Chimera states: Coexistence of coherence and incoherence in networks of coupled oscillators

A chimera state is a spatio-temporal pattern in a network of identical coupled oscillators in which synchronous and asynchronous oscillation coexist. This state of broken symmetry, which usually coexists with a stable spatially symmetric state, has intrigued the nonlinear dynamics community since its discovery in the early 2000s. Recent experiments have led to increasing interest in the origin and dynamics of these states. Here we review the history of research on chimera states and highlight major advances in understanding their behaviour.Comment: 26 pages, 3 figure

Membrane resonance enables stable and robust gamma oscillations

Neuronal mechanisms underlying beta/gamma oscillations (20-80 Hz) are not completely understood. Here, we show that in vivo beta/gamma oscillations in the cat visual cortex sometimes exhibit remarkably stable frequency even when inputs fluctuate dramatically. Enhanced frequency stability is associated with stronger oscillations measured in individual units and larger power in the local field potential. Simulations of neuronal circuitry demonstrate that membrane properties of inhibitory interneurons strongly determine the characteristics of emergent oscillations. Exploration of networks containing either integrator or resonator inhibitory interneurons revealed that: (i) Resonance, as opposed to integration, promotes robust oscillations with large power and stable frequency via a mechanism called RING (Resonance INduced Gamma); resonance favors synchronization by reducing phase delays between interneurons and imposes bounds on oscillation cycle duration; (ii) Stability of frequency and robustness of the oscillation also depend on the relative timing of excitatory and inhibitory volleys within the oscillation cycle; (iii) RING can reproduce characteristics of both Pyramidal INterneuron Gamma (PING) and INterneuron Gamma (ING), transcending such classifications; (iv) In RING, robust gamma oscillations are promoted by slow but are impaired by fast inputs. Results suggest that interneuronal membrane resonance can be an important ingredient for generation of robust gamma oscillations having stable frequency

Time-delayed feedback in neurosystems

The influence of time delay in systems of two coupled excitable neurons is studied in the framework of the FitzHugh-Nagumo model. Time-delay can occur in the coupling between neurons or in a self-feedback loop. The stochastic synchronization of instantaneously coupled neurons under the influence of white noise can be deliberately controlled by local time-delayed feedback. By appropriate choice of the delay time synchronization can be either enhanced or suppressed. In delay-coupled neurons, antiphase oscillations can be induced for sufficiently large delay and coupling strength. The additional application of time-delayed self-feedback leads to complex scenarios of synchronized in-phase or antiphase oscillations, bursting patterns, or amplitude death.Comment: 13 pages, 13 figure

Delay-induced patterns in a two-dimensional lattice of coupled oscillators

We show how a variety of stable spatio-temporal periodic patterns can be created in 2D-lattices of coupled oscillators with non-homogeneous coupling delays. A "hybrid dispersion relation" is introduced, which allows studying the stability of time-periodic patterns analytically in the limit of large delay. The results are illustrated using the FitzHugh-Nagumo coupled neurons as well as coupled limit cycle (Stuart-Landau) oscillators

Role of delay for the symmetry in the dynamics of networks

PACS number(s): 05.45.Xt, 89.75.Kd, 89.75.Hc, 02.30.KsThe symmetry in a network of oscillators determines the spatiotemporal patterns of activity that can emerge. We study how a delay in the coupling affects symmetry-breaking and -restoring bifurcations. We are able to draw general conclusions in the limit of long delays. For one class of networks we derive a criterion that predicts that delays have a symmetrizing effect. Moreover, we demonstrate that for any network admitting a steady-state solution, a long delay can solely advance the first bifurcation point as compared to the instantaneous-coupling regime.This work was partially supported by the Interuniversity Attraction Poles program Photonics@be of the Belgian Science Policy Office under Grant No. IAP VI-10 by MICINN (Spain) under Project No. DeCoDicA (TEC2009-14101) and by the project PHOCUS (EU FET-Open Grant No. 240763). S. Yanchuk and P. Perlikowski are gratefully acknowledged for fruitful discussions.Peer reviewe

Modelling delta-notch perturbations during zebrafish somitogenesis

The discovery over the last 15 years of molecular clocks and gradients in the pre-somitic mesoderm of numerous vertebrate species has added significant weight to Cooke and Zeeman's ‘clock and wavefront’ model of somitogenesis, in which a travelling wavefront determines the spatial position of somite formation and the somitogenesis clock controls periodicity (Cooke and Zeeman, 1976). However, recent high-throughput measurements of spatiotemporal patterns of gene expression in different zebrafish mutant backgrounds allow further quantitative evaluation of the clock and wavefront hypothesis. In this study we describe how our recently proposed model, in which oscillator coupling drives the propagation of an emergent wavefront, can be used to provide mechanistic and testable explanations for the following observed phenomena in zebrafish embryos: (a) the variation in somite measurements across a number of zebrafish mutants; (b) the delayed formation of somites and the formation of ‘salt and pepper’ patterns of gene expression upon disruption of oscillator coupling; and (c) spatial correlations in the ‘salt and pepper’ patterns in Delta-Notch mutants. In light of our results, we propose a number of plausible experiments that could be used to further test the model

