@article {2343,
	title = {Fruit-Surface Flavonoid Accumulation in Tomato Is Controlled by a SIMYB12-Regulated Transcriptional Network},
	journal = {Plos Genetics},
	volume = {5},
	number = {12},
	year = {2009},
	note = {ISI Document Delivery No.: 542CTTimes Cited: 3Cited Reference Count: 58Adato, Avital Mandel, Tali Mintz-Oron, Shira Venger, Ilya Levy, Dorit Yativ, Merav Dominguez, Eva Wang, Zhonghua De Vos, Ric C. H. Jetter, Reinhard Schreiber, Lukas Heredia, Antonio Rogachev, Ilana Aharoni, Asaph},
	month = {Dec},
	pages = {23},
	type = {Article},
	abstract = {The cuticle covering plants{\textquoteright} aerial surfaces is a unique structure that plays a key role in organ development and protection against diverse stress conditions. A detailed analysis of the tomato colorless-peel y mutant was carried out in the framework of studying the outer surface of reproductive organs. The y mutant peel lacks the yellow flavonoid pigment naringenin chalcone, which has been suggested to influence the characteristics and function of the cuticular layer. Large-scale metabolic and transcript profiling revealed broad effects on both primary and secondary metabolism, related mostly to the biosynthesis of phenylpropanoids, particularly flavonoids. These were not restricted to the fruit or to a specific stage of its development and indicated that the y mutant phenotype is due to a mutation in a regulatory gene. Indeed, expression analyses specified three R2R3-MYB-type transcription factors that were significantly down-regulated in the y mutant fruit peel. One of these, SIMYB12, was mapped to the genomic region on tomato chromosome 1 previously shown to harbor the y mutation. Identification of an additional mutant allele that co-segregates with the colorless-peel trait, specific down-regulation of SlMYB12 and rescue of the y phenotype by overexpression of SlMYB12 on the mutant background, confirmed that a lesion in this regulator underlies the y phenotype. Hence, this work provides novel insight to the study of fleshy fruit cuticular structure and paves the way for the elucidation of the regulatory network that controls flavonoid accumulation in tomato fruit cuticle.},
	keywords = {AFFECTS VEGETATIVE DEVELOPMENT, ARABIDOPSIS-THALIANA, EVOLUTION, EXPRESSION, GENE, IDENTIFICATION, LIQUID-CHROMATOGRAPHY, METABOLISM, PHENYLPROPANOID BIOSYNTHESIS, REGULATOR},
	isbn = {1553-7390},
	url = {<Go to ISI>://000273469700027},
	author = {Adato, A. and Mandel, T. and Mintz-Oron, S. and Venger, I. and Levy, D. and Yativ, M. and Dominguez, E. and Wang, Z. H. and De Vos, R. C. H. and Jetter, R. and Schreiber, L. and Heredia, A. and Rogachev, I. and Aharoni, A.}
}
@article {2188,
	title = {Gene expression and metabolism in tomato fruit surface tissues},
	journal = {Plant Physiology},
	volume = {147},
	number = {2},
	year = {2008},
	note = {ISI Document Delivery No.: 308XITimes Cited: 36Cited Reference Count: 115Mintz-Oron, Shira Mandel, Tali Rogachev, Ilana Feldberg, Liron Lotan, Ofra Yativ, Merav Wang, Zhonghua Jetter, Reinhard Venger, Ilya Adato, Avital Aharoni, Asaph},
	month = {Jun},
	pages = {823-851},
	type = {Review},
	abstract = {The cuticle, covering the surface of all primary plant organs, plays important roles in plant development and protection against the biotic and abiotic environment. In contrast to vegetative organs, very little molecular information has been obtained regarding the surfaces of reproductive organs such as fleshy fruit. To broaden our knowledge related to fruit surface, comparative transcriptome and metabolome analyses were carried out on peel and flesh tissues during tomato ( Solanum lycopersicum) fruit development. Out of 574 peel-associated transcripts, 17\% were classified as putatively belonging to metabolic pathways generating cuticular components, such as wax, cutin, and phenylpropanoids. Orthologs of the Arabidopsis ( Arabidopsis thaliana) SHINE2 and MIXTA-LIKE regulatory factors, activating cutin and wax biosynthesis and fruit epidermal cell differentiation, respectively, were also predominantly expressed in the peel. Ultra-performance liquid chromatography coupled to a quadrupole time-of-flight