Bilayers composed of
phosphatidylcholine (PC),
sphingomyelin (SM), and
cholesterol (CHOL) are commonly used as systems to model the raft-
lipid domain structure believed to compartmentalize particular
cell membrane proteins. In this work, micropipette aspiration of giant
unilamellar vesicles was used to test the elasticities, water permeabilities, and
rupture tensions of single-component PC, binary 1:1 PC/CHOL, and 1:1 SM/CHOL, and ternary 1:1:1 PC/SM/CHOL bilayers, one set of measurements with dioleoyl PC (
DOPC; C18:1/C18:1 PC) and the other with stearoyloleoyl PC (
SOPC; C18:0/C18:1 PC). Defining the elastic moduli (K(A)), the initial slopes of the increase in tension (sigma) versus stretch in
lipid surface area (alpha(e)) were determined for all systems at low (15 degrees C) and high (32-33 degrees C) temperatures. The moduli for the single-component PC and binary
phospholipid/CHOL bilayers followed a descending hierarchy of stretch resistance with SM/CHOL >
SOPC/CHOL >
DOPC/CHOL > PC. Although much more resistant to stretch than the single-component PC bilayers, the elastic response of vesicle bilayers made from the ternary
phospholipid/CHOL mixtures showed an abrupt softening (discontinuity in slope), when immediately subjected to a steady ramp of tension at the low temperature (15 degrees C). However, the discontinuities in elastic stretch resistance at low temperature vanished when the bilayers were held at approximately 1 mN/m prestress for long times before a tension ramp and when tested at the higher temperature 32-33 degrees C. The elastic moduli of single-component PC and
DOPC/CHOL bilayers changed very little with temperature, whereas the moduli of the binary
SOPC/CHOL and SM/CHOL bilayers diminished markedly with increase in temperature, as did the ternary
SOPC/SM/CHOL system. For all systems, increasing temperature increased the water permeability but decreased
rupture tension. Concomitantly, the measurements of permeability exhibited a prominent correlation with the
rupture tension across all the systems. Together, these micromechanical tests of binary and ternary
phospholipid/CHOL bilayers demonstrate that PC
hydrocarbon chain unsaturation and temperature are major determinants of the mechanical and permeation properties of membranes composed of raft microdomain-forming
lipids.