BIOCHEMISTRY AND MOLECULAR BIOLOGY OF WAX PRODUCTION IN PLANTS.

The aerial surfaces of plants are covered with a wax layer that is primarily a waterproof barrier but that also provides protection against environmental stresses. The ubiquitous presence of cuticular wax is testimony to its essential function. Genetic and environmental factors influence wax quantity and composition, which suggests that it is an actively regulated process. The basic biochemistry of wax production has been elucidated over the past three decades; however, we still know very little about its regulation. This review presents a discussion along with new perspectives on the regulatory aspects of wax biosynthesis. Among the topics discussed are the partitioning of fatty acid precursors into wax biosynthesis and the elongation of fatty acids with particular emphasis on the nature of the acyl primer, and the role of ATP in fatty acid elongation. The recent cloning of wax biosynthetic genes and the transport of wax to plant surfaces are also discussed.

[1]  M. Lassner,et al.  A jojoba beta-Ketoacyl-CoA synthase cDNA complements the canola fatty acid elongation mutation in transgenic plants. , 1996, The Plant cell.

[2]  P. Kolattukudy.  Lipid-Derived Defensive Polymers and Waxes and Their Role in Plant–Microbe Interaction , 1987 .

[3]  D. W. James,et al.  Directed tagging of the Arabidopsis FATTY ACID ELONGATION1 (FAE1) gene with the maize transposon activator. , 1995, The Plant cell.

[4]  P. Kolattukudy,et al.  Alkane biosynthesis by decarbonylation of aldehyde catalyzed by a microsomal preparation from Botryococcus braunii. , 1991, Archives of biochemistry and biophysics.

[5]  S. Eigenbrode,et al.  Effects of Plant Epicuticular Lipids on Insect Herbivores , 1995 .

[6]  D. Liu,et al.  Discovery of an Epidermal Stearoyl-Acyl Carrier Protein Thioesterase. , 1995, The Journal of Biological Chemistry.

[7]  E. Bermadinger-Stabentheiner.  Physical injury, re‐crystallization of wax tubes and artefacts: identifying some causes of structural alteration to spruce needle wax , 1995 .

[8]  C. Somerville,et al.  A non-specific lipid transfer protein from Arabidopsis is a cell wall protein. , 1993, The Plant journal : for cell and molecular biology.

[9]  P. Gülz,et al.  Seasonal Variation In The Composition Of Epicuticular Waxes Of Quevcus Robur Leaves , 1992 .

[10]  D. L. Cinti,et al.  The fatty acid chain elongation system of mammalian endoplasmic reticulum. , 1992, Progress in lipid research.

[11]  C. Somerville,et al.  Isolation of a cDNA Clone for Spinach Lipid Transfer Protein and Evidence that the Protein Is Synthesized by the Secretory Pathway. , 1991, Plant physiology.

[12]  R. Kalla,et al.  The promoter of the barley aleurone-specific gene encoding a putative 7 kDa lipid transfer protein confers aleurone cell-specific expression in transgenic rice. , 1994, The Plant journal : for cell and molecular biology.

[13]  F. Schneider,et al.  Biosynthesis of very long chain fatty acids in higher plants. , 1994, Progress in lipid research.

[14]  U. Lundqvist,et al.  Phenotypic diversity of barley spike waxes resulting from mutations at locuscer-n , 1983 .

[15]  J. Bessoule,et al.  Involvement of a β-ketoacyl-CoA intermediate in acyl-CoA elongation by an acyl-CoA elongase purified from leek epidermal cells , 1989 .

[16]  P. Kolattukudy,et al.  The gene for the major cuticular wax-associated protein and three homologous genes from broccoli (Brassica oleracea) and their expression patterns. , 1995, The Plant journal : for cell and molecular biology.

[17]  P. Kolattukudy.  Further evidence for an elongation-decarboxylation mechanism in the biosynthesis of paraffins in leaves. , 1968, Plant physiology.

[18]  C. Cassagne,et al.  Long chain fatty acid CoA-activation by microsomes from Allium porrum epidermal cells. , 1979 .

[19]  J. Mikkelsen.  Structure and biosynthesis of β-diketones in barley spike epicuticular wax , 1979 .

[20]  F. Spener,et al.  A phospholipid transfer protein that binds long‐chain fatty acids , 1985 .

[21]  R. Lessire,et al.  Nature of the Fatty Acid Synthetase Systems in Parenchymal and Epidermal Cells of Allium porrum L. Leaves. , 1983, Plant physiology.

