BENDING STIFFNESS OF LIPID BILAYERS .1. BILAYER COUPLE OR SINGLE-LAYER BENDING

To describe the resistance of a bilayer to changes in curvature two mechanisms are distinguished which are termed bilayer couple bending and single-layer bending. In bilayer couple bending, the resistance arises from the 2-D isotropic elasticity of the two layers and their fixed distance. Single-layer bending covers the intrinsic bending stiffness of each monolayer. The two mechanisms are not independent. Even so, the distinction is useful since bilayer couple bending can relax by a slip between the layers from the local to the global fashion. Therefore, the bending stiffness of a bilayer depends on the time scale and on the extent of the deformation imposed on the membrane. Based on experimental data, it is shown by order of magnitude estimates that (a) the bending stiffness determined from thermally induced shape fluctuations of almost spherical vesicles is dominated by single-layer bending; (b) in the tether experiment on lipid vesicles and on red cells, a contribution of local bilayer couple bending can not be excluded; and (c) at the sharp corners at the leading and the trailing edge of tanktreading red cells, local bilayer couple bending appears to be important.