Filament Tension and Phase Locking of Meandering Scroll Waves.

Meandering spiral waves are often observed in excitable media such as the Belousov-Zhabotinsky reaction and cardiac tissue. We derive a theory for drift dynamics of meandering rotors in general reaction-diffusion systems and apply it to two types of external disturbances: an external field and curvature-induced drift in three dimensions. We find two distinct regimes: with small filament curvature, meandering scroll waves exhibit filament tension, whose sign determines the stability and drift direction. In the regimes of strong external fields or meandering motion close to resonance, however, phase locking of the meander pattern is predicted and observed.

Drift of scroll waves in thin layers caused by thickness features: asymptotic theory and numerical simulations.

A scroll wave in a very thin layer of excitable medium is similar to a spiral wave, but its behavior is affected by the layer geometry. We identify the effect of sharp variations of the layer thickness, which is separate from filament tension and curvature-induced drifts described earlier. We outline a two-step asymptotic theory describing this effect, including asymptotics in the layer thickness and calculation of the drift of so-perturbed spiral waves using response functions. As specific examples, we consider drift of scrolls along thickness steps, ridges, ditches, and disk-shaped thickness variations. Asymptotic predictions agree with numerical simulations.

Orbital motion of spiral waves in excitable media.

Spiral waves in active media react to small perturbations as particlelike objects. Here we apply asymptotic theory to the interaction of spiral waves with a localized inhomogeneity, which leads to a novel prediction: drift of the spiral rotation center along circular orbits around the inhomogeneity. The stationary orbits have fixed radii and alternating stability, determined by the properties of the bulk medium and the type of inhomogeneity, while the drift speed along an orbit depends on the strength of the inhomogeneity. Direct numerical simulations confirm the validity and robustness of the theoretical predictions and show that these unexpected effects should be observable in experiment.

Alternative stable scroll waves and conversion of autowave turbulence.

Rotating spiral and scroll waves (vortices) are investigated in the FitzHugh-Nagumo model of excitable media. The focus is on a parameter region in which there exists bistability between alternative stable vortices with distinct periods. Response functions are used to predict the filament tension of the alternative scrolls and it is shown that the slow-period scroll has negative filament tension, while the filament tension of the fast-period scroll changes sign within a hysteresis loop. The predictions are confirmed by direct simulations. Further investigations show that the slow-period scrolls display features similar to delayed after-depolarization and tend to develop into turbulence similar to ventricular fibrillation (VF). Scrolls with positive filament tension collapse or stabilize, similar to monomorphic ventricular tachycardia (VT). Perturbations, such as boundary interaction or shock stimulus, can convert the vortex with negative filament tension into the vortex with positive filament tension. This may correspond to transition from VF to VT unrelated to pinning.

Computation of the response functions of spiral waves in active media.

Rotating spiral waves are a form of self-organization observed in spatially extended systems of physical, chemical, and biological natures. A small perturbation causes gradual change in spatial location of spiral's rotation center and frequency, i.e., drift. The response functions (RFs) of a spiral wave are the eigenfunctions of the adjoint linearized operator corresponding to the critical eigenvalues lambda=0,+/-iomega. The RFs describe the spiral's sensitivity to small perturbations in the way that a spiral is insensitive to small perturbations where its RFs are close to zero. The velocity of a spiral's drift is proportional to the convolution of RFs with the perturbation. Here we develop a regular and generic method of computing the RFs of stationary rotating spirals in reaction-diffusion equations. We demonstrate the method on the FitzHugh-Nagumo system and also show convergence of the method with respect to the computational parameters, i.e., discretization steps and size of the medium. The obtained RFs are localized at the spiral's core.

Response function framework for the dynamics of meandering or large-core spiral waves and modulated traveling waves.

In many oscillatory or excitable systems, dynamical patterns emerge which are stationary or periodic in a moving frame of reference. Examples include traveling waves or spiral waves in chemical systems or cardiac tissue. We present a unified theoretical framework for the drift of such patterns under small external perturbations, in terms of overlap integrals between the perturbation and the adjoint critical eigenfunctions of the linearized operator (i.e., response functions). For spiral waves, the finite radius of the spiral tip trajectory and spiral wave meander are taken into account. Different coordinate systems can be chosen, depending on whether one wants to predict the motion of the spiral-wave tip, the time-averaged tip path, or the center of the meander flower. The framework is applied to analyze the drift of a meandering spiral wave in a constant external field in different regimes.

Control of scroll-wave turbulence using resonant perturbations.

Turbulence of scroll waves is a sort of spatiotemporal chaos that exists in three-dimensional excitable media. Cardiac tissue and the Belousov-Zhabotinsky reaction are examples of such media. In cardiac tissue, chaotic behavior is believed to underlie fibrillation which, without intervention, precedes cardiac death. In this study we investigate suppression of the turbulence using stimulation of two different types, "modulation of excitability" and "extra transmembrane current." With cardiac defibrillation in mind, we used a single pulse as well as repetitive extra current with both constant and feedback controlled frequency. We show that turbulence can be terminated using either a resonant modulation of excitability or a resonant extra current. The turbulence is terminated with much higher probability using a resonant frequency perturbation than a nonresonant one. Suppression of the turbulence using a resonant frequency is up to fifty times faster than using a nonresonant frequency, in both the modulation of excitability and the extra current modes. We also demonstrate that resonant perturbation requires strength one order of magnitude lower than that of a single pulse, which is currently used in clinical practice to terminate cardiac fibrillation. Our results provide a robust method of controlling complex chaotic spatiotemporal processes. Resonant drift of spiral waves has been studied extensively in two dimensions, however, these results show for the first time that it also works in three dimensions, despite the complex nature of the scroll wave turbulence.

