Budding transitions of fluid-bilayer vesicles: the effect of area-difference elasticity

Abstract

Budding and vesiculation are prominent shape transformations of fluid lipid-bilayer vesicles. We discuss these transitions within the context of a curvature model which contains two types of bending energy. In addition to the usual local curvature elasticity κ, we include the effect of a relative areal stretching of the two monolayers. This area-difference elasticity leads to an effective nonlocal curvature energy characterized by another parameter κ¯. We argue that the two contributions to the curvature energy are typically comparable in magnitude. The model interpolates smoothly between the spontaneous-curvature model (κ¯=0) and the bilayer-couple model (κ¯→∞), discussed previously in the literature. Conceptually, this model is not new; however, neither its consequences nor its relation to experiment has previously been explored in detail. In particular, budding is discontinuous (first order) for small κ¯ but changes via a tricritical point to continuous (second order) for large κ¯. The order of the budding transition depends on both the ratio κ¯/κ (which is a material parameter) and the initial area difference between the inner and outer monolayers (which can be modified by appropriate treatment of the vesicle). Estimates suggest that, under typical laboratory conditions, the budding process should be discontinuous, in apparent disagreement with some recent experiments. Possible reasons for this discrepancy are discussed. We propose, in particular, that hysteretic effects are important and that the observed behavior may reflect a spinodal instability.