The influence of cholesterol on phospholipid membrane curvature and bending elasticity

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 (H-II) 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 H-II phases of very large dimension, Increasing levels of cholesterol result in a systematic decrease in the H-II 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 H-II monolayers, The radii of the intrinsic curvature, R-p(o), at a pivotal plane of constant area within the monolayer were determined to be 29.4 Angstrom for DOPE/tetradecane at 22 degrees C, decreasing to 27 Angstrom at 30 mol% cholesterol. For DOPC/tetradecane at 32 degrees C, R-p(o) decreased from 62.5 Angstrom to 40 Angstrom 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 Angstrom. The bending moduli k(c) 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 k(c) 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.