@article {1531,
	title = {CER4 encodes an alcohol-forming fatty acyl-coenzyme A reductase involved in cuticular wax production in Arabidopsis},
	journal = {Plant Physiology},
	volume = {142},
	number = {3},
	year = {2006},
	note = {ISI Document Delivery No.: 103NGTimes Cited: 40Cited Reference Count: 64Rowland, Owen Zheng, Huanquan Hepworth, Shelley R. Lam, Patricia Jetter, Reinhard Kunst, Ljerka},
	month = {Nov},
	pages = {866-877},
	type = {Article},
	abstract = {A waxy cuticle that serves as a protective barrier against uncontrolled water loss and environmental damage coats the aerial surfaces of land plants. It is composed of a cutin polymer matrix and waxes. Cuticular waxes are complex mixtures of very-long-chain fatty acids and their derivatives. We report here the molecular cloning and characterization of CER4, a wax biosynthetic gene from Arabidopsis (Arabidopsis thaliana). Arabidopsis cer4 mutants exhibit major decreases in stem primary alcohols and wax esters, and slightly elevated levels of aldehydes, alkanes, secondary alcohols, and ketones. This phenotype suggested that CER4 encoded an alcohol-forming fatty acyl-coenzyme A reductase (FAR). We identified eight FAR-like genes in Arabidopsis that are highly related to an alcohol-forming FAR expressed in seeds of jojoba (Simmondsia chinensis). Molecular characterization of CER4 alleles and genomic complementation revealed that one of these eight genes, At4g33790, encoded the FAR required for cuticular wax production. Expression of CER4 cDNA in yeast (Saccharomyces cerevisiae) resulted in the accumulation of C24: 0 and C26: 0 primary alcohols. Fully functional green fluorescent protein-tagged CER4 protein was localized to the endoplasmic reticulum in yeast cells by confocal microscopy. Analysis of gene expression by reverse transcription-PCR indicated that CER4 was expressed in leaves, stems, flowers, siliques, and roots. Expression of a beta-glucuronidase reporter gene driven by the CER4 promoter in transgenic plants was detected in epidermal cells of leaves and stems, consistent with a dedicated role for CER4 in cuticular wax biosynthesis. CER4 was also expressed in all cell types in the elongation zone of young roots. These data indicate that CER4 is an alcohol-forming FAR that has specificity for very-long-chain fatty acids and is responsible for the synthesis of primary alcohols in the epidermal cells of aerial tissues and in roots.},
	keywords = {ACID ELONGASE, CELL-DIFFERENTIATION, CUTICLE DEVELOPMENT, ECERIFERUM MUTANTS, EPICUTICULAR WAXES, GENE-EXPRESSION MAP, NORMAL ACCUMULATION, PISUM-SATIVUM, POLLEN FERTILITY, TRANSCRIPTIONAL ACTIVITY},
	isbn = {0032-0889},
	url = {<Go to ISI>://000241892900007},
	author = {Rowland, O. and Zheng, H. Q. and Hepworth, S. R. and Lam, P. and Jetter, R. and Kunst, L.}
}
@article {792,
	title = {The SHINE clade of AP2 domain transcription factors activates wax biosynthesis, alters cuticle properties, and confers drought tolerance when overexpressed in Arabidopsis},
	journal = {Plant Cell},
	volume = {16},
	number = {9},
	year = {2004},
	note = {ISI Document Delivery No.: 854TTTimes Cited: 99Cited Reference Count: 62},
	month = {Sep},
	pages = {2463-2480},
	type = {Article},
	abstract = {The interface between plants and the environment plays a dual role as a protective barrier as well as a medium for the exchange of gases, water, and nutrients. The primary aerial plant surfaces are covered by a cuticle, acting as the essential permeability barrier toward the atmosphere. It is a heterogeneous layer composed mainly of lipids, namely cutin and intracuticular wax with epicuticular waxes deposited on the surface. We identified an Arabidopsis thaliana activation tag gain-of-function mutant shine (shn) that displayed a brilliant, shiny green leaf surface with increased cuticular wax compared with the leaves of wild-type plants. The gene responsible for the phenotype encodes one member of a clade of three proteins of undisclosed function, belonging to the plant-specific family of AP2/EREBP transcription factors. Overexpression of all three SHN clade genes conferred a phenotype similar to that of the original shn mutant. Biochemically, such plants were altered in wax composition (very long fatty acid derivatives). Total cuticular wax levels were increased sixfold in shn compared with the wild type, mainly because of a ninefold increase in alkanes that comprised approximately half of the total waxes in the mutant. Chlorophyll leaching assays and fresh weight loss experiments indicated that overexpression of the SHN genes increased cuticle permeability, probably because of changes in its ultrastructure. Likewise, SHN gene overexpression altered leaf and petal epidermal cell structure, trichome number, and branching as well as the stomatal index. Interestingly, SHN overexpressors displayed significant drought tolerance and recovery, probably related to the reduced stomatal density. Expression analysis using promoter-beta-glucuronidase fusions of the SHN genes provides evidence for the role of the SHN clade in plant protective layers, such as those formed during abscission, dehiscence, wounding, tissue strengthening, and the cuticle. We propose that these diverse functions are mediated by regulating metabolism of lipid and/or cell wall components.},
	keywords = {CER MUTANTS, CHEMICAL-COMPOSITION, CONDENSING ENZYME, CUTICULAR WAX, ECERIFERUM, EPICUTICULAR WAX, EPIDERMAL-CELL DIFFERENTIATION, ORGAN FUSION, POLLEN FERTILITY, RESPONSIVE GENE-EXPRESSION, SEPARATION PROCESSES},
	isbn = {1040-4651},
	url = {<Go to ISI>://000223927000018},
	author = {Aharoni, A. and Dixit, S. and Jetter, R. and Thoenes, E. and van Arkel, G. and Pereira, A.}
}
