Covariant stringlike dynamics of scroll wave filaments in anisotropic cardiac tissue

It has been hypothesized that stationary scroll wave filaments in cardiac tissue describe a geodesic in a curved space whose metric is the inverse diffusion tensor. Several numerical studies support this hypothesis, but no analytical proof has been provided yet for general anisotropy. In this Letter, we derive dynamic equations for the filament in the case of general anisotropy. These equations are covariant under general spatial coordinate transformations and describe the motion of a stringlike object in a curved space whose metric tensor is the inverse diffusion tensor. Therefore the behavior of scroll wave filaments in excitable media with anisotropy is similar to the one of cosmic strings in a curved universe. Our dynamic equations are valid for thin filaments and for general anisotropy. We show that stationary filaments obey the geodesic equation.     

Eikonal formulation of the minimal principle for scroll wave filaments

Recently, Wellner et al. [Proc. Natl. Acad. Sci. U.S.A. 99, 8015 (2002)]] proposed a principle for predicting a stable scroll wave filament shape as a geodesic in a 3D space with a metric determined by the inverse diffusivity tensor of the medium. Using the Hamilton-Jacobi theory we show that this geodesic is the shortest path for a wave propagating through the medium. This allows the use of shortest path algorithms to predict filament shapes, which we confirm numerically for a medium with orthotropic anisotropy. Our method can be used in cardiac tissue experiments since it does not require knowledge of the tissue anisotropy.     

Scroll wave dynamics in a model of the heterogeneous heart

Scroll waves are found in physical, chemical and biological systems and underlie many significant processes including life-threatening cardiac arrhythmias. The theory of scroll waves predicts scroll wave dynamics should be substantially affected by heterogeneity of cardiac tissue together with other factors including shape and anisotropy. In this study, we used our recently developed analytical model of the human ventricle to identify effects of shape, anisotropy, and regional heterogeneity of myocardium on scroll wave dynamics. We found that the main effects of apical-base heterogeneity were an increased scroll wave drift velocity and a shift towards the region of maximum action potential duration. We also found that transmural heterogeneity does not substantially affect scroll wave dynamics and only in extreme cases changes the attractor position.     

Predicting filament drift in twisted anisotropy

Excitable media with twisted anisotropy have recently been attracting significant interest because of their applicability to wave propagation in heart tissue. Here we consider the dynamics of an intramural scroll wave whose filament lies initially within an arbitrary layer of mutually parallel cardiac fibers, and drifts parallel to itself from layer to layer. Earlier simulations have demonstrated that such a filament stabilizes in a layer whose fiber direction is the same as its own. In the present paper we analytically derive the trajectory of the filament, and obtain good agreement with earlier numerical data. For sufficiently sparse scrolls, our analysis predicts an equilibrium alignment perpendicular rather than parallel to the fibers.     

Scroll waves in spherical shell geometries

The evolution of scroll waves in excitable media with spherical shell geometries is studied as a function of shell thickness and outer radius. The motion of scroll wave filaments that are the locii of phaseless points in the medium and organize the wave pattern is investigated. When the inner radius is sufficiently large the filaments remain attached to both the inner and outer surfaces. The minimum size of the sphere that supports spiral waves and the maximum number of spiral waves that can be sustained on a sphere of given size are determined for both regular and random initial distributions. When the inner radius is too small to support spiral waves the filaments detach from the inner surface and form a curved filament connecting the two spiral tips in the surface. In certain parameter domains the filament is an arc of a circle that shrinks with constant shape. For parameter values close to the meandering border, the filament grows and collisions with the sphere walls lead to turbulent filament dynamics. (c) 2001 American Institute of Physics.     

