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Understanding the role of non-equilibrium driving in self-organization iscrucial for developing a predictive description of biological systems, yet itis impeded by their complexity. The actin cytoskeleton serves as a paradigm forhow equilibrium and non-equilibrium forces combine to give rise toself-organization. Motivated by recent experiments that show that actinfilament growth rates can tune the morphology of a growing actin bundlecrosslinked by two competing types of actin binding proteins, we construct aminimal model for such a system and show that the dynamics are subject to a setof thermodynamic constraints that relate the non-equilibrium driving, bundlemorphology, and molecular fluxes. The thermodynamic constraints reveal theimportance of correlations between these molecular fluxes, and offer a route toestimating microscopic driving forces from microscopy experiments.

A strong non-equilibrium bound for sorting of crosslinkers on growing biopolymers