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Postharvest Biology and Technology of Fruits, Vegetables, and Flowers

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BIOCHEMISTRY OF FRUITS 31<br />

content is generally very high in ripe mangoes <strong>and</strong> can reach levels in excess <strong>of</strong> 90% <strong>of</strong><br />

the total soluble solids content. Fructose is the predominant sugar in mangoes. In contrast<br />

to the bananas, the sucrose levels increase with the advancement <strong>of</strong> ripening in mangoes,<br />

potentially due to gluconeogenesis from organic acids (Kumar <strong>and</strong> Selvaraj, 1990). As well,<br />

the levels <strong>of</strong> pentose sugars increase during ripening, <strong>and</strong> could be related to an increase in<br />

the activity <strong>of</strong> the pentose phosphate pathway.<br />

3.3.1.3 Glycolysis<br />

The conversion <strong>of</strong> starch to sugars <strong>and</strong> their subsequent metabolism occur in different compartments.<br />

During the development <strong>of</strong> fruits, photosynthetically fixed carbon is utilized for<br />

both respiration <strong>and</strong> biosynthesis. During this phase, the biosynthetic processes dominate.<br />

As the fruit matures <strong>and</strong> begin to ripen, the pattern <strong>of</strong> sugar utilization changes. Ripening<br />

is a highly energy-intensive process. And this is reflected in the burst in respiratory carbon<br />

dioxide evolution during ripening. As mentioned earlier, the respiratory burst is characteristic<br />

<strong>of</strong> some fruits that are designated as climacteric fruits. The postharvest shelf life <strong>of</strong><br />

fruits can depend on their intensity <strong>of</strong> respiration. <strong>Fruits</strong> such as mango <strong>and</strong> banana possess<br />

high level <strong>of</strong> respiratory activity <strong>and</strong> are highly perishable. The application <strong>of</strong> controlled<br />

atmosphere conditions having low oxygen levels <strong>and</strong> low temperature have thus become a<br />

routine technology for the long-term preservation <strong>of</strong> fruits.<br />

The sugars <strong>and</strong> sugar phosphates generated during the catabolism <strong>of</strong> starch are metabolized<br />

through the glycolysis <strong>and</strong> citric acid cycle (Fig. 3.4). Sugar phosphates can also<br />

be channeled through the pentose phosphate pathway, which is a major metabolic cycle<br />

that provides reducing power for biosynthetic reactions in the form <strong>of</strong> NADPH, as well as<br />

supplying carbon skeletons for the biosynthesis <strong>of</strong> several secondary plant products. The<br />

organic acids stored in the vacuole are metabolized through the functional reversal <strong>of</strong> respiratory<br />

pathway, which is termed as gluconeogenesis. Altogether, sugar metabolism is a<br />

key biochemical characteristic <strong>of</strong> the fruits.<br />

In the glycolytic steps <strong>of</strong> reactions (Fig. 3.4), glucose-6-phosphate is isomerized to<br />

fructose-6-phosphate by the enzyme hexose phosphate isomerase. Glucose-6-phosphate is<br />

derived from glucose-1-phosphate by the action <strong>of</strong> glucose phosphate mutase. Fructose-<br />

6-phosphate is phosphorylated at the C1 position yielding fructose-1,6-bisphosphate. This<br />

reaction is catalyzed by the enzyme phosph<strong>of</strong>ructokinase in the presence <strong>of</strong> ATP. Fructose-<br />

1,6-bisphosphate is further cleaved into two three-carbon intermediates, dihydroxyacetone<br />

phosphate <strong>and</strong> glyceraldehyde-3-phosphate, catalyzed by the enzyme aldolase. These<br />

two compounds are interconvertible through an isomerization reaction mediated by triose<br />

phosphate isomerase. Glyceraldehyde-3-phosphate is subsequently phosphorylated at the<br />

C1 position using orthophosphate, as well as oxidized using nicotinamide adenine dinucleotide<br />

(NAD), to generate 1,3-diphosphoglycerate <strong>and</strong> nicotinamide adenine dinucleotide<br />

plus hydrogen (NADH). In the next reaction, 1,3-diphosphoglycerate is dephosphorylated<br />

by glycerate-3-phosphate kinase in the presence <strong>of</strong> ADP, along with the formation<br />

<strong>of</strong> ATP. Glycerate-3-phosphate formed during this reaction is further isomerized to<br />

2-phosphoglycerate in the presence <strong>of</strong> phosphoglycerate mutase. In the presence <strong>of</strong> the<br />

enzyme enolase, 2-phosphoglycerate is converted to phosphoenol pyruvate (PEP). Dephosphorylation<br />

<strong>of</strong> phosphoenolpyruvate in the presence <strong>of</strong> ADP by pyruvate kinase yields<br />

pyruvate <strong>and</strong> ATP. Metabolic fate <strong>of</strong> pyruvate is highly regulated. Under normal conditions,<br />

it is converted to acetyl coA, which then enters the citric acid cycle. Under anaerobic

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