Topological constraints on spiral wave dynamics in spherical geometries with inhomogeneous excitability.

We analyze the way topological constraints and inhomogeneity in the excitability influence the dynamics of spiral waves on spheres and punctured spheres of excitable media. We generalize the definition of an index such that it characterizes not only each spiral but also each hole in punctured, oriented, compact, two-dimensional differentiable manifolds and show that the sum of the indices is conserved and zero. We also show that heterogeneity and geometry are responsible for the formation of various spiral-wave attractors, in particular pairs of spirals in which one spiral acts as a source and a second as a sink--the latter similar to an antispiral. The results provide a basis for the analysis of the propagation of waves in heterogeneous excitable media in physical and biological systems.