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Postharvest Biology and Technology of Fruits, Vegetables, and Flowers

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ISOPRENOID BIOSYNTHESIS IN FRUITS AND VEGETABLES 285<br />

COOH<br />

CO<br />

C 3 H<br />

+<br />

CH 2 OP<br />

CH 2 OH<br />

CHO<br />

1-Deoxy-D-xylulose-<br />

5-phosphate synthase<br />

CH 2 OP<br />

CHOH<br />

CHOH<br />

C<br />

CH 3<br />

O<br />

Pyruvate<br />

GAP<br />

1-Deoxy-D-xylulose-<br />

5-phosphate<br />

DMAPP CH 2 OH<br />

+<br />

CH 2 OPP<br />

3x<br />

IPP<br />

CH 2 OPP<br />

2x<br />

GGPP<br />

Phytoene synthase<br />

C 40 H 64 phytoene<br />

C 40 H 56 lycopene<br />

C 40 H 56 β-carotene<br />

Fig. 13.2 Structure <strong>of</strong> major plastid-localized DXP/isoprenoid intermediates. Some key enzymes (bold face) <strong>and</strong><br />

intermediates involved in isoprenoid biosynthesis are abbreviated: DMAPP, dimethylallyl pyrophosphate; GAP,<br />

glyceraldehyde-3-phosphate; IPP, isopentenyl pyrophosphate.<br />

<strong>of</strong> plastid- <strong>and</strong> cytosolic-derived isoprenoid intermediates (Lichtenthaler, 1999; Rohmer,<br />

1999; Eisenreich et al., 2001). This is an important point because it means that the entire<br />

carotenoid biosynthetic process—from simple C 3 molecules to complete C 40 structures—<br />

takes place within the plastids <strong>and</strong> is regulated by a set <strong>of</strong> plastid-localized enzymes (Fig.<br />

13.3).<br />

DXP is derived from glyceraldehyde-3-phosphate (GAP) <strong>and</strong> pyruvate through a reaction<br />

catalyzed by the enzyme DXP synthase (DXS). The reaction involves the condensation<br />

<strong>of</strong> hydroxyethyl thiamine (derived by the decarboxylation <strong>of</strong> pyruvate) with the aldehyde<br />

group <strong>of</strong> GAP to form DXP (Lange et al., 1998; Rohmer, 1999). Through several subsequent<br />

enzymatic steps, DXP is converted to isopentenyl pyrophosphate (IPP) <strong>and</strong> its isomer,<br />

dimethylallyl pyrophosphate (DMAPP), both C 5 compounds serve as the basic building<br />

blocks <strong>of</strong> the isoprenoid pathway (Charon et al., 2000; Rodríguez-Concepción et al.,<br />

2000). Geranylgeranyl pyrophosphate (GGPP; C 20 ) is formed by GGPP synthase via the

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