Global organization of dynamics in oscillatory heterogeneous excitable media

An oscillatory heterogeneous excitable medium undergoes a transition from periodic target patterns to a bursting rhythm driven by the spontaneous initiation and termination of spiral waves as coupling or density is reduced. We illustrate these phenomena in monolayers of chick embryonic heart cells using calcium-sensitive fluorescent dyes. These results are modeled in a heterogeneous cellular automaton in which the neighborhood of interaction and cell density is modified. Parameters that give rise to bursting rhythms are organized in distinct zones in parameter space, leading to a global organization that should be applicable to the dynamics in a large class of excitable media.     

Coherence resonance and stochastic synchronization in a nonlinear circuit near a subcritical Hopf bifurcation

We analyze noise-induced phenomena in nonlinear dynamical systems near a subcritical Hopf bifurcation. We investigate qualitative changes of probability distributions (stochastic bifurcations), coherence resonance, and stochastic synchronization. These effects are studied in dynamical systems for which a subcritical Hopf bifurcation occurs. We perform analytical calculations, numerical simulations and experiments on an electronic circuit. For the generalized Van der Pol model we uncover the similarities between the behavior of a self-sustained oscillator characterized by a subcritical Hopf bifurcation and an excitable system. The analogy is manifested through coherence resonance and stochastic synchronization. In particular, we show both experimentally and numerically that stochastic oscillations that appear due to noise in a system with hard excitation, can be partially synchronized even outside the oscillatory regime of the deterministic system.     

Stochastic phase dynamics and noise-induced mixed-mode oscillations in coupled oscillators

Synaptically coupled neurons show in-phase or antiphase synchrony depending on the chemical and dynamical nature of the synapse. Deterministic theory helps predict the phase differences between two phase-locked oscillators when the coupling is weak. In the presence of noise, however, deterministic theory faces difficulty when the coexistence of multiple stable oscillatory solutions occurs. We analyze the solution structure of two coupled neuronal oscillators for parameter values between a subcritical Hopf bifurcation point and a saddle node point of the periodic branch that bifurcates from the Hopf point, where a rich variety of coexisting solutions including asymmetric localized oscillations occurs. We construct these solutions via a multiscale analysis and explore the general bifurcation scenario using the lambda-omega model. We show for both excitatory and inhibitory synapses that noise causes important changes in the phase and amplitude dynamics of such coupled neuronal oscillators when multiple oscillatory solutions coexist. Mixed-mode oscillations occur when distinct bistable solutions are randomly visited. The phase difference between the coupled oscillators in the localized solution, coexisting with in-phase or antiphase solutions, is clearly represented in the stochastic phase dynamics.     

Noise-induced excitability in oscillatory media

A noise-induced phase transition to excitability is reported in oscillatory media with FitzHugh-Nagumo dynamics. This transition takes place via a noise-induced stabilization of a deterministically unstable fixed point of the local dynamics, while the overall phase-space structure of the system is maintained. Spatial coupling is required to prevent oscillations through suppression of fluctuations (via clustering in the case of local coupling). Thus, the joint action of coupling and noise leads to a different type of phase transition and results in a stabilization of the system. The resulting regime is shown to display characteristic traits of excitable media, such as stochastic resonance and wave propagation. This effect thus allows the transmission of signals through an otherwise globally oscillating medium.     

Noise-induced mixed-mode oscillations in a relaxation oscillator near the onset of a limit cycle

A detailed asymptotic study of the effect of small Gaussian white noise on a relaxation oscillator undergoing a supercritical Hopf bifurcation is presented. The analysis reveals an intricate stochastic bifurcation leading to several kinds of noise-driven mixed-mode oscillations at different levels of amplitude of the noise. In the limit of strong time-scale separation, five different scaling regimes for the noise amplitude are identified. As the noise amplitude is decreased, the dynamics of the system goes from the limit cycle due to self-induced stochastic resonance to the coherence resonance limit cycle, then to bursting relaxation oscillations, followed by rare clusters of several relaxation cycles (spikes), and finally to small-amplitude oscillations (or stable fixed point) with sporadic single spikes. These scenarios are corroborated by numerical simulations.     

