Lipid polymorphism of model and cellular membranes as revealed by electron microscopy

This review surveys the current state of knowledge relating to lipid polymorphism within both model lipid membrane and cellular membrane systems. Emphasis is placed upon the contribution of data obtained by transmission electron microscopy of freeze-fractured specimens. Some consideration is also given to the other important methods for the study of lipid polymorphism, namely X-ray diffraction and NMR spectroscopy. A detailed account of the different phases found in lipid mixtures within model membranes (bilayer, cubic or tetragonal, tubular or hexagonal) provides the background to the understanding of the factors involved in polymorphic phase transitions. The sequential steps involved in lipid polymorphism are defined from electron microscopical data and are related to the structural changes that can be detected within biological membranes. It is proposed that the fine structural changes detected at the initial stages of polymorphic transition in vivo may be highly relevant in relation to membrane fusion events, to the formation of tight junctions, and even to physiological transport processes. Since the later stages of polymorphic transition generally destroy the permeability barrier of model and cellular membranes, extensive rather than localized phase transition of the lipid bilayer is not at the moment considered to be compatible with cellular viability.