Influence of cardiac tissue anisotropy on re-entrant activation in computational models of ventricular fibrillation

The aim of this study was to establish the role played by anisotropic diffusion in (i) the number of filaments and epicardial phase singularities that sustain ventricular fibrillation in the heart, (ii) the lifetimes of filaments and phase singularities, and (iii) the creation and annihilation dynamics of filaments and phase singularities. A simplified monodomain model of cardiac tissue was used, with membrane excitation described by a simplified 3-variable model. The model was configured so that a single re-entrant wave was unstable, and fragmented into multiple re-entrant waves. Re-entry was then initiated in tissue slabs with varying anisotropy ratio. The main findings of this computational study are: (i) anisotropy ratio influenced the number of filaments sustaining simulated ventricular fibrillation, with more filaments present in simulations with smaller values of transverse diffusion coefficient, (ii) each re-entrant filament was associated with around 0.9 phase singularities on the surface of the slab geometry, (iii) phase singularities were longer lived than filaments, and (iv) the creation and annihilation of filaments and phase singularities were linear functions of the number of filaments and phase singularities, and these relationships were independent of the anisotropy ratio. This study underscores the important role played by tissue anisotropy in cardiac ventricular fibrillation.     

Effects of reduced discrete coupling on filament tension in excitable media

Wave propagation in the heart has a discrete nature, because it is mediated by discrete intercellular connections via gap junctions. Although effects of discreteness on wave propagation have been studied for planar traveling waves and vortexes (spiral waves) in two dimensions, its possible effects on vortexes (scroll waves) in three dimensions are not yet explored. In this article, we study the effect of discrete cell coupling on the filament dynamics in a generic model of an excitable medium. We find that reduced cell coupling decreases the line tension of scroll wave filaments and may induce negative filament tension instability in three-dimensional excitable lattices.     

The generalized Einstein-Maxwell theory of gravitation

A new Lagrangian theory of gravitation in which the metric and the arbitrary affine connection are regarded as independent field variables has been considered. Making use of the pure geometrical objects only from the variational principle the empty field equations are derived. It is shown that the metric obeys the ordinary Einstein equations of general relativity. However, the covariant derivative of the metric tensor does not vanish, so that the vector's length is generally nonintegrable under the parallel displacement. The torsion trace vector turns out to be the natural dynamical variable, satisfying the Maxwell-like equations with tensor of homothetic curvature as the Maxwell tensor. The equations of motion are explored; they are shown to be identical to the motion of electric charge under the Lorentz force. The conservation laws are discussed.     

Three-dimensional spiral waves in an excitable reaction system: initiation and dynamics of scroll rings and scroll ring pairs

We report experimental results on spiral and scroll waves in the 1,4-cyclohexanedione Belousov-Zhabotinsky reaction. The propagating concentration waves are detected by two-dimensional photometry and optical tomography. Wave pulses can disappear in front-to-front and front-to-back collisions. This anomaly causes the nucleation of vortices from collisions of three nonrotating waves. In three-dimensional systems, these vortices are scroll rings that rotate around initially circular filaments. Depending on reactant concentrations, the filaments shrink or expand indicating positive and negative filament tensions, respectively. Shrinkage results in vortex annihilation. Expansion is accompanied by filament buckling and bending, which is interpreted as developing Winfree turbulence. We also describe the initiation of scroll ring pairs in four-wave collisions. The two filaments are stacked on top of each other and their motion suggests filament repulsion.     

Intermittent self-organization of scroll wave turbulence in three-dimensional excitable media

We study the asymptotic behavior of scroll wave turbulence in large three-dimensional excitable media modeled by FitzHugh-Nagumo equations. The focus is on the type of turbulence caused by negative tension of scroll wave filaments, which is considered to be one of the mechanisms of cardiac fibrillation. We discovered that the initial increase in turbulence complexity can be followed by intermittent self-organization, when complex filament tangles are replaced by a small number of relatively stable triple filament strands. The intermittency is the result of a competition between the destabilizing effect of negative tension and mutual attraction of filaments with similar orientation.     

