A universal description of stochastic oscillators.

Many systems in physics, chemistry, and biology exhibit oscillations with a pronounced random component. Such stochastic oscillations can emerge via different mechanisms, for example, linear dynamics of a stable focus with fluctuations, limit-cycle systems perturbed by noise, or excitable systems in which random inputs lead to a train of pulses. Despite their diverse origins, the phenomenology of random oscillations can be strikingly similar. Here, we introduce a nonlinear transformation of stochastic oscillators to a complex-valued function [Formula: see text](x) that greatly simplifies and unifies the mathematical description of the oscillator's spontaneous activity, its response to an external time-dependent perturbation, and the correlation statistics of different oscillators that are weakly coupled. The function [Formula: see text] (x) is the eigenfunction of the Kolmogorov backward operator with the least negative (but nonvanishing) eigenvalue λ1 = µ1 + iω1. The resulting power spectrum of the complex-valued function is exactly given by a Lorentz spectrum with peak frequency ω1 and half-width µ1; its susceptibility with respect to a weak external forcing is given by a simple one-pole filter, centered around ω1; and the cross-spectrum between two coupled oscillators can be easily expressed by a combination of the spontaneous power spectra of the uncoupled systems and their susceptibilities. Our approach makes qualitatively different stochastic oscillators comparable, provides simple characteristics for the coherence of the random oscillation, and gives a framework for the description of weakly coupled oscillators.

A framework for macroscopic phase-resetting curves for generalised spiking neural networks.

Brain rhythms emerge from synchronization among interconnected spiking neurons. Key properties of such rhythms can be gleaned from the phase-resetting curve (PRC). Inferring the PRC and developing a systematic phase reduction theory for large-scale brain rhythms remains an outstanding challenge. Here we present a theoretical framework and methodology to compute the PRC of generic spiking networks with emergent collective oscillations. We adopt a renewal approach where neurons are described by the time since their last action potential, a description that can reproduce the dynamical feature of many cell types. For a sufficiently large number of neurons, the network dynamics are well captured by a continuity equation known as the refractory density equation. We develop an adjoint method for this equation giving a semi-analytical expression of the infinitesimal PRC. We confirm the validity of our framework for specific examples of neural networks. Our theoretical framework can link key biological properties at the individual neuron scale and the macroscopic oscillatory network properties. Beyond spiking networks, the approach is applicable to a broad class of systems that can be described by renewal processes.

Quantitative comparison of the mean-return-time phase and the stochastic asymptotic phase for noisy oscillators.

Seminal work by A. Winfree and J. Guckenheimer showed that a deterministic phase variable can be defined either in terms of Poincaré sections or in terms of the asymptotic (long-time) behaviour of trajectories approaching a stable limit cycle. However, this equivalence between the deterministic notions of phase is broken in the presence of noise. Different notions of phase reduction for a stochastic oscillator can be defined either in terms of mean-return-time sections or as the argument of the slowest decaying complex eigenfunction of the Kolmogorov backwards operator. Although both notions of phase enjoy a solid theoretical foundation, their relationship remains unexplored. Here, we quantitatively compare both notions of stochastic phase. We derive an expression relating both notions of phase and use it to discuss differences (and similarities) between both definitions of stochastic phase for (i) a spiral sink motivated by stochastic models for electroencephalograms, (ii) noisy limit-cycle systems-neuroscience models, and (iii) a stochastic heteroclinic oscillator inspired by a simple motor-control system.

Perturbations both trigger and delay seizures due to generic properties of slow-fast relaxation oscillators.

The mechanisms underlying the emergence of seizures are one of the most important unresolved issues in epilepsy research. In this paper, we study how perturbations, exogenous or endogenous, may promote or delay seizure emergence. To this aim, due to the increasingly adopted view of epileptic dynamics in terms of slow-fast systems, we perform a theoretical analysis of the phase response of a generic relaxation oscillator. As relaxation oscillators are effectively bistable systems at the fast time scale, it is intuitive that perturbations of the non-seizing state with a suitable direction and amplitude may cause an immediate transition to seizure. By contrast, and perhaps less intuitively, smaller amplitude perturbations have been found to delay the spontaneous seizure initiation. By studying the isochrons of relaxation oscillators, we show that this is a generic phenomenon, with the size of such delay depending on the slow flow component. Therefore, depending on perturbation amplitudes, frequency and timing, a train of perturbations causes an occurrence increase, decrease or complete suppression of seizures. This dependence lends itself to analysis and mechanistic understanding through methods outlined in this paper. We illustrate this methodology by computing the isochrons, phase response curves and the response to perturbations in several epileptic models possessing different slow vector fields. While our theoretical results are applicable to any planar relaxation oscillator, in the motivating context of epilepsy they elucidate mechanisms of triggering and abating seizures, thus suggesting stimulation strategies with effects ranging from mere delaying to full suppression of seizures.

Isostables for Stochastic Oscillators.

Thomas and Lindner [P. J. Thomas and B. Lindner, Phys. Rev. Lett. 113, 254101 (2014).PRLTAO0031-900710.1103/PhysRevLett.113.254101], defined an asymptotic phase for stochastic oscillators as the angle in the complex plane made by the eigenfunction, having a complex eigenvalue with a least negative real part, of the backward Kolmogorov (or stochastic Koopman) operator. We complete the phase-amplitude description of noisy oscillators by defining the stochastic isostable coordinate as the eigenfunction with the least negative nontrivial real eigenvalue. Our results suggest a framework for stochastic limit cycle dynamics that encompasses noise-induced oscillations.

Global phase-amplitude description of oscillatory dynamics via the parameterization method.

In this paper, we use the parameterization method to provide a complete description of the dynamics of an n-dimensional oscillator beyond the classical phase reduction. The parameterization method allows us, via efficient algorithms, to obtain a parameterization of the attracting invariant manifold of the limit cycle in terms of the phase-amplitude variables. The method has several advantages. It provides analytically a Fourier-Taylor expansion of the parameterization up to any order, as well as a simplification of the dynamics that allows for a numerical globalization of the manifolds. Thus, one can obtain the local and global isochrons and isostables, including the slow attracting manifold, up to high accuracy, which offer a geometrical portrait of the oscillatory dynamics. Furthermore, it provides straightforwardly the infinitesimal phase and amplitude response functions, that is, the extended infinitesimal phase and amplitude response curves, which monitor the phase and amplitude shifts beyond the asymptotic state. Thus, the methodology presented yields an accurate description of the phase dynamics for perturbations not restricted to the limit cycle but to its attracting invariant manifold. Finally, we explore some strategies to reduce the dimension of the dynamics, including the reduction of the dynamics to the slow stable submanifold. We illustrate our methods by applying them to different three-dimensional single neuron and neural population models in neuroscience.

The uncoupling limit of identical Hopf bifurcations with an application to perceptual bistability.

We study the dynamics arising when two identical oscillators are coupled near a Hopf bifurcation where we assume a parameter ϵ uncouples the system at [Formula: see text]. Using a normal form for [Formula: see text] identical systems undergoing Hopf bifurcation, we explore the dynamical properties. Matching the normal form coefficients to a coupled Wilson-Cowan oscillator network gives an understanding of different types of behaviour that arise in a model of perceptual bistability. Notably, we find bistability between in-phase and anti-phase solutions that demonstrates the feasibility for synchronisation to act as the mechanism by which periodic inputs can be segregated (rather than via strong inhibitory coupling, as in the existing models). Using numerical continuation we confirm our theoretical analysis for small coupling strength and explore the bifurcation diagrams for large coupling strength, where the normal form approximation breaks down.