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non-cylindrical lipids. Indeed, only 20
50% of the total lipids need be cylindrical to maintain
e
the normal lamellar phase.
A general description of lipid shape is not entirely satisfying as lipids actually exist at all
locations on the continuum, from truncated cone, through cylinder, to inverted cone [24] .
Therefore attempts have been made to quantify lipid shape. Two of these approaches are dis-
cussed below.
The first approach, proposed by Pieter Cullis in 1979 [19] defines lipid shape by a dimen-
sionless parameter S:
¼ v = a o l c
where a o is the optimum cross-sectional area per molecule at the lipid
S
water interface, v is
the volume per molecule and I c is the length of the fully extended acyl chain. Since these
parameters are not easy to obtain, a simplified version was suggested, where a o is the
cross-sectional area of the head group at the aqueous interface and a h is the cross-sectional
area at the bottom of the acyl chains. Therefore if:
e
a o =
a h > 1
then S
> 1/
inverted cone
a o =
a h ¼ 1
then S
¼ 1/
cylindrical
cone
Unfortunately, S is not a fixed parameter of the lipid's shape, as it may vary with the
membrane's environment (i.e. bathing solution pH, ionic strength, atmospheric pressure,
lateral pressure, temperature etc.).
A second approach, the equilibrium curvature (R o ) concept, was later proposed by Sol
Gruner [25] . R o is a quantitative measure of the propensity of a lipid to form a non-lamellar
structure. It is an estimate of the tendency of a monolayer made from a particular lipid to
curl, thus resulting in hydrocarbon packing strain. R o is normally derived experimentally
a o =
a h < 1
then S
< 1/
TABLE 10.2 The Spontaneous Radius of
Curvature (R o ) for Several
Important Membrane Lipids.
Spontaneous radius of
curvature R o ( ˚ )
Lipid
DOPS
þ
150
Lyso PC
þ
38 to
þ
68
DOPE
28.5
DOPC
80 to
200
Cholesterol
22.8
a
-Tocopherol
13.7
DOG
11.5
The table was adapted from [23] .
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