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Food Lipids: Chemistry, Nutrition, and Biotechnology

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diffusion of three different types: lateral, rotational, <strong>and</strong> transversal [11]. Lateral<br />

diffusion of lipids in biological membranes refers to the two-dimensional translocation<br />

of the molecules in the plane of the membrane. Rotational diffusion of lipid<br />

molecules is restricted to the plane of biological membranes, whereas transverse<br />

diffusion is the out-of-plane rotation or redistribution of lipid molecules between the<br />

two leaflets of the bilayer. Transverse diffusion or ‘‘flip-flop’’ motion is very low in<br />

lipid bilayers, <strong>and</strong> specific enzymes are required to mediate the process.<br />

There are two major components of membrane fluidity. The first component is<br />

the order parameter (S), also called the structural, static, or range component of<br />

membrane fluidity. This is a measure of angular range of rotational motion, with<br />

more tightly packed chains resulting in a more ordered or less fluid bilayer. The<br />

second component of membrane fluidity is microviscosity <strong>and</strong> is the dynamic component<br />

of membrane fluidity. This component measures the rate of rotational motion<br />

<strong>and</strong> is a more accurate reflection of membrane microviscosity.<br />

There are many physical <strong>and</strong> chemical factors that regulate the fluidity properties<br />

of biological membranes, including temperature, pressure, membrane potential,<br />

fatty acid composition, protein incorporation, <strong>and</strong> Ca 2� concentration. For example,<br />

calcium influenced the structure of membranes containing acidic phospholipids by<br />

nonspecifically cross-linking the negative charges. Consequently, increasing the calcium<br />

concentration in systems induced structural rearrangements <strong>and</strong> a decrease in<br />

membrane fluidity [12]. Similarly, changes in microfluidity <strong>and</strong> lateral diffusion fluidity<br />

were exhibited when polyunsaturated fatty acids oxidized [13].<br />

Fluidity is an important property of membranes because of its role in various<br />

cellular functions. Activities of integral membrane-bound enzymes, such as Na � ,K � -<br />

ATPase, can be regulated to some extent by changes in the lipid portions of biological<br />

membranes. In turn, changes in enzyme activities tightly connected to ion transport<br />

processes could affect translocations of ions.<br />

C. Phase Transitions<br />

As is the case for triacylglycerols, phospholipids can exist in a frozen gel state or<br />

in a fluid liquid crystalline state, depending on the temperature [14] as illustrated in<br />

Fig. 3. Transitions between the gel <strong>and</strong> liquid crystalline phases can be monitored<br />

by a variety of techniques, including nuclear magnetic resonance (NMR), electron<br />

spin resonance, fluorescence, <strong>and</strong> differential scanning calorimetry (DSC). With<br />

DSC, both enthalpy <strong>and</strong> cooperativity of the transition may be determined, enthalpy<br />

being the energy required to melt the acyl chains <strong>and</strong> cooperativity reflecting the<br />

number of molecules that undergo a transition simultaneously. However, difficulties<br />

in determining membrane transitions have been attributed to entropy/enthalpy com-<br />

Figure 3 The phospholipid gel–liquid crystalline phase transition.<br />

Copyright 2002 by Marcel Dekker, Inc. All Rights Reserved.

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