Dynamical quorum sensing and synchronization in collections of excitable and oscillatory catalytic particles

We present experimental studies of interacting excitable and oscillatory catalytic particles in well-stirred and spatially distributed systems. A number of distinct paths to synchronized oscillatory behavior are described. We present an example of a Kuramoto type transition in a well-stirred system with a collective rhythm emerging on increasing the number density of oscillatory particles. Groups of spatially distributed oscillatory particles become entrained to a common frequency by organizing centers. Quorum sensing type transitions are found in populations of globally and locally coupled excitable particles, with a sharp transition from steady state to fully synchronized behavior at a critical density or group size.     

Control of coherence in excitable systems by the interplay of noise and time-delay

The control of coherence and spectral properties of noise-induced oscillations by time delayed feedback is studied in a FitzHugh-Nagumo system and analyzed by reduced non-Markovian models. A two-state approach is considered as an abstract simplification of this excitable system. Rest and excited states are characterized by different waiting time distributions. This non-Markovian approach allows one to predict quantitatively the increase of coherence measured by the correlation time, and the modulation of the main frequencies of the stochastic dynamics in dependence on the delay time below Hopf bifurcations. Beyond the Hopf bifurcation, bulk oscillations of an ensemble of excitable two-state rotators emerge in the onset of coherent activation in case of delayed mean field coupling of the ensemble.     

Chaotic patterns in a coupled oscillator-excitator biochemical cell system

In this paper we examine dynamical modes resulting from diffusion-like interaction of two model biochemical cells. Kinetics in each of the cells is given by the ICC model of calcium ions in the cytosol. Constraints for one of the cells are set so that it is excitable. One of the constraints in the other cell - a fraction of activated cell surface receptors-is varied so that the dynamics in the cell is either excitable or oscillatory or a stable focus. The cells are interacting via mass transfer and dynamics of the coupled system are studied as two parameters are varied-the fraction of activated receptors and the coupling strength. We find that (i) the excitator-excitator interaction does not lead to oscillatory patterns, (ii) the oscillator-excitator interaction leads to alternating phase-locked periodic and quasiperiodic regimes, well known from oscillator-oscillator interactions; torus breaking bifurcation generates chaos when the coupling strength is in an intermediate range, (iii) the focus-excitator interaction generates compound oscillations arranged as period adding sequences alternating with chaotic windows; the transition to chaos is accompanied by period doublings and folding of branches of periodic orbits and is associated with a Shilnikov homoclinic orbit. The nature of spontaneous self-organized oscillations in the focus-excitator range is discussed. (c) 1999 American Institute of Physics.     

Singular Hopf bifurcations and mixed-mode oscillations in a two-cell inhibitory neural network

Recent studies of a firing rate model for neural competition as observed in binocular rivalry and central pattern generators [R. Curtu, A. Shpiro, N. Rubin, J. Rinzel, Mechanisms for frequency control in neuronal competition models, SIAM J. Appl. Dyn. Syst. 7 (2) (2008) 609-649] showed that the variation of the stimulus strength parameter can lead to rich and interesting dynamics. Several types of behavior were identified such as: fusion, equivalent to a steady state of identical activity levels for both neural units; oscillations due to either an escape or a release mechanism; and a winner-take-all state of bistability. The model consists of two neural populations interacting through reciprocal inhibition, each endowed with a slow negative-feedback process in the form of spike frequency adaptation. In this paper we report the occurrence of another complex oscillatory pattern, the mixed-mode oscillations (MMOs). They exist in the model at the transition between the relaxation oscillator dynamical regime and the winner-take-all regime. The system distinguishes itself from other neuronal models where MMOs were found by the following interesting feature: there is no autocatalysis involved (as in the examples of voltage-gated persistent inward currents and/or intrapopulation recurrent excitation) and therefore the two cells in the network are not intrinsic oscillators; the oscillations are instead a combined result of the mutual inhibition and the adaptation. We prove that the MMOs are due to a singular Hopf bifurcation point situated in close distance to the transition point to the winner-take-all case. We also show that in the vicinity of the singular Hopf other types of bifurcations exist and we construct numerically the corresponding diagrams.     

