Mechanical properties of model membranes studied from shape transformations of giant vesicles.

Membrane deformations occur frequently in cell functioning. From the physical point of view, the understanding of such shape changes requires the introduction of mechanical parameters like bending elasticity. In this article it is shown how this physical property can be obtained from the analysis of small or large shape transformations from giant vesicles. Then it is demonstrated that the bending modulus is strongly dependent on the membrane composition and environmental conditions. This is the case for one-component bilayers (dilauroylphosphatidylcholine (DLPC), dimyristoylphosphatidylcholine (DMPC), dipalmitoylphosphatidylcholine and stearoyloleoyl-phosphatidylcholine (SOPC) and for two-component lipid mixtures (DMPC/cholesterol, DLPC/dilauroylphosphatidic acid). Further it is shown that the bending elasticity of natural lipid extracts (egg phosphatidylcholine, digalactosyl diglyceride and red blood cell lipid extracts) is generally smaller than that of comparable synthetic model membranes. The role of transmembrane proteins is examined by measuring the bending elasticity of SOPC/gramicidin mixtures. Finally, larger scale shape transformations of giant vesicles under an alternative electric field are discussed.

Bending elasticities of model membranes: influences of temperature and sterol content.

Giant liposomes obtained by electroformation and observed by phase-contrast video microscopy show spontaneous deformations originating from Brownian motion that are characterized, in the case of quasispherical vesicles, by two parameters only, the membrane tension sigma and the bending elasticity k(c). For liposomes containing dimyristoyl phosphatidylcholine (DMPC) or a 10 mol% cholesterol/DMPC mixture, the mechanical property of the membrane, k(c), is shown to be temperature dependent on approaching the main (thermotropic) phase transition temperature T(m). In the case of DMPC/cholesterol bilayers, we also obtained evidence for a relation between the bending elasticity and the corresponding temperature/cholesterol molecular ratio phase diagram. Comparison of DMPC/cholesterol with DMPC/cholesterol sulfate bilayers at 30 degrees C containing 30% sterol ratio shows that k(c) is independent of the surface charge density of the bilayer. Finally, bending elasticities of red blood cell (RBC) total lipid extracts lead to a very low k(c) at 37 degrees C if we refer to DMPC/cholesterol bilayers. At 25 degrees C, the very low bending elasticity of a cholesterol-free RBC lipid extract seems to be related to a phase coexistence, as it can be observed by solid-state (31)P-NMR. At the same temperature, the cholesterol-containing RBC lipid extract membrane shows an increase in the bending constant comparable to the one observed for a high cholesterol ratio in DMPC membranes.