Postharvest Biology and Technology of Fruits, Vegetables, and Flowers

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CHANGES IN NUTRITIONAL QUALITY OF FRUITS AND VEGETABLES 449 Phenylalanine PAL Cinnamic acid C4H p-Coumaric acid 4CL p-Coumaryl CoA Phenyl propanoid pathway Acetyl-CoA Malonyl-CoA CHS Flavonoid pathway Chalcone CHI Flavanone (naringenin) F3H Dihydroflavonol DFR Leucoanthocyanidin ANS Anthocyanidin UFGT Anthocyanin Glycosylated anthocyanindins Petunidin Peonidin Malvidin Cyanindin Delphinidin Pelargonidin Fig. 21.2 A simplified presentation of anthocyanin biosynthesis in plants. PAL, phenylalanine ammonia lyase; C4H, cinnamate-4-hyroxylase; 4CL, 4-coumaryl CoA ligase; CHS, chalcone synthase; CHI, chalcone isomerase; F3H, flavanone-3-hydroxylase; DFR, dihydroflavonol-4-reductase; ANS, anthocyanin synthase; UFGT, UDPglucose flavonoid 3-oxy-glucosyltransferase. (Adapted from Jaakola et al., 2002.) Anthocyanins are synthesized through flavonoid biosynthetic pathway, which is shown in Fig. 21.2. The first step of anthocyanin biosynthesis is the condensation of three molecules of malonyl-CoA with p-coumaroyl-CoA in the presence of the enzyme CHS, which produces chalcone. In the presence of chalcone isomerase, chalcone is converted to flavanone (naringenin) (Jaakola et al., 2002). Flavanone is hydroxylated through flavanone- 3-hydroxylase (F3H) and forms dihydroflavonols, which differ in the number of hydroxyl groups. Dihydroflavonol-4-reductase (DFR) transforms dihydroflavonols to colorless leucoanthocyanidin, which yields colored anthocyanidins in the presence of the enzyme anthocyanin synthase (ANS). The glycosylation of anthocyanidins leads to the formation of anthocyanins by the enzyme UDP-glucose flavonoid 3-oxy-glucosyltransferase (UFGT).
450 POSTHARVEST BIOLOGY & TECHNOLOGY OF FRUITS, VEGETABLES, & FLOWERS Different anthocyanidins give different colors to fruits (Jaakola et al., 2002; Paliyath and Murr, 2006). In red and nonred apples, differential expression of anthocyanin biosynthetic genes was observed. The expression of CHS, F3H, DFR, and ANS genes was found in red apples when anthocyanin was not detected (before ripening stages). However, UFGT gene expression was observed only during the ripening stages with the formation of anthocyanins. This suggests that UFGT gene expression regulates the production of anthocyanins during ripening in apples (Kondo and Hiraoka, 2002). Environmental factors such as light and temperature affect anthocyanin accumulation in fruits. No anthocyanin was found in apples in the absence of light (Proctor, 1974). 21.4.5 Interrelationships of metabolic pathways In fruits, quality-attributing pathways are interrelated. During starch degradation, glucose-1- phosphate is generated, which enters into several metabolic pathways such as glycolysis and PPP. Glycolysis converts glucose to acetyl CoA after a series of reactions. The acetyl CoA is a precursor for the synthesis of fatty acids, volatile esters, isoprenoids, and organic acids as show in Fig. 21.3. The PPP is an important pathway, which provides carbon skeletons to several biosynthetic pathways including amino acids, secondary metabolites, and nucleic acids. Besides providing carbon skeleton, it also gives reducing energy (NADPH) to various pathways (Fig. 21.3). The carbon skeletons for the synthesis of anthocyanins come from p-coumaroyl-CoA and malonyl-CoA. The phenylpropanoid pathway donates p-coumaroyl- CoA, which derives from erythrose-4-phosphate and pyruvate that are generated during PPP. Therefore, PPP plays a significant role in the fruit quality regulation. In flavonoid biosynthesis, the hydroxylation of flavonoids by F3H requires NADPH, which comes from PPP. Isoprenoid synthesis also needs NADPH from PPP (Fig. 21.3). Moreover, it is an important component for the antioxidant enzyme system. The enzymes of ascorbate–glutathione cycle, glutathione reductase (GR), and monodehydroascorbate reductase (MDHAR), require NADPH to inactivate reactive oxygen species (ROS) generated during stress and senescence (Fig. 21.3). In order to maintain fruit quality and shelf life, the precise functioning of the antioxidant enzyme system is necessary. During membrane lipid catabolism, several fatty acid intermediates are formed that are required for the synthesis of aroma volatile components. The precursors of volatile esters, including hexanal, hexanol, and short-chain alcohols, are also derived from metabolism of lipids. The effect of hexanal has been investigated on PLD activity of membrane, and soluble fractions isolated from corn kernel (Paliyath et al., 1999). Inclusion of hexanal in assay mixture resulted in over 75% inhibition of PLD activity. Other technologies have also been used for the inhibition of PLD activity. A naturally occurring lipid, lysophosphatidylethanolamine (LPE), showed potent inhibition of PLD activity in leaves, flowers, and postharvest fruits. Ryu et al. (1997) noticed a reduction in PLD activity by increasing the length and unsaturation of LPE acyl chain. In addition, decreased ethylene production was found in LPE-treated fruits. Antisense PLD tomatoes showed PLD inhibition. Furthermore, they also possessed higher levels of lycopene, firmness, and soluble solids in the fruits (Pinhero et al., 2003), suggesting that by inhibiting certain metabolic pathways, quality components and shelf life of fruits can be increased.
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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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RHIZOSPHERE MICROORGANISMS 371 Gian
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Chapter 18 Biotechnological Approac
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BIOTECHNOLOGICAL APPROACHES 375 tec
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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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