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Postharvest Biology and Technology of Fruits, Vegetables, and Flowers

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

endotransglycosylase (XTH, EC 2.4.1.207), the endo-β-1,4-glucanases (EGases, EC<br />

3.2.1.4; previously known as cellulases), endo-β-mannanase (EC 3.2.1.78) (Bewley et al.,<br />

2000), β-D-xylosidases (EC 3.2.1.37) (Itai et al., 2003), <strong>and</strong> expansins. Mannan transglycosylases<br />

regulated during ripening are very recent additions to the list <strong>of</strong> c<strong>and</strong>idates for roles<br />

in glycan network disassembly (Schroder et al., 2004). Ripening-associated modifications<br />

in hemicellulose-cellulose structure appear to greatly influence the accessibility <strong>of</strong> multiple<br />

enzymes to their substrates. Expansins target components <strong>of</strong> the glycan superstructure<br />

in ripening fruits, indirectly facilitating hydrolysis <strong>of</strong> the HGA substrate by one or more<br />

pectinase (Brummell et al., 1999).<br />

Efficient distribution <strong>of</strong> C-14-starch into glucose, fructose, <strong>and</strong> sucrose revealed considerable<br />

sugar interconversions indicating active gluconeogenesis during mango fruit ripening<br />

(Yashoda et al., 2006). But the texture <strong>of</strong> Kent mango is most likely moderated by changes in<br />

solubility <strong>of</strong> insoluble pectin or by nonpectin components in the cell wall as the temperature<br />

gradient infusion <strong>of</strong> fresh-cut mangoes with PME <strong>and</strong>/or calcium chloride had no impact<br />

on hardness <strong>and</strong> adhesiveness (Banjongsinsiri et al., 2004).<br />

Biochemical changes during ripening <strong>of</strong> capsicum are characterized as an increase in<br />

free sugar levels, an increase in in situ hydrolysis <strong>of</strong> some hemicellulose fractions (Hem a,<br />

b, <strong>and</strong> c), <strong>and</strong> a general increase in the activity <strong>of</strong> cellulase, α-mannosidase, laminarinase,<br />

polygalacturanase, galactanase, mannanase, β-galactosidase, <strong>and</strong> hemicellulase (HCe) activity<br />

on Hem b <strong>and</strong> c. But the activity <strong>of</strong> xylanase, PME, <strong>and</strong> HCe on Hem a decreased<br />

during ripening (Prabha et al., 1998). Jagadeesh et al. (2004a) observed ripening specificity<br />

for β-hexosaminidase <strong>and</strong> an interrelationship between this <strong>and</strong> α-mannosidase activities,<br />

which may be responsible for the textural s<strong>of</strong>tening associated with capsicum fruit<br />

ripening.<br />

The amount <strong>of</strong> cellulose does not decline during ripening in most fruits (Maclachlan<br />

<strong>and</strong> Brady, 1994; Sakurai <strong>and</strong> Nevins, 1997), <strong>and</strong> it is presumed that depolymerization <strong>of</strong><br />

cellulose is not a major feature <strong>of</strong> ripening. In avocado, a loss <strong>of</strong> fibrillar material from the<br />

wall occurred during ripening that was due to the depolymerization <strong>of</strong> large unbranched<br />

molecules <strong>of</strong> cellulose <strong>and</strong> an increase in the proportion <strong>of</strong> crystalline cellulose, suggesting<br />

a preferential degradation <strong>of</strong> peripheral amorphous cellulose chains (Platt-Aloia et al., 1980;<br />

O’Donoghue et al., 1994).<br />

During peach fruit development <strong>and</strong> ripening, cell wall undergoes several structural<br />

<strong>and</strong> biochemical changes driven by several hydrolases (Bonghi et al., 1998). Among these,<br />

the endo-β-1,4-β-glucanase (EGase), or cellulase, may play a crucial role in cell wall<br />

hydrolysis. During the four stages <strong>of</strong> peach growth, EGase activity was high during S1 <strong>and</strong><br />

early S2, declined during S3, <strong>and</strong> increased with the onset <strong>of</strong> ripening (S4), implying that<br />

the EGases is involved in early fruit growth <strong>and</strong> the initial phases <strong>of</strong> s<strong>of</strong>tening. The presence<br />

<strong>of</strong> two is<strong>of</strong>orms <strong>and</strong> the dual effect <strong>of</strong> propylene on enzyme activity suggest that different<br />

EGase genes operate during the early <strong>and</strong> late developmental stages in peach.<br />

Detectable endo-β-1,4-glucanases (EGases) enzyme activity is first observed in large<br />

green fruits <strong>of</strong> strawberry, but a steep increase occurs in white fruits when the ripening<br />

process starts (Trainotti et al., 1999). This process is then accompanied by a further increase<br />

in EGase activity, which appears to be doubled in red ripe fruits.<br />

An increase in cellulase (endo-1,4-β-D-glucanase) accompanies progressive s<strong>of</strong>tening,<br />

loss <strong>of</strong> skin strength, <strong>and</strong> a breakdown <strong>of</strong> cell walls in the mesocarp <strong>of</strong> raspberry, indicating<br />

its involvement in fruit separation as well as s<strong>of</strong>tening (Sexton et al., 1997). The initial

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