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Postharvest Biology and Technology of Fruits, Vegetables, and Flowers

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450 POSTHARVEST BIOLOGY & TECHNOLOGY OF FRUITS, VEGETABLES, & FLOWERS<br />

Different anthocyanidins give different colors to fruits (Jaakola et al., 2002; Paliyath <strong>and</strong><br />

Murr, 2006).<br />

In red <strong>and</strong> nonred apples, differential expression <strong>of</strong> anthocyanin biosynthetic genes<br />

was observed. The expression <strong>of</strong> CHS, F3H, DFR, <strong>and</strong> ANS genes was found in red<br />

apples when anthocyanin was not detected (before ripening stages). However, UFGT gene<br />

expression was observed only during the ripening stages with the formation <strong>of</strong> anthocyanins.<br />

This suggests that UFGT gene expression regulates the production <strong>of</strong> anthocyanins during<br />

ripening in apples (Kondo <strong>and</strong> Hiraoka, 2002). Environmental factors such as light <strong>and</strong><br />

temperature affect anthocyanin accumulation in fruits. No anthocyanin was found in apples<br />

in the absence <strong>of</strong> light (Proctor, 1974).<br />

21.4.5 Interrelationships <strong>of</strong> metabolic pathways<br />

In fruits, quality-attributing pathways are interrelated. During starch degradation, glucose-1-<br />

phosphate is generated, which enters into several metabolic pathways such as glycolysis <strong>and</strong><br />

PPP. Glycolysis converts glucose to acetyl CoA after a series <strong>of</strong> reactions. The acetyl CoA<br />

is a precursor for the synthesis <strong>of</strong> fatty acids, volatile esters, isoprenoids, <strong>and</strong> organic acids<br />

as show in Fig. 21.3. The PPP is an important pathway, which provides carbon skeletons to<br />

several biosynthetic pathways including amino acids, secondary metabolites, <strong>and</strong> nucleic<br />

acids. Besides providing carbon skeleton, it also gives reducing energy (NADPH) to various<br />

pathways (Fig. 21.3). The carbon skeletons for the synthesis <strong>of</strong> anthocyanins come from<br />

p-coumaroyl-CoA <strong>and</strong> malonyl-CoA. The phenylpropanoid pathway donates p-coumaroyl-<br />

CoA, which derives from erythrose-4-phosphate <strong>and</strong> pyruvate that are generated during PPP.<br />

Therefore, PPP plays a significant role in the fruit quality regulation.<br />

In flavonoid biosynthesis, the hydroxylation <strong>of</strong> flavonoids by F3H requires NADPH,<br />

which comes from PPP. Isoprenoid synthesis also needs NADPH from PPP (Fig. 21.3).<br />

Moreover, it is an important component for the antioxidant enzyme system. The enzymes<br />

<strong>of</strong> ascorbate–glutathione cycle, glutathione reductase (GR), <strong>and</strong> monodehydroascorbate reductase<br />

(MDHAR), require NADPH to inactivate reactive oxygen species (ROS) generated<br />

during stress <strong>and</strong> senescence (Fig. 21.3). In order to maintain fruit quality <strong>and</strong> shelf life, the<br />

precise functioning <strong>of</strong> the antioxidant enzyme system is necessary.<br />

During membrane lipid catabolism, several fatty acid intermediates are formed that are<br />

required for the synthesis <strong>of</strong> aroma volatile components. The precursors <strong>of</strong> volatile esters,<br />

including hexanal, hexanol, <strong>and</strong> short-chain alcohols, are also derived from metabolism<br />

<strong>of</strong> lipids. The effect <strong>of</strong> hexanal has been investigated on PLD activity <strong>of</strong> membrane, <strong>and</strong><br />

soluble fractions isolated from corn kernel (Paliyath et al., 1999). Inclusion <strong>of</strong> hexanal in<br />

assay mixture resulted in over 75% inhibition <strong>of</strong> PLD activity. Other technologies have<br />

also been used for the inhibition <strong>of</strong> PLD activity. A naturally occurring lipid, lysophosphatidylethanolamine<br />

(LPE), showed potent inhibition <strong>of</strong> PLD activity in leaves, flowers,<br />

<strong>and</strong> postharvest fruits. Ryu et al. (1997) noticed a reduction in PLD activity by increasing<br />

the length <strong>and</strong> unsaturation <strong>of</strong> LPE acyl chain. In addition, decreased ethylene production<br />

was found in LPE-treated fruits. Antisense PLD tomatoes showed PLD inhibition. Furthermore,<br />

they also possessed higher levels <strong>of</strong> lycopene, firmness, <strong>and</strong> soluble solids in<br />

the fruits (Pinhero et al., 2003), suggesting that by inhibiting certain metabolic pathways,<br />

quality components <strong>and</strong> shelf life <strong>of</strong> fruits can be increased.

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