Generalized measures of population synchrony.

Synchronized behavior among individuals, broadly defined, is a ubiquitous feature of populations. Understanding mechanisms of (de)synchronization demands meaningful, interpretable, computable quantifications of synchrony, relevant to measurements that can be made of complex, dynamic populations. Despite the importance to analyzing and modeling populations, existing notions of synchrony often lack rigorous definitions, may be specialized to a particular experimental system and/or measurement, or may have undesirable properties that limit their utility. Here we introduce a notion of synchrony for populations of individuals occupying a compact metric space that depends on the Fréchet variance of the distribution of individuals across the space. We establish several fundamental and desirable mathematical properties of our proposed measure of synchrony, including continuity and invariance to metric scaling. We establish a general approximation result that controls the disparity between synchrony in the true space and the synchrony observed through a discretization of state space, as may occur when observable states are limited by measurement constraints. We develop efficient algorithms to compute synchrony for distributions in a variety of state spaces, including all finite state spaces and empirical distributions on the circle, and provide accessible implementations in an open-source Python module. To demonstrate the usefulness of the synchrony measure in biological applications, we investigate several biologically relevant models of mechanisms that can alter the dynamics of population synchrony over time, and reanalyze published experimental and model data concerning the dynamics of the intraerythrocytic developmental cycles of Plasmodium parasites. We anticipate that the rigorous definition of population synchrony and the mathematical and biological results presented here will be broadly useful in analyzing and modeling populations in a variety of contexts.