Postharvest Biology and Technology of Fruits, Vegetables, and Flowers

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PHOSPHOLIPASE D, MEMBRANE DETERIORATION, AND SENESCENCE 217 1 2 3 β-1 β-2 β-3 β-4 LePLDα1 6 LHGTLHVTIFEVDNLQGEEEGGHFFSKIKQHFEETVGIGKGTPKLYATIDLEKARVGRTRIIENE 70 FaPLDα1 9 LHGTLHATIYEVDKLHGS--SGNFLRKITGKLEETVGLGKGVSKLYATVDLERARVGRTRVIEKE 71 CmPLDα1 6 LHGTLHATIYEIDRLHTGG-SSNVFSMLRQNFEEAVGIGKGQTKLYATIDLEKARVGRTRILESE 69 CsPLDα1 6 LHGTLHATIYEIDRLHTGG-SSNVFSMLRQNFEEAVGIGKGQTKLYATIDLEKARVGRTRILESD 69 CmPLDα2 6 LHGTLHVTIYEVDKLHSGG--RNFLKQLVENVEEAVGFGRGITRLYATIDLEKARVGRTRRLERE 68 LePLDα2 6 LHGTLHVTIFEVDKLRTNF-GREIFNKVVQGIEGAIGFNKTASTLYATIDLGKARVGRTRLLD-E 68 LePLDα3 6 LHGTLHVTIFEVDRLHTNF-GRDFFNKVVQGIEGAIGFNKAASRLYATIDLGKARVGRTRLLD-D 68 4 5 β-1 β-6 β-7 β-8 LePLDα1 71 PKNPRWYESFHIYCAHMASNVIFTIKDDNPFGASLIGRAYVPVEELLEGEEIDKWVEIMDKEMNP 135 FaPLDα1 72 PSNPNWSESFHIYCAHVAANVIFTVKESNPIGASLIGRAYVPVEQLIEGEEVDTWAKILDDKKNP 136 CmPLDα1 70 PSNPRWYESFHIYCAHKASNVIFTVKDDNPIGATLIGHAYVPVEDIVDGEEVDRWVPILDENQNP 134 CsPLDα1 70 PSNPKWNESFHIYCAHKASNVIFTVKDDNPIGATLIGRAYVPVEDIVDGEEVDKWVPILDENQNP 134 CmPLDα2 69 HSNPKWYETFHIYCAHMASNIIFTVKDDNPIGATLIGRAYLPVREIIRGDEVDKWVPILDEQKKP 133 LePLDα2 69 HKNPRWYESFHIYCAHMASDVVFTVKADNPIGAELIGRAYLPVEQLIVGEVVDEWLEILDTERKP 133 LePLDα3 69 HKNPRWYESFHIYCAHMAANVIITVKFDNPIGAEVIGRAYFPVQQLLDGEEVDEWLEILNTERKP 133 (a) LePLDα1 FaPLDα1 CmPLDα1 CsPLDα1 CmPLDα2 LePLDα2 LePLDα3 (b) LePLDα1 FaPLDα1 CmPLDα1 CsPLDα1 CmPLDα2 LePLDα2 LePLDα3 (c) ↑ ↑ ↑ ↑ AS-1 300 QGTDVNCVLCPRNPDNGGSFVQDIQISTMFTHHQKIIVVDSALPSGESEKRRILSFVGGID 360 301 QNTDVNCVLCPRNPDGGGSIVQGAQISTMFTHHQKIVVVDSEMPNGGSQSRRIVSFVGGLD 361 299 QDTDVHCVLCPRNPDDGGSIVQDLQISTMFTHHQKIVVVDSPMPNGDSDRRRIVSFVGGID 359 299 QDTDVHCVLCPRNPDDGGSIVQDLQISTMFTHHQKIVVVDSPMPNGDSDKRRIVSFVGGID 359 298 ANTDVHCVLCPRNPDDGANVIQDIAVGTMFTHHQKIVVVDSALPNGDPSKRRIVSFVGGLD 358 298 ENSEVHCVLCPRNPDDGRSIIQNIEIGTMFTHHQKIVVVDGELPNGDTERRRIVSYIGGID 358 298 RGTQVSCVLCPRNPDDGRSIIQNIEIGTMFTHHQKIVIVDGEMPNGDRERRRIVSYIGGLD 358 PEST-rich sequence AS-2 629 KKSGEYEPSESPEPDSNYMRAQEARRFMIYVHSKMMIVDDEYIIVGSANINQRSMDGARD 688 630 KKDGEYEPSEAPEADSDYIRAQEARRFMIYVHTKMMIVDDEYIIIGSANINQRSMDGARD 689 628 KRSGEYEPSEAPEEDSDYLRAQQARRFMIYVHTKMMIVDDEYIIVGSANINQRSMDGARD 687 628 KRSGEYEPSEAPEEDSDYLRAQQARRFMIYVHTKMMIVDDEYIIVGSANINQRSMDGARD 687 627 KKPGEYEPSESPDENSDYLRAQQSRRFMIYVHSKMMIVDDEYIIVGSANINQRSMDGARD 686 627 KKTGEYEPSESPEPDSDYQKAQEARRFMIYVHAKMMIVDDEYIIIGSANINQRSMDGARD 686 627 KKRGEYEPCEPPEPNSGYHKAQEARRFMIYVHSKMMIVDDEYIIIGSANINQRSMDGARD 686 ↑ Fig. 9.12 Alignment of the encoded amino acid sequences of PLD alphas from tomato (LePLDα1, LePLDα2, and LePLDα3), strawberry (FaPLDα1), honeydew melon (CmPLDα1 and CmPLDα2), and cucumber (CsPLDα1) comparing (a) the C2 Ca 2+ /phospholipid-binding domain, (b) the first HKD active site motif (AS-1), and (c) a PEST-rich proteolysis-targeting sequence plus the second HKD-GSN active site motif (AS-2). ↑ ↑ ↑ ↑ ↑ ↑ website showed the presence of a C2 Ca 2+ /phospholipids-binding domain and two PLD active site motifs, HxKxxxxD and HxKxxxxDxxxxxxGSxN (Pappan et al., 1997; Stuckey and Dixon, 1999; Wang, 2000), in each of the seven PLD alphas from tomato, strawberry, melon, and cucumber (Fig. 9.12). Despite the relatively poor conservation between PLD alphas and other plant PLDs (about 40–44% identity with PLDs beta, gamma, and delta), these features are present in all types of plant PLDs except PLD zeta, which includes plextrin and phox homology domains and lacks the C2 domain (Wang et al., 2006). C2 domains occur in a variety of proteins that in one way or another interact with membranes. Calcium binding to specific sites in the C2 domain confers a charge to the protein, facilitating electrostatic association with anionic sites on a membrane (Ponting and Kerr, 1996; Nalefski et al., 1997). Thus, in plant PLDs including a C2 domain, calcium-activated membrane association of an otherwise soluble, cytosolic enzyme appears to involve an