[22]  K. Feldmann,et al.  Leaf Epicuticular Waxes of the Eceriferum Mutants in Arabidopsis , 1995, Plant physiology.

[23]  U. Lundqvist,et al.  Dominant mutations atCer-yy change barley spike wax into leaf blade wax , 1982 .

[24]  P. McCourt,et al.  A mutant of Arabidopsis deficient in c(18:3) and c(16:3) leaf lipids. , 1986, Plant physiology.

[25]  C. Cassagne,et al.  Very long chain fatty acids: Occurrence and biosynthesis in membrane fractions from etiolated maize coleoptiles , 1982 .

[26]  P. Sterk,et al.  Cell-specific expression of the carrot EP2 lipid transfer protein gene. , 1991, The Plant cell.

[27]  E. Cahoon,et al.  Analysis of Glucocerebrosides of Rye (Secale cereale L. cv Puma) Leaf and Plasma Membrane. , 1991, Plant physiology.

[28]  P. Kolattukudy,et al.  Alkane biosynthesis by decarbonylation of aldehydes catalyzed by a particulate preparation from Pisum sativum. , 1984, Proceedings of the National Academy of Sciences of the United States of America.

[29]  B. Søgaard,et al.  Thecer-cqu region in barley: Gene cluster or multifunctional gene , 1980 .

[30]  The genetics of cuticular wax biosynthesis , 1994 .

[31]  B. Svensson,et al.  A 10 kD barley basic protein transfers phosphatidylcholine from liposomes to mitochondria , 1989 .

[32]  F. Salamini,et al.  Glossy mutants of maize , 1979, Heredity.

[33]  J. Jaworski,et al.  Oleoyl‐CoA is not an immediate substrate for fatty acid elongation in developing seeds of Brassica napus , 1995 .

[34]  E. Ashworth,et al.  Involvement of Cork Cells in the Secretion of Epicuticular Wax Filaments on Sorghum bicolor (L.) Moench , 1994, International Journal of Plant Sciences.

[35]  V. P. Agrawal,et al.  Characterization and solubilization of an acyl chain elongation system in microsomes of leek epidermal cells. , 1985, Archives of biochemistry and biophysics.

[36]  K. Evenson,et al.  Fatty Acid-Elongating Activity in Rapidly Expanding Leek Epidermis , 1995, Plant physiology.

[37]  P. Gülz.  Epicuticular Leaf Waxes in the Evolution of the Plant Kingdom , 1994 .

[38]  K. Feldmann,et al.  Changes in epicuticular waxes on wildtype and eceriferum mutants in Arabidopsis during development , 1996 .

[39]  J. Ohlrogge,et al.  Fat metabolism in higher plants. Characterization of plant acyl-ACP and acyl-CoA hydrolases. , 1978, Archives of biochemistry and biophysics.

[40]  A. Zarrabi,et al.  Epicuticular Lipids of Alfalfa Relative to Its Susceptibility to Spotted Alfalfa Aphids (Homoptera: Aphididae) , 1991 .

[41]  Watanabe Shinichiro,et al.  Purification and characterization of a non-specific lipid transfer protein from germinated castor bean endosperms which transfers phospholipids and galactolipids , 1986 .

[42]  A. Culbreath,et al.  Cuticular lipids from wild and cultivated peanuts and the relative resistance of these peanut species to fall armyworm and thrips , 1993 .

[43]  R. McDonald,et al.  Relationship of epicuticular wax composition of grapefruit to chilling injury , 1991 .

[44]  R. W. Davis,et al.  A conditional sterile mutation eliminates surface components from Arabidopsis pollen and disrupts cell signaling during fertilization. , 1993, Genes & development.

[45]  M. Delseny,et al.  Spatial and temporal expression of a maize lipid transfer protein gene. , 1991, The Plant cell.

[46]  J. Mikkelsen,et al.  Biosynthesis of β-diketones and hydrocarbons in barley spike epicuticular wax☆ , 1978 .

[47]  C. Jeffree,et al.  ORIGINS OF THE FINE STRUCTURE OF PLANT EPICUTICULAR WAXES , 1976 .

[48]  J. Kader,et al.  Phospholipid transfer protein from maize seedlings is partly membrane-bound , 1986 .

[49]  M. Yamada.  Lipid Transfer Proteins in Plants and Microorganisms , 1992 .

[50]  G. Bianchi,et al.  Chemical structure of long-chain esters from “sansa” olive oil , 1994 .

[51]  R. Poethig,et al.  Heterochronic effects of glossy15 mutations on epidermal cell identity in maize. , 1994, Development.

