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Postharvest Biology and Technology of Fruits, Vegetables, and Flowers

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THE ROLE OF POLYPHENOLS 269<br />

3'<br />

OH<br />

HO O + O-Sugar<br />

5'<br />

OH<br />

Anthocyanin<br />

Pelargonidin<br />

Cyanidin<br />

Delphinidin<br />

Peonidin<br />

Petunidin<br />

Malvidin<br />

Substituents<br />

3′-OH<br />

3′-OH, 5′-OH<br />

3′-OCH 3<br />

3′-OCH 3 , 5′-OH<br />

3′-OCH 3 , 5′- OCH 3<br />

Color<br />

Orange red<br />

Purplish red<br />

Bluish purple<br />

Rosy red<br />

Purple<br />

Pink/red<br />

Fig. 12.5 Chemical structures <strong>of</strong> selected anthocyanins <strong>of</strong> fruits. The major sugars <strong>of</strong> C-3 glycosylation include<br />

glucose, galactose, arabinose, rhamnose, <strong>and</strong> rutinose.<br />

activity <strong>of</strong> PAL, which in turn is regulated by ethylene (Blankenship <strong>and</strong> Unrath, 1988;<br />

Gomez-Cordoves et al., 1996). Exposure <strong>of</strong> fruit to UV light in combination with low-night<br />

<strong>and</strong> high-day temperatures can also regulate anthocyanin biosynthesis (Reay, 1999). For<br />

example, it was found that both UV light <strong>and</strong> advancing maturity promoted anthocyanin<br />

biosynthesis in “Jonathan” apple (Reay <strong>and</strong> Lancaster, 2001). It has been found that the<br />

concentration <strong>of</strong> anthocyanins was higher on the blush side than on the shaded side <strong>of</strong><br />

“Elstar” <strong>and</strong> “Jonagold” apples, <strong>and</strong> in fruit at the top <strong>and</strong> sides <strong>of</strong> the apple tree canopy<br />

(Awad <strong>and</strong> de Jager, 2000). Fine regulation <strong>of</strong> the PAL activity may be dependent on the<br />

extent <strong>of</strong> light exposure to the fruit (tree position) <strong>and</strong> climatic conditions during fruit<br />

maturation. It has also been demonstrated that UV light <strong>and</strong> low temperatures can induce<br />

the production <strong>of</strong> other flavonoids <strong>and</strong> phenolic acids (Lancaster et al., 2000). During<br />

cold storage <strong>of</strong> apple (“Granny Smith,” “Cr<strong>of</strong>ton,” <strong>and</strong> “Lady Williams”), flavonoid levels<br />

have been shown to remain constant (Golding et al., 2001), although some studies have<br />

reported minor fluctuations in levels. A similar study using “Jonagold” <strong>and</strong> “Elstar” cultivars<br />

found that most flavonoids <strong>and</strong> hydroxycinnamic acid derivatives were stable throughout<br />

cold or controlled atmosphere storage (Awad <strong>and</strong> de Jager, 2000). In “Delicious” <strong>and</strong><br />

“Ralls” apple fruit, flavan-3-ols <strong>and</strong> flavonols were generally stable during storage; however,<br />

an increase in anthocyanin concentration with a concomitant decrease in simple phenols<br />

has also been reported (Ju et al., 1996). Others have reported a proportional decrease in<br />

anthocyanin concentration in storage (Lin et al., 1989). It seems that anthocyanin content<br />

remained constant during storage, while the carotenoid levels increased <strong>and</strong> chlorophyll<br />

levels decreased (Reay, 1998). It has also been suggested that factors other than the level<br />

<strong>of</strong> ethylene in the tissue contribute to the de novo biosynthesis <strong>of</strong> anthocyanins, including<br />

low temperature (Arawaka, 1991), fruit maturity (Murphey <strong>and</strong> Dilley, 1988), <strong>and</strong> storage

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