Dynamic field theory and equations of motion in cosmology

We discuss a field-theoretical approach based on general-relativistic variational principle to derive the covariant field equations and hydrodynamic equations of motion of baryonic matter governed by cosmological perturbations of dark matter and dark energy. The action depends on the gravitational and matter Lagrangian. The gravitational Lagrangian depends on the metric tensor and its first and second derivatives. The matter Lagrangian includes dark matter, dark energy and the ordinary baryonic matter which plays the role of a bare perturbation. The total Lagrangian is expanded in an asymptotic Taylor series around the background cosmological manifold defined as a solution of Einstein’s equations in the form of the Friedmann–Lemaître–Robertson–Walker (FLRW) metric tensor. The small parameter of the decomposition is the magnitude of the metric tensor perturbation. Each term of the series expansion is gauge-invariant and all of them together form a basis for the successive post-Friedmannian approximations around the background metric. The approximation scheme is covariant and the asymptotic nature of the Lagrangian decomposition does not require the post-Friedmannian perturbations to be small though computationally it works the most effectively when the perturbed metric is close enough to the background FLRW metric. The temporal evolution of the background metric is governed by dark matter and dark energy and we associate the large scale inhomogeneities in these two components as those generated by the primordial cosmological perturbations with an effective matter density contrast δρ/ρ≤1$\delta \rho /\rho \le 1$. The small scale inhomogeneities are generated by the condensations of baryonic matter considered as the bare perturbations of the background manifold that admits δρ/ρ?1$\delta \rho /\rho ?1$. Mathematically, the large scale perturbations are given by the homogeneous solution of the linearized field equations while the small scale perturbations are described by a particular solution of these equations with the bare stress–energy tensor of the baryonic matter. We explicitly work out the covariant field equations of the successive post-Friedmannian approximations of Einstein’s equations in cosmology and derive equations of motion of large and small scale inhomogeneities of dark matter and dark energy. We apply these equations to derive the post-Friedmannian equations of motion of baryonic matter comprising stars, galaxies and their clusters.     

Accurate eikonal-curvature relation for wave fronts in locally anisotropic reaction-diffusion systems

The dependency of wave velocity in reaction-diffusion (RD) systems on the local front curvature determines not only the stability of wave propagation, but also the fundamental properties of other spatial configurations such as vortices. This Letter gives the first derivation of a covariant eikonal-curvature relation applicable to general RD systems with spatially varying anisotropic diffusion properties, such as cardiac tissue. The theoretical prediction that waves which seem planar can nevertheless possess a nonvanishing geometrical curvature induced by local anisotropy is confirmed by numerical simulations, which reveal deviations up to 20% from the nominal plane wave speed.     

Organizing Filament of Small Amplitude Scroll Waves

We theoretically analyze the organizing filament of small amplitude scroll waves in general excitable media by perturbation method and explicitly give the expressions of coefficients in Keener theory.In particular for the excitable media with equal diffusion,we obtain a close system for the motion of the filament.With an example of the Oregonator Model,our results are in good agreement with those simulated by Windree.     

Cookbook asymptotics for spiral and scroll waves in excitable media

Algebraic formulas predicting the frequencies and shapes of waves in a reaction-diffusion model of excitable media are presented in the form of four recipes. The formulas themselves are based on a detailed asymptotic analysis (published elsewhere) of the model equations at leading order and first order in the asymptotic parameter. The importance of the first order contribution is stressed throughout, beginning with a discussion of the Fife limit, Fife scaling, and Fife regime. Recipes are given for spiral waves and detailed comparisons are presented between the asymptotic predictions and the solutions of the full reaction-diffusion equations. Recipes for twisted scroll waves with straight filaments are given and again comparisons are shown. The connection between the asymptotic results and filament dynamics is discussed, and one of the previously unknown coefficients in the theory of filament dynamics is evaluated in terms of its asymptotic expansion. (c) 2002 American Institute of Physics.     

Vortex Turbulence due to the Interplay of Filament Tension and Rotational Anisotropy

The mechanism of scroll wave turbulence is investigated in excitable media with rotational anisotropy. We adopt the Barkley model with heterogeneity in the diffusion constants. Through comparative numerical studies, we demonstrate the vortex turbulence results from the rotational anisotropy＇s cooperation with negative filament tension or competition with positive filament tension. The presence of rotational anisotropy can enlarge the parameter region leading to negative-tension induced wave turbulence in isotropic media.     

