence of external stimulation. The main goal is to analyse a reaction-diffusion system that models the dynamical behaviour where adjacent cells are coupled through passive electrical coupling. We first analyse the dynamics of an isolated muscle cell for which the model consists of three first-order ordinary differential equations. The cell is either excitable, nonexcitable, or oscillatory depending on the model parameters. To understand this we reduce the model to two equations, nondimensionalise, then perform a detailed numerical bifurcation analysis of the nondimensionalised model. One parameter bifurcation diagrams reveal that even though there is no external stimulus the cell can exhibit two fundamentally distinct types of excitability. By computing two-parameter bifurcation diagrams we are able to explain how the cell transitions between the two types of excitability as parameters are varied. We then study the full reaction-diffusion system first through numerical integration. We show that the system is capable of exhibiting a wide variety of spatiotemporal behaviours such as travelling pulses, travelling fronts, and spatiotemporal chaos. Through a linear stability analysis we are able to show that the spatiotemporal patterns are not due to diffusion-driven instability as is often the case for reaction-diffusion systems. It is as a consequence of the nonlinear dynamics of the reaction terms and coupling effect of diffusion. The precise mechanism is not yet well understood, this will be subject of future work. We then examine travelling wave solutions in detail. In particular we show how they relate to homoclinic and heteroclinic solutions in travelling wave coordinates. Finally we review spectral stability analysis for travelling waves and compute the essential spectrum of travelling waves in our system

Astrocytic Ion Dynamics: Implications for Potassium Buffering and Liquid Flow

We review modeling of astrocyte ion dynamics with a specific focus on the implications of so-called spatial potassium buffering, where excess potassium in the extracellular space (ECS) is transported away to prevent pathological neural spiking. The recently introduced Kirchoff-Nernst-Planck (KNP) scheme for modeling ion dynamics in astrocytes (and brain tissue in general) is outlined and used to study such spatial buffering. We next describe how the ion dynamics of astrocytes may regulate microscopic liquid flow by osmotic effects and how such microscopic flow can be linked to whole-brain macroscopic flow. We thus include the key elements in a putative multiscale theory with astrocytes linking neural activity on a microscopic scale to macroscopic fluid flow.Comment: 27 pages, 7 figure