Regulation of lipid composition in biological membranes - biophysical studies of lipids and lipid synthesizing enzymes

The study of the role played by membrane lipids, functional lipidomics, has become increasingly important in membrane biology. The physico-chemical properties of the lipids in biological membranes are subject to some fundamental requirements. In general, the acyl chains shall be in a liquid-like state to keep the membrane proteins active, and the lipids must form a bilayer structure in order for the membrane to be an insulating barrier. However, a potential ability of the lipids to form nonbilayer structures seems to be a prequisite for several membrane-associated cell processes. Therefore, organisms exposed to changes in the environmental conditions, such as temperature and uptake of fatty acids, adjust their membrane lipid composition. Examples of prokaryotic organisms that have been studied in this respect are the cell wall-less bacterium Acholeplasma laidlawii, the Gram-negative bacterium Escherichia coli, and the Gram-positive bacteria Bacillus megaterium and Clostridium butyricum, and among eukaroytic organisms are found fungi, higher plants, and poikilothermic animals. By synthesizing a proper combination of acyl chain and polar head group structures, the organisms modify the phase transition temperatures of the membrane lipids so that they are maintained in a lamellar liquid crystalline phase, and the formation of a lamellar gel phase as well as reserved nonlamellar phases is avoided. It has been shown that A. laidlawii and E. coli maintain a balance between lamellar-formed and nonlamellar-forming lipids. A growing body of evidence shows that nonlamellar-forming membrane lipids play essential roles in many aspects of membrane functioning. Short-lived nonbilayer structures are probably formed in the processes of fusion and fission of lipid bilayers, and long-lived bilayer structures with a small radius of curvature occur in several types of biological membranes (e.g. smooth endoplasmic reticulum, inner mitochondrial membrane, and prolamellar bodies). The activity of membrane-associated proteins can be modulated by adding detergents or nonlamellar-forming lipids to bilayers. Some examples are the regeneration of denatured bacteriorphodopsin, and the activities of protein kinase C, some phospholipases, and some key lipid synthases involved in the lipid metabolism of eukaryotic cells, A. laidlawii, and E. coli. The physico-chemical properties of the lipid matrix can be a direct feed-back signal on the activity of the lipid synthases. Finally, nonlamellar-forming lipids are essential for a proper cell division, and an efficient translocation of proteins across the plasma membrane, of E. coli cells. (C) 2002 Elsevier Science B.V. All rights reserved.