Anchoring of drifting spiral and scroll waves to impermeable inclusions in excitable media

Anchoring of spiral and scroll waves in excitable media has attracted considerable interest in the context of cardiac arrhythmias. Here, by bombarding inclusions with drifting spiral and scroll waves, we explore the forces exerted by inclusions onto an approaching spiral and derive the equations of motion governing spiral dynamics in the vicinity of inclusion. We demonstrate that these forces nonmonotonically depend on distance and can lead to complex behavior: (a)?anchoring to small but circumnavigating larger inclusions; (b)?chirality-dependent anchoring.     

The Quantum Regularization of Singular Black-Hole Solutions in Covariant Quantum Gravity

An excruciating issue that arises in mathematical, theoretical and astro-physics concerns the possibility of regularizing classical singular black hole solutions of general relativity by means of quantum theory. The problem is posed here in the context of a manifestly covariant approach to quantum gravity. Provided a non-vanishing quantum cosmological constant is present, here it is proved how a regular background space-time metric tensor can be obtained starting from a singular one. This is obtained by constructing suitable scale-transformed and conformal solutions for the metric tensor in which the conformal scale form factor is determined uniquely by the quantum Hamilton equations underlying the quantum gravitational field dynamics.     

Vortex dynamics in three-dimensional continuous myocardium with fiber rotation: Filament instability and fibrillation

Wave propagation in ventricular muscle is rendered highly anisotropic by the intramural rotation of the fiber. This rotational anisotropy is especially important because it can produce a twist of electrical vortices, which measures the rate of rotation (in degree/mm) of activation wavefronts in successive planes perpendicular to a line of phase singularity, or filament. This twist can then significantly alter the dynamics of the filament. This paper explores this dynamics via numerical simulation. After a review of the literature, we present modeling tools that include: (i) a simplified ionic model with three membrane currents that approximates well the restitution properties and spiral wave behavior of more complex ionic models of cardiac action potential (Beeler-Reuter and others), and (ii) a semi-implicit algorithm for the fast solution of monodomain cable equations with rotational anisotropy. We then discuss selected results of a simulation study of vortex dynamics in a parallelepipedal slab of ventricular muscle of varying wall thickness (S) and fiber rotation rate (theta(z)). The main finding is that rotational anisotropy generates a sufficiently large twist to destabilize a single transmural filament and cause a transition to a wave turbulent state characterized by a high density of chaotically moving filaments. This instability is manifested by the propagation of localized disturbances along the filament and has no previously known analog in isotropic excitable media. These disturbances correspond to highly twisted and distorted regions of filament, or "twistons," that create vortex rings when colliding with the natural boundaries of the ventricle. Moreover, when sufficiently twisted, these rings expand and create additional filaments by further colliding with boundaries. This instability mechanism is distinct from the commonly invoked patchy failure or wave breakup that is not observed here during the initial instability. For modified Beeler-Reuter-like kinetics with stable reentry in two dimensions, decay into turbulence occurs in the left ventricle in about one second above a critical wall thickness in the range of 4-6 mm that matches experiment. However this decay is suppressed by uniformly decreasing excitability. Specific experiments to test these results, and a method to characterize the filament density during fibrillation are discussed. Results are contrasted with other mechanisms of fibrillation and future prospects are summarized. (c)1998 American Institute of Physics.     

Filament instability and rotational tissue anisotropy: A numerical study using detailed cardiac models

The role of cardiac tissue anisotropy in the breakup of vortex filaments is studied using two detailed cardiac models. In the Beeler-Reuter model, modified to produce stable spiral waves in two dimensions, we find that anisotropy can destabilize a vortex filament in a parallelepipedal slab of tissue. The mechanisms of the instability are similar to the ones reported in previous work on a simplified cardiac model by Fenton and Karma [Chaos 8, 20 (1998)]. In the Luo-Rudy model, also modified to produce stable spiral waves in two dimensions, we find that anisotropy does not destabilize filaments. A possible explanation for this model-dependent behavior based on spiral tip trajectories is offered. (c) 2001 American Institute of Physics.