Effect of Distributed Delays in Systems of Coupled Phase Oscillators

Communication delays are common in many complex systems. It has been shown that these delays cannot be neglected when they are long enough compared to other timescales in the system. In systems of coupled phase oscillators discrete delays in the coupling give rise to effects such as multistability of steady states. However, variability in the communication times inherent to many processes suggests that the description with discrete delays maybe insufficient to capture all effects of delays. An interesting example of the effects of communication delays is found during embryonic development of vertebrates. A clock based on biochemical reactions inside cells provides the periodicity for the successive and robust formation of somites, the embryonic precursors of vertebrae, ribs and some skeletal muscle. Experiments show that these cellular clocks communicate in order to synchronize their behavior. However, in cellular systems, fluctuations and stochastic processes introduce a variability in the communication times. Here we account for such variability by considering the effects of distributed delays. Our approach takes into account entire intervals of past states, and weights them according to a delay distribution. We find that the stability of the fully synchronized steady state with zero phase lag does not depend on the shape of the delay distribution, but the dynamics when responding to small perturbations about this steady state do. Depending on the mean of the delay distribution, a change in its shape can enhance or reduce the ability of these systems to respond to small perturbations about the phase-locked steady state, as compared to a discrete delay with a value equal to this mean. For synchronized steady states with non-zero phase lag we find that the stability of the steady state can be altered by changing the shape of the delay distribution. We conclude that the response to a perturbation in systems of phase oscillators coupled with discrete delays has a sharper functional dependence on the mean delay than in systems with distributed delays in the coupling. The strong dependence of the coupling on the mean delay time is partially averaged out by distributed delays that take into account intervals of the past.:Abstract i Acknowledgement iii I. INTRODUCTION 1. Coupled Phase Oscillators Enter the Stage 5 1.1. Adjusting rhythms – synchronization . . . . . . . . . . . . . . . . . . 5 1.2. Historical remarks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 1.3. Reducing variables – phase models . . . . . . . . . . . . . . . . . . . . 9 1.4. The Kuramoto order parameter . . . . . . . . . . . . . . . . . . . . . . 10 1.5. Who talks to whom – coupling topologies . . . . . . . . . . . . . . . . 12 2. Coupled Phase Oscillators with Delay in the Coupling 15 2.1. Communication needs time – coupling delays . . . . . . . . . . . . . . 15 2.1.1. Discrete delays consider one past time . . . . . . . . . . . . . . 16 2.1.2. Distributed delays consider multiple past times . . . . . . . . 17 2.2. Coupled phase oscillators with discrete delay . . . . . . . . . . . . . . 18 2.2.1. Phase locked steady states with no phase lags . . . . . . . . . 18 2.2.2. m-twist solutions: phase-locked steady states with phase lags 21 3. The Vertebrate Segmentation Clock – What Provides the Rhythm? 25 3.1. The clock and wavefront mechanism . . . . . . . . . . . . . . . . . . . 26 3.2. Cyclic gene expression on the cellular and the tissue level . . . . . . 27 3.3. Coupling by Delta-Notch signalling . . . . . . . . . . . . . . . . . . . . 29 3.4. The Delayed Coupling Theory . . . . . . . . . . . . . . . . . . . . . . . 30 3.5. Discrete delay is an approximation – is it sufficient? . . . . . . . . . 32 4. Outline of the Thesis 33 II. DISTRIBUTED DELAYS 5. Setting the Stage for Distributed Delays 37 5.1. Model equations with distributed delays . . . . . . . . . . . . . . . . . 37 5.2. How we include distributed delays . . . . . . . . . . . . . . . . . . . . 38 5.3. Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40 6. The Phase-Locked Steady State Solution 41 6.1. Global frequency of phase-locked steady states . . . . . . . . . . . . . 41 6.2. Linear stability of the steady state . . . . . . . . . . . . . . . . . . . . 42 6.3. Linear dynamics of the perturbation – the characteristic equation . 43 6.4. Summary and application to the Delayed Coupling Theory . . . . . . 50 7. Dynamics Close to the Phase-Locked Steady State 53 7.1. The response to small perturbations . . . . . . . . . . . . . . . . . . . 53 7.2. Relation between order parameter and perturbation modes . . . . . 54 7.3. Perturbation dynamics in mean-field coupled systems . . . . . . . . 56 7.4. Nearest neighbour coupling with periodic boundary conditions . . . 62 7.4.1. How variance and skewness influence synchrony dynamics . 73 7.4.2. The dependence of synchrony dynamics on the number of oscillators . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83 7.5. Synchrony dynamics in systems with arbitrary coupling topologies . 88 7.6. Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93 8. The m-twist Steady State Solution on a Ring 95 8.1. Global frequency of m-twist steady states . . . . . . . . . . . . . . . . 95 8.2. Linear stability of m-twist steady states . . . . . . . . . . . . . . . . . 97 8.3. Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103 9. Dynamics Approaching the m-twist Steady States 105 9.1. Relation between order parameter and perturbation modes . . . . . 105 9.2. Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109 10.Conclusions and Outlook 111 vi III. APPENDICES A. 119 A.1. Distribution composed of two adjacent boxcar functions . . . . . . . 119 A.2. The gamma distribution . . . . . . . . . . . . . . . . . . . . . . . . . . 124 A.3. Distribution composed of two Dirac delta peaks . . . . . . . . . . . . 125 A.4. Gerschgorin’s circle theorem . . . . . . . . . . . . . . . . . . . . . . . . 127 A.5. The Lambert W function . . . . . . . . . . . . . . . . . . . . . . . . . . 127 A.6. Roots of unity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 127 B. Simulation methods 12