mass spectrometer and gas chromatography-mass spectrometry using a flame ionization detector identified 100 metabolites that are enriched in the peel tissue during development. These included flavonoids, glycoalkaloids, and amyrin-type pentacyclic triterpenoids as well as polar metabolites associated with cuticle and cell wall metabolism and protection against photooxidative stress. Combined results at both transcript and metabolite levels revealed that the formation of cuticular lipids precedes phenylpropanoid and flavonoid biosynthesis. Expression patterns of reporter genes driven by the upstream region of the wax-associated SlCER6 gene indicated progressive activity of this wax biosynthetic gene in both fruit exocarp and endocarp. Peel-associated genes identified in our study, together with comparative analysis of genes enriched in surface tissues of various other plant species, establish a springboard for future investigations of plant surface biology.},
	keywords = {ABC, ARABIDOPSIS-THALIANA, CELL ELONGATION, COTTON FIBER, CUTICULAR STRUCTURE, CUTIN BIOSYNTHESIS, FUNCTIONAL-CHARACTERIZATION, MASS-SPECTROMETRY, MYB TRANSCRIPTION FACTOR, TRANSPIRATION BARRIER PROPERTIES, TRANSPORTER GENE},
	isbn = {0032-0889},
	url = {<Go to ISI>://000256423600034},
	author = {Mintz-Oron, S. and Mandel, T. and Rogachev, I. and Feldberg, L. and Lotan, O. and Yativ, M. and Wang, Z. and Jetter, R. and Venger, I. and Adato, A. and Aharoni, A.}
}
@article {2140,
	title = {Mechanism and Active Site Residues of GDP-Fucose Synthase},
	journal = {Journal of the American Chemical Society},
	volume = {130},
	number = {51},
	year = {2008},
	note = {ISI Document Delivery No.: 406UOTimes Cited: 3Cited Reference Count: 37Lau, Stephen T. B. Tanner, Martin E.},
	month = {Dec},
	pages = {17593-17602},
	type = {Article},
	abstract = {L-Fucose, 6-deoxy-L-galactose, is a key component of many important glycoconjugates including the blood group antigens and the Lewis(x) ligands. The biosynthesis of GDP-L-fucose begins with the action of a dehydratase that converts GDP-D-mannose into GDP-4-keto-6-deoxy-mannose. The enzyme GDPfucose synthase, GFS, (also known as GDP-4-keto-6-deoxy-D-mannose epimerase/reductase, GMER) then converts GDP-4-keto-6-deoxy-D-mannose into GDP-L-fucose. The GFS reaction involves epimerizations at both C-3 {\textquoteright}{\textquoteright} and C-5 {\textquoteright}{\textquoteright} followed by an NADPH-dependent reduction of the carbonyl at C-4. This manuscript describes studies that elucidate the order of the epimerization steps and the roles of the active site acid/base residues responsible for the epimerizations. An active site mutant, Cys109Ser, produces GDP-6-deoxy-D-altrose as its major product indicating that C-3 {\textquoteright}{\textquoteright} epimerization occurs first and premature reduction of the GDP-4-keto-6-deoxy-D-altrose intermediate becomes competitive with GDP-L-fucose production. The same mutation results in the appearance of a kinetic isotope effect when [3 {\textquoteright}{\textquoteright}-H-2]-GDP-6-deoxy-4-keto-mannose is used as a substrate. This indicates that Cys109 is the base responsible for the deprotonation of the substrate at C-3 {\textquoteright}{\textquoteright}. The Cys109Ser mutant also catalyzes a rapid wash-in of solvent derived deuterium into the C-5 {\textquoteright}{\textquoteright} position of GDP-fucose in the presence of NADP(+). This confirms the order of epimerizations and the role of Cys109. Finally, the inactive His179Gln mutant readily catalyzes the wash-out of deuterium from the C-3 {\textquoteright}{\textquoteright} position of [3 {\textquoteright}{\textquoteright}-H-2]-GDP-6-deoxy-4-keto-mannose. Together these results strongly implicate an ordered sequence of epimerizations (C-3 {\textquoteright}{\textquoteright} followed by C-5 {\textquoteright}{\textquoteright}) and suggest that Cys109 acts as a base and His179 acts as an acid in both epimerization steps.},
	keywords = {6-DEHYDRATASE, ARABIDOPSIS-THALIANA, BIOLOGICAL FUNCTION, CATALYTIC MECHANISM, DTDP-GLUCOSE 4, ESCHERICHIA-COLI, GDP-4-KETO-6-DEOXY-D-MANNOSE EPIMERASE/REDUCTASE, HELICOBACTER-PYLORI, LEUKOCYTE ADHESION, MANNOSE 4, UDP-GALACTOSE 4-EPIMERASE},
