Chaotic dynamics in an ionic model of the propagated cardiac action potential

We simulate the effect of periodic stimulation on a strand of ventricular muscle by numerically integrating the one-dimensional cable equation using the Beeler-Reuter model to represent the transmembrane currents. As stimulation frequency is increased, the rhythms of synchronization [1:1----2:2----2:1----4:2---- irregular----3:1----6:2----irregular----4:1----8:2----...----1:0] are successively encountered. We show that this sequence of rhythms can be accounted for by considering the response of the strand to premature stimulation. This involves deriving a one-dimensional finite-difference equation or "map" from the response to premature stimulation, and then iterating this map to predict the response to periodic stimulation. There is good quantitative agreement between the results of iteration of the map and the results of the numerical integration of the cable equation. Calculation of the Lyapunov exponent of the map yields a positive value, indicating sensitive dependence on initial conditions ("chaos"), at stimulation frequencies where irregular rhythms are seen in the corresponding numerical cable simulations. The chaotic dynamics occurs via a previously undescribed route, following two period-doubling bifurcations. Bistability (the presence of two different synchronization rhythms at a fixed stimulation frequency) is present both in the simulations and the map. Thus, we have been able to directly reduce consideration of the dynamics of a partial differential equation (which is of infinite dimension) to that of a one-dimensional map, incidentally demonstrating that concepts from the field of non-linear dynamics--such as period-doubling bifurcations, bistability, and chaotic dynamics--can account for the phenomena seen in numerical simulations of the cable equation. Finally, we sketch out how the one-dimensional description can be extended, and point out some implications of our work for the generation of malignant ventricular arrhythmias.