Beyond a pacemaker's entrainment limit: Phase walk-through

Desynchronization occurs when a pacemaker is externally stimulated at too high a rate, beyond the entrainment limit. One may then observe phase walk-through: the pacemaker and stimulus phase difference, rather than lock to a fixed pattern, may cycle repeatedly through all values. By means of a simple but nonlinear one-variable model we describe qualitatively the properties of entrainment, loss of entrainment, and phase walk-through for rhythmically stimulated pacemakers. We obtain an expression for the cycle length (beat period) of the repetitive phase walk-through. This shows, in contrast to the simplest treatment, that in general beat frequency is not a simple linear function of the difference between the pacemaker and stimulus frequencies. Our results are illustrated in the context of a model experimental pacemaker, the rhythmically flashing firefly. We discuss how these modeling results apply in a much more general setting.