Dominant phase-advanced driving analysis of self-sustained oscillations in biological networks

Oscillatory behaviors can be ubiquitously observed in various systems. Biological rhythms are significant in governing living activities of all units. The emergence of biological rhythms is the consequence of large numbers of units. In this paper we discuss several important examples of sustained oscillations in biological media, where the unit composed in the system does not possess the oscillation behavior. The dominant phase-advanced driving method is applied to study the skeletons and oscillatory organizing motifs in excitable networks and gene regulatory networks.     

From simple to complex oscillatory behavior in metabolic and genetic control networks

We present an overview of mechanisms responsible for simple or complex oscillatory behavior in metabolic and genetic control networks. Besides simple periodic behavior corresponding to the evolution toward a limit cycle we consider complex modes of oscillatory behavior such as complex periodic oscillations of the bursting type and chaos. Multiple attractors are also discussed, e.g., the coexistence between a stable steady state and a stable limit cycle (hard excitation), or the coexistence between two simultaneously stable limit cycles (birhythmicity). We discuss mechanisms responsible for the transition from simple to complex oscillatory behavior by means of a number of models serving as selected examples. The models were originally proposed to account for simple periodic oscillations observed experimentally at the cellular level in a variety of biological systems. In a second stage, these models were modified to allow for complex oscillatory phenomena such as bursting, birhythmicity, or chaos. We consider successively (1) models based on enzyme regulation, proposed for glycolytic oscillations and for the control of successive phases of the cell cycle, respectively; (2) a model for intracellular Ca(2+) oscillations based on transport regulation; (3) a model for oscillations of cyclic AMP based on receptor desensitization in Dictyostelium cells; and (4) a model based on genetic regulation for circadian rhythms in Drosophila. Two main classes of mechanism leading from simple to complex oscillatory behavior are identified, namely (i) the interplay between two endogenous oscillatory mechanisms, which can take multiple forms, overt or more subtle, depending on whether the two oscillators each involve their own regulatory feedback loop or share a common feedback loop while differing by some related process, and (ii) self-modulation of the oscillator through feedback from the system's output on one of the parameters controlling oscillatory behavior. However, the latter mechanism may also be viewed as involving the interplay between two feedback processes, each of which might be capable of producing oscillations. Although our discussion primarily focuses on the case of autonomous oscillatory behavior, we also consider the case of nonautonomous complex oscillations in a model for circadian oscillations subjected to periodic forcing by a light-dark cycle and show that the occurrence of entrainment versus chaos in these conditions markedly depends on the wave form of periodic forcing. (c) 2001 American Institute of Physics.     

Chaotically spiking canards in an excitable system with 2D inertial fast manifolds

We introduce a new class of excitable systems with two-dimensional fast dynamics that includes inertia. A novel transition from excitability to relaxation oscillations is discovered where the usual Hopf bifurcation is followed by a cascade of period doubled and chaotic small excitable attractors and, as they grow, by a new type of canard explosion where a small chaotic background erratically but deterministically triggers excitable spikes. This scenario is also found in a model for a nonlinear Fabry-Perot cavity with one pendular mirror.     

Noise-induced synchronous stochastic oscillations small scale cultured heart-cell networks

This paper reports that the synchronous integer multiple oscillations of heart-cell networks or clusters are observed in the biology experiment. The behaviour of the integer multiple rhythm is a transition between super- and sub- threshold oscillations, the stochastic mechanism of the transition is identified. The similar synchronized oscillations are theoretically reproduced in the stochastic network composed of heterogeneous cells whose behaviours are chosen as excitable or oscillatory states near a Hopf bifurcation point. The parameter regions of coupling strength and noise density that the complex oscillatory rhythms can be simulated are identified. The results show that the rhythm results from a simple stochastic alternating process between super- and sub-threshold oscillations. Studies on single heart cells forming these clusters reveal excitable or oscillatory state nearby a Hopf bifurcation point underpinning the stochastic alternation. In discussion, the results are related to some abnormal heartbeat rhythms such as the sinus arrest.     

