Membrane-induced interactions between curvature-generating protein domains: the role of area perturbation

Membrane deformation by asymmetric crescent-shaped proteins such as BAR-domains is calculated, using a mean field model that accounts for both bending and area stretch deformations. The penalties associated with membrane bending and area perturbation lead to moderately long-ranged (order 10 nm), non-monotonic, membrane-induced interactions between proteins that may prevent the formation of closely packed aggregates. As a result, BAR-domain proteins may favor the formation of an ordered array with a specific separation between domains whose spacing is set by the ratio between the bending and area stretch moduli