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• Articles by S. Leikin
• Articles by R.P. Rand

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• The influence of cholesterol on phospholipid membrane curvat...

  The influence of cholesterol on phospholipid membrane curvature and bending elasticity Biophysical Journal, Volume 73, Issue 1, 1 July 1997, Pages 267-276 Z. Chen and R.P. Rand Abstract The behavior of dioleoylphosphatidylethanolamine (DOPE)/cholesterol/tetradecane and dioleoylphosphatidylcholine (DOPC)/cholesterol/tetradecane were examined using x-ray diffraction and the osmotic stress method. DOPE/tetradecane, with or without cholesterol, forms inverted hexagonal (HII) phases in excess water. DOPC/tetradecane forms lamellar phases without cholesterol at lower temperatures. With tetradecane, as little as 5 mol% cholesterol in DOPC induced the formation of HII phases of very large dimension. Increasing levels of cholesterol result in a systematic decrease in the HII lattice dimension for both DOPE and DOPC in excess water. Using osmotic pressure to control hydration, we applied a recent prescription to estimate the intrinsic curvature and bending modulus of the HII monolayers. The radii of the intrinsic curvature, RPO, at a pivotal plane of constant area within the monolayer were determined to be 29.4 A for DOPE/tetradecane at 22 degrees C, decreasing to 27 A at 30 mol% cholesterol. For DOPC/tetradecane at 32 degrees C, RPO decreased from 62.5 A to 40 A as its cholesterol content increased from 30 to 50 mol%. These data yielded an estimate of the intrinsic radius of curvature for pure DOPC of 87.3 A. The bending moduli kc of DOPE/tetradecane and DOPC/tetradecane, each with 30 mol% cholesterol, are 15 and 9 kT, respectively. Tetradecane itself was shown to have little effect on the bending modulus in the cases of DOPE and cholesterol/DOPE. Surprisingly, cholesterol effected only a modest increase in the kc of these monolayers, which is much smaller than estimated from its effect on the area compressibility modulus in bilayers. We discuss possible reasons for this difference. Abstract | PDF (872 kb)

• Lipid Intermediates in Membrane Fusion: Formation, Structure...

  Lipid Intermediates in Membrane Fusion: Formation, Structure, and Decay of Hemifusion Diaphragm Biophysical Journal, Volume 83, Issue 5, 1 November 2002, Pages 2634-2651 Yonathan Kozlovsky, Leonid V. Chernomordik and Michael M. Kozlov Abstract Lipid bilayer fusion is thought to involve formation of a local hemifusion connection, referred to as a fusion stalk. The subsequent fusion stages leading to the opening of a fusion pore remain unknown. The earliest fusion pore could represent a bilayer connection between the membranes and could be formed directly from the stalk. Alternatively, fusion pore can form in a single bilayer, referred to as hemifusion diaphragm (HD), generated by stalk expansion. To analyze the plausibility of stalk expansion, we studied the pathway of hemifusion theoretically, using a recently developed elastic model. We show that the stalk has a tendency to expand into an HD for lipids with sufficiently negative spontaneous splay, <0. For different experimentally relevant membrane configurations we find two characteristic values of the spontaneous splay. and , determining HD dimension. The HD is predicted to have a finite equilibrium radius provided that the spontaneous splay is in the range <<, and to expand infinitely for <. In the case of common lipids, which do not fuse spontaneously, an HD forms only under action of an external force pulling the diaphragm rim apart. We calculate the dependence of the HD radius on this force. To address the mechanism of fusion pore formation, we analyze the distribution of the lateral tension emerging in the HD due to the establishment of lateral equilibrium between the deformed and relaxed portions of lipid monolayers. We show that this tension concentrates along the HD rim and reaches high values sufficient to rupture the bilayer and form the fusion pore. Our analysis supports the hypothesis that transition from a hemifusion to a fusion pore involves radial expansion of the stalk. Abstract | Full Text | PDF (308 kb)

• Comparative Study of the Effects of Several n-Alkanes on Pho...

  Comparative Study of the Effects of Several n-Alkanes on Phospholipid Hexagonal Phases Biophysical Journal, Volume 74, Issue 2, 1 February 1998, Pages 944-952 Z. Chen and R.P. Rand Abstract The effects of a series of normal alkanes (decane, dodecane, tetradecane, hexadecane, and octadecane) on the hexagonal H structures containing dioleoylphosphatidylethanolamine (DOPE) and dioleoylphosphatidylcholine (DOPC) were studied using x-ray diffraction and osmotic stress. The alkanes affect structural dimensions and the monolayer intrinsic curvature and bending modulus. The alkane effects are chain-length dependent and are attributed to their different distribution within the H structure. The data suggest that short-chain alkanes are more uniformly distributed within the H hydrocarbon regions and change the curvature and bending modulus of the monolayer, whereas longer-chain alkanes appear confined more to the interstitial region and do not change the curvature and bending modulus. Abstract | Full Text | PDF (222 kb)

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Abstract

Diacylglycerol, a biological membrane second messenger, is a strong perturber of phospholipid planar bilayers. It converts multibilayers to the reverse hexagonal phase (HII), composed of highly curved monolayers. We have used x-ray diffraction and osmotic stress of the HII phase to measure structural dimensions, spontaneous curvature, and bending moduli of dioleoylphosphatidylethanolamine (DOPE) monolayers doped with increasing amounts of dioleoylglycerol (DOG). The diameter of the HII phase cylinders equilibrated in excess water decreases significantly with increasing DOG content. Remarkably, however, all structural dimensions at any specific water/lipid ratio that is less than full hydration are insensitive to DOG. By plotting structural parameters of the HII phase with changing water content in a newly defined coordinate system, we show that the elastic deformation of the lipid monolayers can be described as bending around a pivotal plane of constant area. This dividing surface includes 30% of the lipid volume independent of the DOG content (polar heads and a small fraction of hydrocarbon chains). As the mole fraction of DOG increases to 0.3, the radius of spontaneous curvature defined for the pivotal surface decreases from 29 A to 19 A, and the bending modulus increases from approximately 11 to 14 (+/-0.5) kT. We derive the conversion factors and estimate the spontaneous curvatures and bending moduli for the neutral surface which, unlike the pivotal plane parameters, are intrinsic properties that apply to other deformations and geometries. The spontaneous curvature of the neutral surface differs from that of the pivotal plane by less than 10%, but the difference in the bending moduli is up to 40%. Our estimate shows that the neutral surface bending modulus is approximately 9kT and practically does not depend on the DOG content.