Self-organized dynamics and emergent shape spaces of active isotropic fluid surfaces

Abstract

Theories of self-organized active fluid surfaces have emerged as an important class of minimal models for the shape dynamics of biological membranes, cells and tissues. However, due to their inherent geometric nonlinearities and the absence of general minimization principles in active systems, it remains a major challenge to systematically study the emergent shape spaces such theories give rise to. Here, we introduce a novel variational approach that allows for a direct computation of stationary surface geometries and flows, which enables the classification of non-equilibrium phase transitions in shape spaces described by active surface theories. To achieve this, we construct a dissipation functional systematically from the entropy production in active surfaces and show how generic symmetries imposed by Onsager relations can be exploited to also account for reactive non-dissipative terms in constitutive laws. This functional is supplemented by Lagrange multipliers that relax nonlinear geometric constraints, which leads to a tractable variational problem suitable for implicit dynamic simulations and explicit calculations of non-trivial steady state geometries and flows. We apply this framework to study the dynamics of open fluid membranes and closed active fluid surfaces, and characterize the space of stationary solutions that corresponding surfaces and flows occupy. These analyses rationalize the interplay of first-order shape transitions of internally and externally forced fluid membranes, reveal degenerate regions in stationary shape spaces of mechanochemically active surfaces and identify a mechanism by which hydrodynamic screening controls the geometry of active surfaces undergoing cell division-like shape transformations.