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Postharvest Biology and Technology of Fruits, Vegetables, and Flowers

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BIOCHEMISTRY OF FRUITS 41<br />

3.3.3 Proteolysis <strong>and</strong> structure breakdown in chloroplasts<br />

The major proteinaceous compartments in fruits are the chloroplasts that are distributed in<br />

the epidermal <strong>and</strong> hypodermal layers <strong>of</strong> fruits. The chloroplasts are not very abundant in<br />

fruits. During senescence, the chloroplast structure is gradually disassembled with a decline<br />

in chlorophyll levels due to degradation <strong>and</strong> disorganization <strong>of</strong> the grana lamellar stacks<br />

<strong>of</strong> the chloroplast. With the disorganization <strong>of</strong> the thylakoid, globular structures termed<br />

as plastoglobuli accumulate within the chloroplast stroma <strong>and</strong> are rich in degraded lipids.<br />

The degradation <strong>of</strong> chloroplasts <strong>and</strong> chlorophyll results in the unmasking <strong>of</strong> other colored<br />

pigments <strong>and</strong> is a prelude to the state <strong>of</strong> ripening <strong>and</strong> development <strong>of</strong> organoleptic qualities.<br />

Mitochondria, which are also rich in protein, are relatively stable <strong>and</strong> undergo disassembly<br />

during the latter part <strong>of</strong> ripening <strong>and</strong> senescence.<br />

Chlorophyll degradation is initiated by the enzyme chlorophyllase that splits chlorophyll<br />

into chlorophyllide <strong>and</strong> the phytol chain. Phytol chain is made up <strong>of</strong> isoprenoid units<br />

(methyl-1,3-butadiene) <strong>and</strong> its degradation products accumulate in the plastoglobuli. Flavor<br />

components such as 6-methyl-5-heptene-2-one, a characteristic component <strong>of</strong> tomato<br />

flavor, are also produced by the catabolism <strong>of</strong> phytol chain. The removal <strong>of</strong> magnesium<br />

from chlorophyllide results in the formation <strong>of</strong> pheophorbide. Pheophorbide, which possesses<br />

a tetrapyrole structure, is converted to a straight-chain colorless tetrapyrrole by the<br />

action <strong>of</strong> pheophorbide oxidase. Action <strong>of</strong> several other enzymes is necessary for the full<br />

catabolism <strong>of</strong> chlorophyll. The protein complexes that organize the chlorophyll, the lightharvesting<br />

complexes, are degraded by the action <strong>of</strong> several proteases. The enzyme ribulosebis-phosphate<br />

carboxylase/oxygenase (Rubisco), the key enzyme in photosynthetic carbon<br />

fixation, is the most abundant protein in chloroplast. Rubisco levels also decline during<br />

ripening/senescence due to proteolysis. The amino acids resulting from the catabolism <strong>of</strong><br />

proteins may be translocated to regions where they are needed for biosynthesis. In fruits,<br />

they may just enrich the soluble fraction with amino acids.<br />

3.4 Secondary plant products<br />

Secondary plant products are regarded as metabolites that are derived from primary<br />

metabolic intermediates through well-defined biosynthetic pathways. The importance <strong>of</strong><br />

the secondary plant products to the plant or organ in question may not readily be obvious,<br />

but these compounds appear to have a role in the interaction <strong>of</strong> the plant with the<br />

environment. The secondary plant products may include nonprotein amino acids, alkaloids,<br />

isoprenoid components (terpenes, carotenoids, etc.), flavonoids <strong>and</strong> anthocyanins,<br />

ester volatiles, <strong>and</strong> several other organic compounds with diverse structure. The number<br />

<strong>and</strong> types <strong>of</strong> secondary plant products are enormous, but, with the perspective <strong>of</strong> fruit quality,<br />

the important secondary plant products include isoprenoids, anthocyanins, <strong>and</strong> ester<br />

volatiles.<br />

3.4.1 Isoprenoid biosynthesis<br />

In general, isoprenoids possess a basic five-carbon skeleton in the form <strong>of</strong> 2-methyl-1,<br />

3-butadiene (isoprene), which undergoes condensation to form larger molecules. There are<br />

two distinct pathways for the formation <strong>of</strong> isoprenoids: the acetate/mevalonate pathway

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