	isbn = {0002-7863},
	url = {<Go to ISI>://000263320600076},
	author = {Lau, S. T. B. and Tanner, M. E.}
}
@article {2113,
	title = {Plant surface lipid biosynthetic pathways and their utility for metabolic engineering of waxes and hydrocarbon biofuels},
	journal = {Plant Journal},
	volume = {54},
	number = {4},
	year = {2008},
	note = {ISI Document Delivery No.: 299KRTimes Cited: 12Cited Reference Count: 122Jetter, Reinhard Kunst, Ljerka},
	month = {May},
	pages = {670-683},
	type = {Review},
	abstract = {Due to their unique physical properties, waxes are high-value materials that are used in a variety of industrial applications. They are generated by chemical synthesis, extracted from fossil sources, or harvested from a small number of plant and animal species. As a result, the diversity of chemical structures in commercial waxes is low and so are their yields. These limitations can be overcome by engineering of wax biosynthetic pathways in the seeds of high-yielding oil crops to produce designer waxes for specific industrial end uses. In this review, we first summarize the current knowledge regarding the genes and enzymes generating the chemical diversity of cuticular waxes that accumulate at the surfaces of primary plant organs. We then consider the potential of cuticle biosynthetic genes for biotechnological wax production, focusing on selected examples of wax ester chain lengths and isomers. Finally, we discuss the genes/enzymes of cuticular alkane biosynthesis and their potential in future metabolic engineering of plants for the production of renewable hydrocarbon fuels.},
	keywords = {ARABIDOPSIS-THALIANA, BRASSICA-OLERACEA, chain lengths, CONDENSING ENZYME, CUTICULAR WAX, cuticular waxes, ECERIFERUM MUTANTS, EPICUTICULAR WAX, ESTERS, fatty acid elongation, FATTY ACYL-COENZYME, HYDROCARBONS, industrial products, LEAVES PISUM-SATIVUM, MOLECULAR CHARACTERIZATION, SACCHAROMYCES-CEREVISIAE},
	isbn = {0960-7412},
	url = {<Go to ISI>://000255755000012},
	author = {Jetter, R. and Kunst, L.}
}
@article {2238,
	title = {Sealing plant surfaces: Cuticular wax formation by epidermal cells},
	journal = {Annual Review of Plant Biology},
	volume = {59},
	year = {2008},
	note = {ISI Document Delivery No.: 311IFTimes Cited: 55Cited Reference Count: 148Samuels, Lacey Kunst, Ljerka Jetter, Reinhard},
	pages = {683-707},
	type = {Review},
	abstract = {The vital importance of plant surface wax in protecting tissue from environmental stresses is reflected in the huge commitment of epidermal cells to cuticle formation. During cuticle deposition, a massive flux of lipids occurs from the sites of lipid synthesis in the plastid and the endoplasmic reticulum to the plant surface. Recent genetic studies in Arabidopsis have improved our understanding of fatty acid elongation and of the subsequent modification of the elongated products into primary alcohols, wax esters, secondary alcohols, and ketones, shedding light on the enzymes involved in these pathways. In contrast, the biosynthesis of alkanes is still poorly understood, as are the mechanisms of wax transport from the site of biosynthesis to the cuticle. Currently, nothing is known about wax trafficking from the endoplasmic reticulum to the plasma membrane, or about translocation through the cell wall to the cuticle. However, a first breakthrough toward an understanding of wax export recently came with the discovery of ATP binding cassette (ABC) transporters that are involved in releasing wax from the plasma membrane into the apoplast. An overview of our present knowledge of wax biosynthesis and transport and the regulation of these processes during cuticle assembly is presented, including the evidence for coordination of cutin polyester and wax production.},
	keywords = {ABC, ABC transporters, ACYL-COA SYNTHETASE, ALKANES, ARABIDOPSIS, ARABIDOPSIS-THALIANA, CHAIN FATTY-ACID, COENZYME-A SYNTHETASE, DEEP-WATER RICE, ECERIFERUM CER MUTANTS, ENDOPLASMIC-RETICULUM, fatty acid elongation, LEAVES PISUM-SATIVUM, LIPID-TRANSFER PROTEIN, primary alcohols, regulation of wax biosynthesis, TRANSPORTER},
	isbn = {1040-2519},