[52]  P. Kolattukudy.  Enzymatic synthesis of fatty alcohols in Brassica oleracea. , 1971, Archives of biochemistry and biophysics.

[53]  P. Sterk,et al.  Expression pattern of a tobacco lipid transfer protein gene within the shoot apex. , 1992, The Plant journal : for cell and molecular biology.

[54]  G. J. Blomquist,et al.  Unusual mechanism of hydrocarbon formation in the housefly: cytochrome P450 converts aldehyde to the sex pheromone component (Z)-9-tricosene and CO2. , 1994, Proceedings of the National Academy of Sciences of the United States of America.

[55]  V. P. Agrawal,et al.  Biosynthesis of very long chain fatty acids in microsomes from epidermal cells of Allium porrum L. , 1984, Archives of biochemistry and biophysics.

[56]  P. Kolattukudy,et al.  Biopolyester Membranes of Plants: Cutin and Suberin , 1980, Science.

[57]  G. Bianchi,et al.  Maize pollen lipids , 1990 .

[58]  D. Murphy,et al.  Sub-cellular localization of fatty acid elongase in developing seeds of Lunaria annua and Brassica napus , 1993 .

[59]  G. Bianchi,et al.  Chemosystematics of surface lipids from maize and some related species , 1990 .

[60]  M. Koornneef,et al.  A Genetic and Phenotypic Description of Eceriferum (cer) Mutants in Arabidopsis thaliana , 1989 .

[61]  E. Sutter.  Chemical composition of epicuticular wax in cabbage plants grown in vitro , 1984 .

[62]  H. Matsubara,et al.  A frxC Homolog Exists in the Chloroplast DNAs from Various Pteridophytes and in Gymnosperms , 1992 .

[63]  D. Taylor,et al.  Fatty acid elongation in developing seeds of Arabidopsis thaliana , 1992 .

[64]  F. Salamini,et al.  Absence of long chain aldehydes in the wax of the Glossy II mutant of maize , 1985 .

[65]  J. Ohlrogge,et al.  Acetyl-acyl carrier protein is not a major intermediate in fatty acid biosynthesis in spinach. , 1993, European journal of biochemistry.

[66]  Frans,et al.  Genes Galore: A Summary of Methods for Accessing Results from Large-Scale Partial Sequencing of Anonymous Arabidopsis cDNA Clones , 1994, Plant physiology.

[67]  Kunisuke Tanaka,et al.  Acyl-CoA Synthetase in Maturing Safflower Seeds , 1993 .

[68]  J. Ohlrogge,et al.  Cloning and expression in Escherichia coli of a novel thioesterase from Arabidopsis thaliana specific for long-chain acyl-acyl carrier proteins. , 1995, Archives of biochemistry and biophysics.

[69]  P. Kolattukudy,et al.  Identification of a lipid transfer protein as the major protein in the surface wax of broccoli (Brassica oleracea) leaves. , 1994, Archives of biochemistry and biophysics.

[70]  P. Wettstein-knowles.  Genes, Elongases and Associated Enzyme Systems in Epicuticular Wax Synthesis , 1987 .

[71]  W. Stiekema,et al.  Molecular characterization of the CER1 gene of arabidopsis involved in epicuticular wax biosynthesis and pollen fertility. , 1995, The Plant cell.

[72]  C. Cassagne,et al.  Biosynthesis of saturated very long chain fatty acids by purified membrane fractions from leek epidermal cells. , 1978, Archives of biochemistry and biophysics.

[73]  P. Stumpf,et al.  Isolation and function of spinach leaf beta-ketoacyl-[acyl-carrier-protein] synthases. , 1982, Proceedings of the National Academy of Sciences of the United States of America.

[74]  R. Langhans,et al.  Formation of epicuticular wax and its effect on water loss in cabbage plants regenerated from shoot-tip culture , 1982 .

[75]  E. Ashworth,et al.  Chemically Induced Cuticle Mutation Affecting Epidermal Conductance to Water Vapor and Disease Susceptibility in Sorghum bicolor (L.) Moench , 1994, Plant physiology.

[76]  F. Salamini,et al.  Early event in maize leaf epidermis formation as revealed by cell lineage studies , 1994 .

[77]  A. Hannoufa,et al.  Epicuticular waxes of eceriferum mutants of Arabidopsis thaliana , 1993 .

[78]  P. Kolattukudy.  Cutin, suberin, and waxes , 1980 .

[79]  G. Bianchi,et al.  Composition and structure of maize epicuticular wax esters , 1989 .