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.Comment: 5 pages, 6 Figures, major revisions, accepted for publication in
Phys. Rev.