	url = {<Go to ISI>://000256593200027},
	author = {Samuels, L. and Kunst, L. and Jetter, R.}
}
@article {963,
	title = {Plant cuticular lipid export requires an ABC transporter},
	journal = {Science},
	volume = {306},
	number = {5696},
	year = {2004},
	note = {ISI Document Delivery No.: 866ENTimes Cited: 103Cited Reference Count: 10},
	month = {Oct},
	pages = {702-704},
	type = {Article},
	abstract = {A waxy protective cuticle coats all primary aerial plant tissues. Its synthesis requires extensive export of lipids from epidermal cells to the plant surface. Arabidopsis cer5 mutants had reduced stem cuticular wax loads and accumulated sheetlike inclusions in the cytoplasm of wax-secreting cells. These inclusions represented abnormal deposits of cuticular wax and resembled inclusions found in a human disorder caused by a defective peroxisomal adenosine triphosphate binding cassette (ABC) transporter. We found that the CER5 gene encodes an ABC transporter localized in the plasma membrane of epidermal cells and conclude that it is required for wax export to the cuticle.},
	keywords = {ARABIDOPSIS-THALIANA, CANCER RESISTANCE PROTEIN, MUTANTS},
	isbn = {0036-8075},
	url = {<Go to ISI>://000224756700056},
	author = {Pighin, J. A. and Zheng, H. Q. and Balakshin, L. J. and Goodman, I. P. and Western, T. L. and Jetter, R. and Kunst, L. and Samuels, A. L.}
}
@article {1026,
	title = {Tomato fruit cuticular waxes and their effects on transpiration barrier properties: functional characterization of a mutant deficient in a very-long-chain fatty acid beta-ketoacyl-CoA synthase},
	journal = {Journal of Experimental Botany},
	volume = {55},
	number = {401},
	year = {2004},
	note = {ISI Document Delivery No.: 829GETimes Cited: 49Cited Reference Count: 44},
	month = {Jun},
	pages = {1401-1410},
	type = {Article},
	abstract = {Cuticular waxes play a pivotal role in limiting transpirational water loss across the plant surface. The correlation between the chemical composition of the cuticular waxes and their function as a transpiration barrier is still unclear. In the present study, intact tomato fruits (Lycopersicon esculentum) are used, due to their astomatous surface, as a novel integrative approach to investigate this composition-function relationship: wax amounts and compositions of tomato were manipulated before measuring unbiased cuticular transpiration. First, successive mechanical and extractive wax-removal steps allowed the selective modification of epi- and intracuticular wax layers. The epicuticular film consisted exclusively of very-long-chain aliphatics, while the intracuticular compartment contained large quantities of pentacyclic triterpenoids as well. Second, applying reverse genetic techniques, a loss-of-function mutation with a transposon insertion in a very-long-chain fatty acid elongase beta-ketoacyl-CoA synthase was isolated and characterized. Mutant leaf and fruit waxes were deficient in n-alkanes and aldehydes with chain lengths beyond C-30, while shorter chains and branched hydrocarbons were not affected. The mutant fruit wax also showed a significant increase in intracuticular triterpenoids. Removal of the epicuticular wax layer, accounting for one-third of the total wax coverage on wild-type fruits, had only moderate effects on transpiration. By contrast, reduction of the intracuticular aliphatics in the mutant to approximately 50\% caused a 4-fold increase in permeability. Hence, the main portion of the transpiration barrier is located in the intracuticular wax layer, largely determined by the aliphatic constituents, but modified by the presence of triterpenoids, whereas epicuticular aliphatics play a minor role.},
	keywords = {ARABIDOPSIS-THALIANA, BARRIER, BICOLOR L MOENCH, CITRUS LEAF CUTICLES, CONDENSING ENZYME, CONDUCTANCE, cuticle, ECERIFERUM CER MUTANTS, EPICUTICULAR WAX, EPICUTICULAR WAXES, EPIDERMAL, intracuticular wax, PRUNUS-LAUROCERASUS L, tomato, transpiration, TRANSPORT-PROPERTIES, WATER PERMEABILITY},
	isbn = {0022-0957},
	url = {<Go to ISI>://000222034600015},
	author = {Vogg, G. and Fischer, S. and Leide, J. and Emmanuel, E. and Jetter, R. and Levy, A. A. and Riederer, M.}
}
