Postharvest Biology and Technology of Fruits, Vegetables, and Flowers

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CHANGES IN NUTRITIONAL QUALITY OF FRUITS AND VEGETABLES 447 Sucrose Photosynthesis Glucose Glucose-1-phosphate Starch Glycolysis ATP Hexokinase NADP NADPH Pentose phosphate pathway ADP Glucose-6-phosphate Glucose-6-phosphate dehydrogenase 6-Phosphogluconate NADP NADPH 6-Phosphogluconate dehydrogenase Glyceraldehyde-3-phosphate CO 2 Ribulose-5-phosphate NADPH pool Pyruvate dehydrogenase Pyruvate NAD NADH Isoprenoid pathway Glyceraldehyde-3- phosphate Sedoheptulose-7- phosphate Acetyl CoA Transaldolase Citric acid cycle Fructose-6-phosphate Erythrose-4-phosphate Phenyl propanoid pathway Fig. 21.1 Carbohydrate metabolism in fruits. Starch is catabolized to sugars that undergo glycolysis and pentose phosphate pathway. Pentose phosphate pathway provides carbon and reducing power (NADPH) for the synthesis of flavonoids through phenyl propanoid pathway. 21.4.2 Biosynthesis of isoprenoids Isoprenoids are secondary plant products that are derived from primary metabolites. Although there are several kinds of secondary plant products, the levels of isoprenoids, ester volatiles, and anthocyanin components determine the fruit quality to a large extent. Isoprenoids are synthesized from two pathways: classical mevalonate (MVA) pathway and 1-deoxyxylulose-5-phosphate (DXP) pathway. These two pathways provide precursors, isopentenyl pyrophosphate (IPP) and its isomer dimethylallyl pyrophosphate (DMAPP), for the synthesis of isoprene (Liu et al., 2005b; Paliyath and Murr, 2006). Mevalonate pathway is localized in the cytosol, and DXP pathway occurs in the chloroplast. In mevalonate pathway, three acetyl-CoA molecules undergo condensation to produce 3-hydroxy-3-methyl-glutaryl CoA (HMG-CoA), which forms mevalonic acid (MVA) in the presence of the enzyme
448 POSTHARVEST BIOLOGY & TECHNOLOGY OF FRUITS, VEGETABLES, & FLOWERS HMG-CoA reductase (HMGR). MVA undergoes phosphorylation, and produces IPP, a five-carbon condensational unit of many terpenes. DMAPP is an isomer of IPP, which is generated in the presence of the enzyme IPP isomerase. These components undergo condensation and produce geranyl (C 10 ), farnesyl (C 15 ), and geranylgeranyl (C 20 ) pyrophosphate compounds, which form monoterpenes, sesquiterpenes, and diterpenes. Monoterpenes and their derivatives (aldehydes, esters, and alcohols) are major volatile compounds in fruits (Paliyath et al., 1997; Paliyath and Murr, 2006). The DXP pathway starts with the condensation of pyruvate and glyceraldehyde-3-P, which generates DXP. Further, a series of reactions produce IPP and DMAPP simultaneously in this pathway (Rohmer et al., 1993). 21.4.3 Biosynthesis of ester volatiles Every fruit has its own characteristic aroma, which is due to the presence of unique monoterpenes, esters, organic acids, aldehydes, and ketones. During lipid catabolism, several alcohols and acids are generated that combine with fatty acyl CoA to form ester volatiles. Fruits, including banana, apples, and strawberry, are enriched in ester volatiles. With the advancement of ripening, production of volatile components increases. The key enzyme in the synthesis of ester volatiles is alcohol acyl CoA transferase (AAT) that transfers an acyl moiety of the acyl-CoA to a corresponding alcohol (Paliyath and Murr, 2006). In fruits such as apple, tomato and melon, the aroma formation depends on ethylene production (Bauchot et al., 1998; Fan et al., 1998; Griffiths et al., 1999). It has been reported that repression of ACC synthase and ACC oxidase reduces aroma components in tomato and melon (El-Sharkawy et al., 2005; Zhu et al., 2005). Lipoxygenase activity resulting in the production of hexanal and hexanol is also affected by ethylene biosynthesis in tomato and apple fruits. It is also noted that the use of AVG, an inhibitor of ethylene synthesis, and diazocyclopentadiene, an ethylene action inhibitor, also reduces the biosynthesis of aroma components in apples (Fan et al., 1998). The application of 1-MCP, an ethylene action inhibitor, also decreases aroma volatile production in apples (Fan et al., 1999). The activity of AAT enzyme was lower in transgenic apples with suppressed ethylene biosynthesis. Furthermore, the exposure of transgenic fruits to 80 μL/L ethylene increased AAT activity to the same level observed in nontransformed apples (Defilippi et al., 2005), suggesting that biosynthesis of flavor volatiles depends on ethylene production. However, it is an open question how ethylene affects the activities of volatile components. 21.4.4 Biosynthesis of anthocyanins Anthocyanins give red, purple, and blue colors to many fruits, vegetables, cereal, and flowers. The stability of anthocyanins depends on the pH and the chelation of metal ions. They are stable under acidic conditions and are rapidly broken down under neutral conditions. Anthocyanins are glycosylated anthocyanidins, and sugars are mostly attached to the 3-hydroxyl position of the anthocyanidins (sometimes to the 5 or 7 position). There are several hundred types of anthocyanins depending on the number of hydroxyl groups, their methylation patterns, and the nature and number of sugars that are attached the molecule. In higher plants, six anthocyanidins are common that include cyanidin (Cy), delphinidin (Dp), peonidin (Pn), petunidin (Pt), malvidin (Mv), and pelargonidin (Pg).