Wave-particle dualism of spiral waves dynamics.

We demonstrate and explain a wave-particle dualism of such classical macroscopic phenomena as spiral waves in active media. That means although spiral waves appear as nonlocal processes involving the whole medium, they respond to small perturbations as effectively localized entities. The dualism appears as an emergent property of a nonlinear field and is mathematically expressed in terms of the spiral waves response functions, which are essentially nonzero only in the vicinity of the spiral wave core. Knowledge of the response functions allows quantitatively accurate prediction of the spiral wave drift due to small perturbations of any nature, which makes them as fundamental characteristics for spiral waves as mass is for the condensed matter.

Computation of the drift velocity of spiral waves using response functions.

Rotating spiral waves are a form of self-organization observed in spatially extended systems of physical, chemical, and biological nature. In the presence of a small perturbation, the spiral wave's center of rotation and fiducial phase may change over time, i.e., the spiral wave drifts. In linear approximation, the velocity of the drift is proportional to the convolution of the perturbation with the spiral's response functions, which are the eigenfunctions of the adjoint linearized operator corresponding to the critical eigenvalues λ=0,±iω . Here, we demonstrate that the response functions give quantitatively accurate prediction of the drift velocities due to a variety of perturbations: a time dependent, periodic perturbation (inducing resonant drift); a rotational symmetry-breaking perturbation (inducing electrophoretic drift); and a translational symmetry-breaking perturbation (inhomogeneity induced drift) including drift due to a gradient, stepwise, and localized inhomogeneity. We predict the drift velocities using the response functions in FitzHugh-Nagumo and Barkley models, and compare them with the velocities obtained in direct numerical simulations. In all cases good quantitative agreement is demonstrated.

Asymptotic properties of mathematical models of excitability.

We analyse small parameters in selected models of biological excitability, including Hodgkin-Huxley (Hodgkin & Huxley 1952 J. Physiol.117, 500-544) model of nerve axon, Noble (Noble 1962 J. Physiol.160, 317-352) model of heart Purkinje fibres and Courtemanche et al. (Courtemanche et al. 1998 Am. J. Physiol.275, H301-H321) model of human atrial cells. Some of the small parameters are responsible for differences in the characteristic time-scales of dynamic variables, as in the traditional singular perturbation approaches. Others appear in a way which makes the standard approaches inapplicable. We apply this analysis to study the behaviour of fronts of excitation waves in spatially extended cardiac models. Suppressing the excitability of the tissue leads to a decrease in the propagation speed, but only to a certain limit; further suppression blocks active propagation and leads to a passive diffusive spread of voltage. Such a dissipation may happen if a front propagates into a tissue recovering after a previous wave, e.g. re-entry. A dissipated front does not recover even when the excitability restores. This has no analogy in FitzHugh-Nagumo model and its variants, where fronts can stop and then start again. In two spatial dimensions, dissipation accounts for breakups and self-termination of re-entrant waves in excitable media with Courtemanche et al. kinetics.

Resonant drift of spiral waves in the complex ginzburg-landau equation.

Weak periodic external perturbations of an autowave medium can cause large-distance directed motion of the spiral wave. This happens when the period of the perturbation coincides with, or is close to the rotation period of a spiral wave, or its multiple. Such motion is called resonant or parametric drift. It may be used for low-voltage defibrillation of heart tissue. Theory of the resonant drift exists, but so far was used only qualitatively. In this paper, we show good quantitative agreement of the theory with direct numerical simulations. This is done for Complex Ginzburg-Landau Equation, one of the simplest autowave models.

Effects of shear flows on nonlinear waves in excitable media.

If an excitable medium is moving with relative shear, the waves of excitation may be broken by the motion. We consider such breaks for the case of a constant linear shear flow. The mechanisms and conditions for the breaking of solitary waves and wavetrains are essentially different: the solitary waves require the velocity gradient to exceed a certain threshold, whilst the breaking of repetitive wavetrains happens for arbitrarily small velocity gradients. Since broken waves evolve into new spiral wave sources, this leads to spatio-temporal irregularity.

Spatiotemporal irregularity in an excitable medium with shear flow.

We consider an excitable medium moving with relative shear, subjected to a localized disturbance that in a stationary medium would produce a pair of spiral waves. The spiral waves so created are distorted and then broken by the motion of the medium. Such breaks generate new spiral waves, and so a "chain reaction" of spiral wave births and deaths is observed. This leads to a complicated spatiotemporal pattern, the "frazzle gas" [term suggested by Markus et al., Nature (London) 371, 402 (1994)], which eventually fills the whole medium. In this paper, we display and interpret the main features of the pattern.