Postharvest Biology and Technology of Fruits, Vegetables, and Flowers

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ISOPRENOID BIOSYNTHESIS IN FRUITS AND VEGETABLES 283 and their processed products because consumers prefer tomatoes and products with intense red color. As well, the carotenoids are nutraceuticals because they play a beneficial role in regulating health. Inverse associations have been observed between blood lycopene levels and the risk of cancer development at several sites in the human body (Giovannucci, 1999). The most beneficial effects have been observed in the reduction of the incidence of prostate, lung, and stomach cancers, with slightly lower benefits at other sites that include pancreas, colon, rectum esophagus, oral cavity, and cervix (Giovannucci, 1999). β-Carotene (provitamin A) is a vitamin A precursor, and considerable efforts have been made to increase dietary levels of β-carotene by engineering the genes for the carotenoid biosynthetic pathway enzymes into rice endosperm (Ye et al., 2000). Vitamin A deficiency causes xerophthalmia and blindness (Mayne, 1996) and has led to over a million deaths annually among children 1–4 years of age (Humphrey et al., 1992). Finally, levels of dietary lutein (Seddon et al., 1994; Curran-Celentano et al., 2001) and serum lutein (EDCCSG, 1992, 1993) are related to a reduced risk of age-related macular degeneration. For these reasons, there is a tremendous interest among the industry and grower sectors to obtain tomatoes with the highest carotenoid levels possible. 13.3 Carotenoid biosynthesis Carotenoids are biosynthesized by the chloroplast-localized isoprenoid pathway. The accumulation of lycopene (C 40 ) during fruit ripening results from increased lycopene synthesis as well as decreased conversion of lycopene to more complex carotenoids—with the notable exception being the formation of small amounts of β-carotene at the expense of lycopene (Ronen et al., 2000). At least two enzymes are known to regulate carotenoid synthesis: 1-deoxy-D-xylulose-5-phosphate synthase (DXS) and phytoene synthase (PSY, which is coded by PSY1 in ripe fruit). DXS appears to be the paramount of several enzymes involved in the synthesis of isopentenyl pyrophosphate, which is the key 5-carbon unit of the isoprenoid pathway (Fig. 13.1). Within this pathway, PSY forms phytoene (C 40 ), which is an immediate precursor to lycopene (Fig. 13.2). Increased expression of DXS and PSY1 genes during ripening correlates with the pattern of lycopene accumulation, thus demonstrating transcriptional regulation of the pathway during ripening (Lois et al., 2000). As expected, PSY activity also correlates with these patterns of mRNA transcription and lycopene levels (Fraser et al., 1994). The accumulation of lycopene in tomato fruit has been found to be regulated by light, and this is mediated by phytochromes (Alba et al., 2000a). Phytochromes are light receptors involved in responses regulated by red light (R) and far-red light (FR) resulting in alteration of growth and development. In particular, these modified proteins change conformations depending on whether they have received R or FR; however, it is R that activates the protein within the cell to cause physiological responses (Quail, 2002). Most phytochrome responses occur at low-photon fluence rates (1–1,000 μmol/m 2 ) once a given number of photons have been intercepted, irrespective of the duration of the light exposure (Neff et al., 2000). As well, they are commonly characterized by R/FR reversibility. 13.4 Composition of tomato pigments The ripening of tomato fruit is characterized by a large shift in pigment composition. Pigmentation of immature and mature green fruit is similar to that of other photosynthetic
284 POSTHARVEST BIOLOGY & TECHNOLOGY OF FRUITS, VEGETABLES, & FLOWERS GAP + Pyruvate DXS 1-Deoxy-D-xylulose-5-P PDS DMAPP +IPP GGPP (C 20 ) 2x PSY Phytoene (C 40 ) ζCDS ζ-Carotene Lycopene IPP (C 5 ) ε-Cyclase γ-Carotene β-Carotene Zeaxanthin β-Cyclase δ-Carotene α-Carotene Lutein Fig. 13.1 Plastid-localized DXP/isoprenoid pathway in plants. Some key enzymes (bold face) and intermediates involved in isoprenoid biosynthesis are abbreviated: β-cyclase, ε-cyclase; ζ CDS, ζ -carotene desaturase; DMAPP, dimethylallyl pyrophosphate; DXS, 1-deoxy-D-xylulose-5-phosphate synthase; GA-3P, glyceraldehyde- 3-phosphate; GGPP, geranylgeranyl pyrophosphate; IPP, isopentenyl pyrophosphate; PDS, phytoene desaturase; PSY, phytoene synthase. tissue (i.e., the leaves): chlorophyll accounts for the green color that masks the effects of most other chromophores. The major role of the carotenoids in these tissues is to act as light receptors and protectors of the photosynthetic apparatus. The major carotenoids that accumulate in ripe tomato fruits are lycopene (∼90%), β-carotene (5–10%), and lutein (1–5%), with trace amounts (
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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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Page 248 and 249: PHOSPHOLIPASE D, MEMBRANE DETERIORA
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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 369 Such
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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 389 Kik
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BIOTECHNOLOGICAL APPROACHES 391 Tat
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POSTHARVEST FACTORS AFFECTING POTAT
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BIOSENSOR-BASED TECHNOLOGIES 419 20
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BIOSENSOR-BASED TECHNOLOGIES 421 Ta
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BIOSENSOR-BASED TECHNOLOGIES 425 Li
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BIOSENSOR-BASED TECHNOLOGIES 427 So
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BIOSENSOR-BASED TECHNOLOGIES 429 Pr
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BIOSENSOR-BASED TECHNOLOGIES 435 st
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Cl O O O OH Cl O OH Cl Cl Cl 2,4-Di
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BIOSENSOR-BASED TECHNOLOGIES 439 O
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BIOSENSOR-BASED TECHNOLOGIES 441 Le
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Chapter 21 Changes in Nutritional Q
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CHANGES IN NUTRITIONAL QUALITY OF F
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Index Abscisic acid (ABA), 65, 210,
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INDEX 469 Biosensor-based technolog
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INDEX 471 Cryptochlorogenic acid (4
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INDEX 473 French bean, 95 Fresh-cut
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INDEX 475 LePLDα3 (AY013253), 213-
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INDEX 477 Pectin methylesterase (PM
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INDEX 479 PSY1 expression, 289 PSY1
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INDEX 481 Sugars, biosynthesis of,
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