218 POSTHARVEST BIOLOGY & TECHNOLOGY OF FRUITS, VEGETABLES, & FLOWERS electrostatic rather than a hydrophobic interaction. It was also recently shown that the C2 domain of PLD alphas possesses binding sites that enable Ca 2+ -independent association with an aspartic protease, cardosin A, involving RGD and KGE motifs in the protease sequence (Simões et al., 2005). The C2 domain in plant PLD alphas comprises the range of amino acids from about 6 to 150 (Fig. 9.12), and as in other C2 superfamily proteins is characterized by the presence of eight beta strand motifs that form an antiparallel beta sandwich (Nalefski et al., 1997; Rizo and Südhof, 1998). The proposed locations of the eight beta strands in C2 of plant PLD alphas and betas appear to vary depending on whether modeling is based on the crystal structure of rat synaptotagmin or PLC delta 1 and cPLA 2 (Pappan et al., 1997; Rizo and Südhof, 1998; Zheng et al., 2000). Nevertheless, a common feature in both models is the presence of several critical amino acids that participate in binding of up to four Ca 2+ ions and are located at specific sites on exposed loops connecting the beta sheets. In Arabidopsis PLD beta 1 and rat synaptotagmin, four acidic amino acid residues (glutamic acid; E or aspartic acid; D) were identified as those most involved in Ca 2+ binding (Pappan et al., 1997). A subsequent alignment of Arabidopsis PLD alpha and beta with PLC delta 1 and cPLA 2 confirmed three of those four amino acids and identified a single asparagine (N) residue as also being important (Zheng et al., 2000). The five corresponding residues in LePLDα1 occur at AA 38, 49, 69, 97, and 99 (Fig. 9.12), and notably, acidic AAs at 49 and 99 have been substituted by basic lysine (K) and polar but uncharged asparagine (N), respectively. These, or similar substitutions seen in the seven fruit PLD alphas were thought to account for the requirement of alpha-type PLDs for millimolar rather than micromolar concentrations of calcium for activity at neutral pH (Wang, 2000, 2001; Zheng et al., 2000). However, other studies (Yuan et al., 2006a; Tiwari and Paliyath, 2007) have confirmed that strawberry PLD alpha and a tomato PLD alpha C2 domain show activation at physiologically activated, low micromolar levels of calcium suggesting the presence of high-affinity calcium-binding sites. It is interesting that the third of the four critical acidic AAs, at position 95 in LePLDα2 and LePLDα3, is substituted by nonpolar and hydrophobic alanine or phenylalanine (Fig. 9.12), raising the possibility that Ca 2+ binding is severely compromised in these two PLD alphas. As well, LePLDα1 is the only PLD alpha of the seven to have asparagine (AA 69) at the nonacidic Ca 2+ -binding position found in PLC delta 1 and cPLA 2 , although all the PLD alphas have two acidic AAs (E or D) adjacent to this residue that could take part in binding calcium ions. Conceivably the C2 domains in the seven fruit PLD alphas could exist in different topologies, thus exposing different sets of calcium-binding sites, but nevertheless exhibiting conformational changes required for phospholipid (membrane) binding, some of these at physiologically activated cytosolic calcium levels. C2 in Arabidopsis PLD beta was shown to have three high-affinity Ca 2+ -binding sites with dissociation constants (K d s) of 0.8, 2.4, and 24 μM, whereas calorimetry data indicated that C2 of Arabidopsis PLD alpha has 1–3 low-affinity sites with K d in the range of 470– 590 μM (Zheng et al., 2000). However, this property in Arabidopsis PLD alpha is a unique situation and may not be reflective of several PLD alphas that show high-affinity calciumbinding sites and are activated at micromolar calcium levels. Aside from in the C2 domain, Pappan et al. (2004) demonstrated that specific binding of calcium ions also occurs in the catalytic portion of Arabidopsis PLD beta 1. A bacterially expressed PLD beta 1 lacking the C2 region had much lower activity than the complete enzyme, required higher levels of calcium for maximum activity, and was activated by phosphatidylserine much more than
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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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THE BREAKDOWN OF CELL WALL COMPONEN
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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 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 377 pro
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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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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 431 e
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BIOSENSOR-BASED TECHNOLOGIES 433 el
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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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