Interactions of Anisotropic Inclusions on a Fluid Membrane

Abstract

Biological cells and membranes need to be properly shaped to fulfill many fundamental functions. This shaping is often aided by the aggregation of membrane-bound proteins that both sense the membrane curvature and shape it. Therefore, these protein inclusions interact with each other through the deformation of the membrane that they influence, and a key question is to understand the law that governs their interaction. Whereas the theoretical case of isotropic proteins is well understood, an important feature of many such proteins is their anisotropic interaction with the membrane. Here, we derive an interaction law for rigid circular membrane inclusions which impose an anisotropic contact angle on the surrounding membrane. We include the effects of both membrane bending and tension. Using asymptotic analysis, we identify two distinguished limits corresponding to weak anisotropy/weak tension and strong anisotropy/strong tension, respectively. The resulting laws exhibit a bistability in the equilibrium separation of inclusions. Inclusions with very weak anisotropy equilibrate with large separation, those with very strong anisotropy equilibrate with small separation, while there is a range of anisotropies for which both equilibria are stable. Our results provide a theoretical mechanism for the global aggregation of inclusions seen both in experiments and simulations.