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Biophys J, July 2000, p. 328-339, Vol. 79, No. 1
and
§Department of Physics, University of British
Columbia, Vancouver, British Columbia V6T 1Z1, Canada; *Department of
Pathology, University of British Columbia, Vancouver, British Columbia
V6T 1W5, Canada; Department of Cell Biology, Duke
University School of Medicine, Durham, North Carolina 27710 USA; and
Department of Mechanical Engineering and Materials
Science, Duke University, Durham, North Carolina 27708 USA
Micropipette pressurization of giant bilayer vesicles was
used to measure both elastic bending kc and
area stretch KA moduli of fluid-phase
phosphatidylcholine (PC) membranes. Twelve diacyl PCs were chosen:
eight with two 18 carbon chains and degrees of unsaturation from one
double bond (C18:1/0, C18:0/1) to six double bonds per lipid (diC18:3),
two with short saturated carbon chains (diC13:0, diC14:0), and two with
long unsaturated carbon chains (diC20:4, diC22:1). Bending moduli were
derived from measurements of apparent expansion in vesicle surface area
under very low tensions (0.001-0.5 mN/m), which is dominated by
smoothing of thermal bending undulations. Area stretch moduli were
obtained from measurements of vesicle surface expansion under high
tensions (>0.5 mN/m), which involve an increase in area per molecule
and a small
but importantcontribution from smoothing of residual
thermal undulations. The direct stretch moduli varied little (< ±10%) with either chain unsaturation or length about a mean of 243 mN/m. On the other hand, the bending moduli of
saturated/monounsaturated chain PCs increased progressively with chain
length from 0.56 × 10
19 J for diC13:0 to 1.2 × 1019 J for diC22:1. However, quite unexpectedly for
longer chains, the bending moduli dropped precipitously to ~0.4 × 1019 J when two or more cis double
bonds were present in a chain (C18:0/2, diC18:2, diC18:3, diC20:4).
Given nearly constant area stretch moduli, the variations in bending
rigidity with chain length and polyunsaturation implied significant
variations in thickness. To test this hypothesis, peak-to-peak
headgroup thicknesses hpp of bilayers were
obtained from x-ray diffraction of multibilayer arrays at controlled
relative humidities. For saturated/monounsaturated chain bilayers, the
distances hpp increased smoothly from
diC13:0 to diC22:1 as expected. Moreover, the distances and elastic
properties correlated well with a polymer brush model of the bilayer
that specifies that the elastic ratio
(kc/KA)1/2 = (hpp 
ho)/24, where
ho 
1 nm accounts for separation of
the headgroup peaks from the deformable hydrocarbon region. However, the elastic ratios and thicknesses for diC18:2, diC18:3, and diC20:4 fell into a distinct group below the correlation, which showed that
poly-cis unsaturated chain bilayers are thinner and more flexible than saturated/monounsaturated chain bilayers.
Biophys J, July 2000, p. 328-339, Vol. 79, No. 1
© 2000 by the Biophysical Society 0006-3495/00/07/328/12 $2.00
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