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Food Lipids: Chemistry, Nutrition, and Biotechnology

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olive (Olea europea), <strong>and</strong> avocado (Persea americana)] <strong>and</strong> seed lipids account for<br />

over 80% of world production. In the United States, plant oils account for over 94%<br />

of the annual dem<strong>and</strong> of edible oils (8). Furthermore, the plant sources included in<br />

Table 1 are useful because of their suitability for organized agriculture (5) <strong>and</strong> acceptible<br />

yield of biomass (<strong>and</strong>, therefore, lipid). Less than 0.1% of about 300,000<br />

known plant species are cultivated in organized agriculture, <strong>and</strong> fewer than 20 species<br />

account for the principal lipid products of commerce (5). However, the great diversity<br />

among the plant kingdom offers potential for exploitation in view of the current<br />

ability to genetically transfer <strong>and</strong> manipulate specific plant traits. For example, a<br />

species that produces an unusual, precious, or otherwise noteworthy fatty acid or<br />

assembled lipid but is not suitable for commercial cultivation can serve as germ<br />

plasm for incorporating these valuable traits into species that are subject to commercial<br />

culture. Thus, in view of the relative contribution to commerce <strong>and</strong> exploitable<br />

diversity of plant lipids, it seems appropriate to restrict this chapter to a discussion<br />

of storage lipid biosynthesis in plants. Reviews on fatty acid <strong>and</strong> glycerolipid<br />

biosynthesis in animal <strong>and</strong> microbial systems are available elsewhere (9–12).<br />

II. LIPID DIVERSITY AMONG PLANTS<br />

As diverse as lipids are in plants (<strong>and</strong> other living organisms), they can be easily<br />

classified into two categories based on their physiological role. Functional lipids are<br />

those involved in vital <strong>and</strong> metabolic processes. Typically, these are isoprenyl-derived<br />

lipids (e.g., sterols, carotenoids), <strong>and</strong> polar glycerolipids (e.g., galactosyl- <strong>and</strong> phospholipids),<br />

components that comprise, <strong>and</strong> contribute to the functioning of,<br />

(sub)cellular membranes (10,13). In view of the role of functional lipids in membranes,<br />

their fatty acid composition in plants is essentially restricted to palmitic<br />

(16:0),* palmitoleic (16:1�9), trans-�3-hexadecenoic (16:1�3t), hexadecatrienoic<br />

(16:3�7,10,13), stearic (18:0), oleic (18:1�9), linoleic (18:2�9,12), <strong>and</strong> linolenic<br />

(18:3�9,12,15) acids (10,13,14). Functional lipids are not discussed in this chapter, except<br />

as they relate to the biosynthesis of fatty acids <strong>and</strong> the assembly of triacylglycerols<br />

(Sec. V). (Some functional lipids, such as soybean lecithin, are products of<br />

commerce, but many products of these types are simply by-products of the edible<br />

oil industry <strong>and</strong> would probably not alone justify the use of the host plant tissue in<br />

commerce.) The interested reader is encouraged to consult reviews on biosynthesis<br />

of polar lipids (15,16) <strong>and</strong> isoprenyl lipids, including sterols, carotenoids, tocopherol,<br />

<strong>and</strong> lower isoprenoids (17–19).<br />

Storage lipids are deposited as oil bodies in fruit or seeds <strong>and</strong> are intended to<br />

serve as a carbon (energy) source for growth during germination of seeds. In fruit<br />

tissues, the raison d’être of the copious quantities of oil may be to attract predators<br />

<strong>and</strong> facilitate seed dispersal. Invariably, storage lipids predominantly take the form<br />

of triacylglycerols (20), <strong>and</strong> their fatty acyl groups are first mobilized for catabolic<br />

processes by endogenous lipases (21) <strong>and</strong>, ultimately converted into metabolic carbon<br />

(acetate) via � oxidation in peroxisomes (22). There is a much greater diversity<br />

in fatty acid composition (range of 2–24 acyl carbons) of storage triacylglycerols<br />

*Fatty acid (or acyl group) identification is formalized throughout the chapter as X:Y, where X is the<br />

acyl carbon number <strong>and</strong> Y is the number of double bonds, all of which are cis unless otherwise noted.<br />

Subscript � i identifies position of the double bond, counting from the carboxy terminus.<br />

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

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