A perspective on systems containing many action potential waves that, individually, are prone to spiral wave breakup is proposed. The perspective is based on two quantities, "predator" and "prey," which we define as the fraction of the system in the excited state and in the excitable but unexcited state, respectively. These quantities exhibited a number of properties in both simulations and fibrillating canine cardiac tissue that were found to be consistent with a proposed theory that assumes the existence of regions we call "domains of influence," each of which is associated with the activity of one action potential wave. The properties include (i) a propensity to rotate in phase space in the same sense as would be predicted by the standard Volterra-Lotka predator-prey equations, (ii) temporal behavior ranging from near periodic oscillation at a frequency close to the spiral wave rotation frequency ("type-1" behavior) to more complex oscillatory behavior whose power spectrum is composed of a range of frequencies both above and, especially, below the spiral wave rotation frequency ("type-2" behavior), and (iii) a strong positive correlation between the periods and amplitudes of the oscillations of these quantities. In particular, a rapid measure of the amplitude was found to scale consistently as the square root of the period in data taken from both simulations and optical mapping experiments. Global quantities such as predator and prey thus appear to be useful in the study of multiple spiral wave systems, facilitating the posing of new questions, which in turn may help to provide greater understanding of clinically important phenomena such as ventricular fibrillation.
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