Sorting of proteins with shape and curvature anisotropy on a lipid bilayer tube.

Curvature induced sorting of lipid membrane bound proteins has been widely studied through experiments that induce curvature variation in a giant unilamellar lipid-bilayer vesicle with adsorbed proteins by pulling thin cylindrical tethers. In the theoretical space, this has been supplemented with models that capture curvature dependent interaction between membrane and idealized protein particles, through free energy contributions. Many membrane proteins such as the BAR domain proteins are known to have extremely anisotropic shapes and soft interacting potentials, whereas the idealizations of protein particles explored in models have only assumed them as hard disk-like particles with curvature anisotropy. Here, we present a model of sorting of the proteins while including the effects of softness in their interaction potentials, shape anisotropy in the protein structure, and curvature anisotropy in the interactions with the membrane. This is based on a clean separation of free energy contributions from non-ideal fluid behavior of soft anisotropic particles and curvature interactions between proteins and membranes. We probe the behavior of the sorting function under limiting conditions and show that it converges to the previously derived models. In addition to this, we present a comparison of the variation in sorting ratio due to the observed variation in the shape parameter values in known membrane proteins. Finally, using published experimental data for membrane proteins, we perform fitting and derive model parameters. We observe that shape anisotropy adversely affects the sorting of proteins to a high curvature region, whereas curvature anisotropy and softer interaction between proteins favor sorting.