Sustainment and Controlment of Noise-Induced Circadian Oscillations in <Emphasis Type="Italic">Neurospora</Emphasis>: Noise and External Signal Effects

The effect of light noise on a Neurospora circadian clock system in the steady states is investigated. It is found that the circadian oscillations could be induced by light noise, leading to various resonance phenomena including internal signal stochastic resonance (ISSR) and ISSR without tuning in the system. The strength of ISSR could be significantly reinforced with the decrease of the distance of the control parameter to the Hopf bifurcation point of the system. The fundamental frequency of noise-induced circadian oscillations almost does not change with the increment of light noise intensity, which implies that the Neurospora system could sustain intrinsic circadian rhythms. In addition, the ISSR and ISSR without tuning could be both amplified, suppressed or destroyed by tuning the frequency or amplitude of external signal.     

Bifurcation analysis of a normal form for excitable media: are stable dynamical alternans on a ring possible?

We present a bifurcation analysis of a normal form for traveling waves in one-dimensional excitable media. The normal form that has been recently proposed on phenomenological grounds is given in the form of a differential delay equation. The normal form exhibits a symmetry-preserving Hopf bifurcation that may coalesce with a saddle node in a Bogdanov-Takens point, and a symmetry-breaking spatially inhomogeneous pitchfork bifurcation. We study here the Hopf bifurcation for the propagation of a single pulse in a ring by means of a center manifold reduction, and for a wave train by means of a multiscale analysis leading to a real Ginzburg-Landau equation as the corresponding amplitude equation. Both the center manifold reduction and the multiscale analysis show that the Hopf bifurcation is always subcritical independent of the parameters. This may have links to cardiac alternans, which have so far been believed to be stable oscillations emanating from a supercritical bifurcation. We discuss the implications for cardiac alternans and revisit the instability in some excitable media where the oscillations had been believed to be stable. In particular, we show that our condition for the onset of the Hopf bifurcation coincides with the well known restitution condition for cardiac alternans.     

Noise induced complexity: from subthreshold oscillations to spiking in coupled excitable systems

We study the stochastic dynamics of an ensemble of N globally coupled excitable elements. Each element is modeled by a FitzHugh-Nagumo oscillator and is disturbed by independent Gaussian noise. In simulations of the Langevin dynamics we characterize the collective behavior of the ensemble in terms of its mean field and show that with the increase of noise the mean field displays a transition from a steady equilibrium to global oscillations and then, for sufficiently large noise, back to another equilibrium. In the course of this transition diverse regimes of collective dynamics ranging from periodic subthreshold oscillations to large-amplitude oscillations and chaos are observed. In order to understand the details and mechanisms of these noise-induced dynamics we consider the thermodynamic limit N-->infinity of the ensemble, and derive the cumulant expansion describing temporal evolution of the mean field fluctuations. In Gaussian approximation this allows us to perform the bifurcation analysis; its results are in good qualitative agreement with dynamical scenarios observed in the stochastic simulations of large ensembles.     

From oscillations to excitability: A case study in spatially extended systems

This volume is devoted to the presentation of the main contributions to the workshop "From oscillations to excitability: A case study in spatially extended systems," organized by the authors in Nice in June 1993. It gives an overview of the current research on spatiotemporal patterns in a wide range of systems that display self-oscillatory or excitable behavior. It tries to give a better understanding of the transition from the oscillatory to the excitable regime and of its effect on the properties of spiral waves, and to fill the gap between the theories and concepts used to describe both regimes in the so-called "active media."     

Stochastic Signal Induced Multiple Spatial Coherence Resonances and Spiral Waves in Excitable Media

Multiple spatial coherence resonances and spiral waves with various temporal-spatial structures are simulated in a two-dimensional network of excitable cells driven by a stochastic signal. The relationship between the multiple resonances and correspondingly different transitions of the spiral wave are elucidated. The results further provide a possible approach of applications of stochastic signal to evoke pattern transitions in excitable media.