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Postharvest Biology and Technology
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Edition first published 2008 c○ 2
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vi CONTENTS 9 Structural Deteriorat
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Contributors Ishan Adyanthaya Depar
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x CONTRIBUTORS Gopinadhan Paliyath
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xii PREFACE difficult to find a boo
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Chapter 1 Postharvest Biology and T
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POSTHARVEST BIOLOGY AND TECHNOLOGY
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COMMON FRUITS, VEGETABLES, FLOWERS,
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Chapter 3 Biochemistry of Fruits Go
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BIOCHEMISTRY OF FRUITS 21 et al., 2
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BIOCHEMISTRY OF FRUITS 23 3.2.2 Lip
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BIOCHEMISTRY OF FRUITS 25 exposure
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BIOCHEMISTRY OF FRUITS 27 isoform o
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BIOCHEMISTRY OF FRUITS 29 Maltose
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BIOCHEMISTRY OF FRUITS 31 content i
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BIOCHEMISTRY OF FRUITS 33 control.
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BIOCHEMISTRY OF FRUITS 35 range fro
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BIOCHEMISTRY OF FRUITS 37 six (gluc
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BIOCHEMISTRY OF FRUITS 39 Ethylene
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BIOCHEMISTRY OF FRUITS 41 3.3.3 Pro
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BIOCHEMISTRY OF FRUITS 43 component
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Phenylalanine Phenylalanine ammonia
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BIOCHEMISTRY OF FRUITS 47 coloratio
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BIOCHEMISTRY OF FRUITS 49 Fluhr, R.
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Chapter 4 Biochemistry of Flower Se
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BIOCHEMISTRY OF FLOWER SENESCENCE 5
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PROGRAMMED CELL DEATH DURING PLANT
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(a) (a′) (b) (b′) (c) (d) Fig.
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Chapter 6 Ethylene Perception and G
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ETHYLENE PERCEPTION AND GENE EXPRES
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Chapter 7 Enhancing Postharvest She
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ENHANCING POSTHARVEST SHELF LIFE AN
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THE BREAKDOWN OF CELL WALL COMPONEN
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Table 8.3 Transgenic genetics to de
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Chapter 9 Structural Deterioration
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PHOSPHOLIPASE D, MEMBRANE DETERIORA
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Fig. 9.3 Fracture face of a fully o
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PHOSPHOLIPASE D INHIBITION TECHNOLO
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HEAT TREATMENT FOR ENHANCING POSTHA
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THE ROLE OF POLYPHENOLS 261 mainly
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THE ROLE OF POLYPHENOLS 263 Table 1
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THE ROLE OF POLYPHENOLS 265 Pentose
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THE ROLE OF POLYPHENOLS 267 Phenyla
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THE ROLE OF POLYPHENOLS 269 3' OH H
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THE ROLE OF POLYPHENOLS 271 OH OH O
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THE ROLE OF POLYPHENOLS 273 compoun
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THE ROLE OF POLYPHENOLS 275 washing
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THE ROLE OF POLYPHENOLS 277 (Fujita
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THE ROLE OF POLYPHENOLS 279 Boyer,
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THE ROLE OF POLYPHENOLS 281 Peschel
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ISOPRENOID BIOSYNTHESIS IN FRUITS A
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Chapter 14 Postharvest Treatments A
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POSTHARVEST TREATMENTS AFFECTING SE
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Chapter 15 Polyamines and Regulatio
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POLYAMINES AND REGULATION OF RIPENI
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Chapter 16 Postharvest Enhancement
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POSTHARVEST ENHANCEMENT OF PHENOLIC
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RHIZOSPHERE MICROORGANISMS 361 the
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RHIZOSPHERE MICROORGANISMS 363 Rese
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RHIZOSPHERE MICROORGANISMS 365 Tabl
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RHIZOSPHERE MICROORGANISMS 367 Toma
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Chapter 18 Biotechnological Approac
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BIOTECHNOLOGICAL APPROACHES 375 tec
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BIOTECHNOLOGICAL APPROACHES 379 suc
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BIOTECHNOLOGICAL APPROACHES 381 Fla
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BIOTECHNOLOGICAL APPROACHES 383 adm
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BIOTECHNOLOGICAL APPROACHES 387 Bar
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BIOTECHNOLOGICAL APPROACHES 389 Kik
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BIOTECHNOLOGICAL APPROACHES 391 Tat
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POSTHARVEST FACTORS AFFECTING POTAT
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POSTHARVEST FACTORS AFFECTING POTAT
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Page 482 and 483: Index Abscisic acid (ABA), 65, 210,
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