New aspects on membrane lipid regulation in Acholeplasma laidlawii A and phase equilibria of monoacyldiglucosyldiacylglycerol

A new membrane lipid, monoacyldiglucosyldiacylglycerol (MADGlcDAG), was recently discovered in Acholeplasma laidlawii strain A-EF22 demanding a new study of the biosynthetic regulation, and the phase behavior, of the glucolipids in this organism. The only liquid-crystalline phase formed by MADGlcDAG is a reversed hexagonal phase, A. laidlawii A-EF22 synthesizes four lipids that have the ability to induce the formation of reversed nonlamellar phases: MADGlcDAG, monoglucosyldiacylglycerol (MGlcDAG), monoacylmonoglucosyldiacylglycerol (MAMGlcDAG), and diacylglycerol (DAG), A C-n value of approximately 16 seems to be a critical value for the fractions of these lipids in the membrane: the fractions of MADGlcDAG and MGlcDAG are largest when the C-n values are lower than 16, while the fractions of MAMGlcDAG and DAG are largest when the C-n values are higher than 16, The fraction of nonlamellar-forming liqids was 55 mol % when the C-n value was 14.8 and the degree of unsaturation was 33 mol %, This fraction was reduced to 7 mol % when the C-n value and the degree of unsaturation were increased to 17.8 and 92 mol %, respectively, i.e., at conditions that markedly favor the formation of reversed nonlamellar phases. These observations convincingly show that a balance between lamellar- and nonlamellar-forming lipids is maintained in the membrane and strongly support the validity of the lipid regulation model proposed by us. From earlier biochemical data, obtained with short acyl chains, that were difficult to reconcile with our regulation model, it could be predicted that a lipid ought to be synthesized that assists MGlcDAG to maintain the nonlamellar-forming properties with the short chains. It is shown in the present work that this lipid is MADGlcDAG and that the regulation of the balance between lamellar- and nonlamellar-forming lipids is even more complex and sophisticated in A. laidlawii A-EF22 than previously proposed.