植物表皮蜡质合成及运输的研究进展
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摘要
蜡质是覆盖在大多数陆生植物表面,能够保护植物免受生物和非生物胁迫的疏水表层。目前通过突变体分离鉴定和分子生物学等技术,对蜡质的合成及运输途径的研究都有了较大的进展。本文综述了蜡质合成运输中已经研究清楚的途径及相关酶和基因,并提出了存在的问题和展望,以期能够对蜡质的合成和运输有一个全面的了解并对进一步的研究提供指导。
The plant cuticle is a hydrophobic layer that covers the aerial epidermis of most land plants, providing protection against biotic stresses and abiotic stresses. Currently, the research of plant cuticles biosynthesis and export pathway have had a great progress by the technologies of isolation and identification of mutants and molecular biology. Here, we reviewed recent progress of plant cuticles biosynthesis and transport with the related enzymes and genes and highlight some of major questions to have a comprehensive understanding that will drive future research in this field.
The plant cuticle is a hydrophobic layer that covers the aerial epidermis of most land plants, providing protection against biotic stresses and abiotic stresses. Currently, the research of plant cuticles biosynthesis and export pathway have had a great progress by the technologies of isolation and identification of mutants and molecular biology. Here, we reviewed recent progress of plant cuticles biosynthesis and transport with the related enzymes and genes and highlight some of major questions to have a comprehensive understanding that will drive future research in this field.
引文
Bach L,Michaelson LV,Haslam R,Bellec Y,Gissot L,Marion J,Da Costa M,Boutin JP,Miquel M,Tellier F,et al(2008).The verylong-chain hydroxy fatty acyl-Co A dehydratase PASTICCINO2is essential and limiting for plant development.Proc Natl Acad Sci USA,105:14727-14731
Barthlott W,Neinhuis C,Cutler D,Ditsch F,Meusel I,Theisen I,Wilhelmi H(1998).Classification and term inology of plant epicuticular waxes.Bot J Linn Soc,126(3):237-260
Beaudoin F,Wu X,Li F,Haslam RP,Markham JE,Zheng H,Napier JA,Kunst L(2009).Functional characterization of the Arabidopsisβ-ketoacyl-coenzyme A reductase candidates of the fatty acid elongase.Plant Physiol,150:1174-1191
Bernard A,Domergue F,Pascal S,Jetter R,Renne C,Faure JD,Haslam RP,Napier JA,Lessire R,Joubès J(2012).Reconstitution of plant alkane biosynthesis in yeast demonstrates that Arabidopsis ECERIFERUM1 and ECERIFERUM3 are core components of a very-long-chain alkane synthesis complex.Plant Cell,24:3106-3118
Bessire M,Borel S,Fabre G,Carra?a L,Efremova N,Yephremov A,Cao Y,Jetter R,Jacquat AC,Métraux JP,et al(2011).A member of the PLEIOTROPIC DRUG RESISTANCE family of ATPbinding cassette transporters is required for the formation of a functional cuticle in Arabidopsis.Plant Cell,23:1958-1970
Bird D,Beisson F,Brigham A,Shin J,Greer S,Jetter R,Kunst L,Wu X,Yephremov A,Samuels L(2007).Characterization of Arabidopsis ABCG11/WBC11,an ATP binding cassette(ABC)transporter that is required for cuticular lipid secretion.Plant J,52:485-498
Bohnert HJ,Nelson DE,Jensen RJ(1995).Adaptations to environmental stresses.Plant Cell,7(7):1099-1111
Bourdenx B,Bernard A,Domergue F,Pascal S,Léger A,Roby D,Pervent M,Vile D,Haslam RP,Napier JA,et al(2011).Overexpression of Arabidopsis ECERIFERUM1 promotes wax verylong-chain alkane biosynthesis and influences plant response to biotic and abiotic stresses.Plant Physiol,156:29-44
Cheesbrough TM,Kolattukudy PE(1984).Alkane biosynthesis by decarbonylation of aldehydes catalyzed by a particulate preparation from Pisum sativum.Proc Natl Acad Sci USA,81:6613-6617
Debono A,Yeats TH,Rose JKC,Bird D,Jetter R,Kunst L,Samuels L(2009).Arabidopsis LTPG is a glycosylphosphatidylinositol-anchored lipid transfer protein required for export of lipids to the plant surface.Plant Cell,21:1230-1238
Fiebig A,Mayfield JA,Miley NL,Chau S,Fischer RL,Preuss D(2000).Alterations in CER6,a gene identical to CUT1,differentially affect longchain lipid content on the surface of pollen and stems.Plant Cell,12:2001-2008
Girard AL,Mounet F,Lemaire-Chamley M,Gaillard C,Elmorjani K,Vivancos J,Runavot JL,Quemener B,Petit J,Germain V,et al(2012).Tomato GDSL1 is required for cutin deposition in the fruit cuticle.Plant Cell,24:3119-3134
Greer S,Wen M,Bird D,Wu X,Samuels L,Kunst L,Jetter R(2007).The cytochrome P450 enzyme CYP96A15 is the midchain alkane hydroxylase responsible for formation of secondary alcohols and ketones in stem cuticular wax of Arabidopsis.Plant Physiol,145:653-667
Haslam TM,Ma?as-Fernández A,Zhao L,Kunst L(2012).Arabidopsis ECERIFERUM2 is a component of the fatty acid elongation machinery required for fatty acid extension to exceptional lengths.Plant Physiol,160:1164-1174
Isaacson T,Kosma DK,Matas AJ,Buda GJ,He Y,Yu B,Pravitasari A,Batteas JD,Stark RE,Jenks MA,et al(2009).Cutin deficiency in the tomato fruit cuticle consistently affects resistance to microbial infection and biomechanical properties,but not transpirational water loss.Plant J,60:363-377
Jenks MA,Tuttle HA,Eigenbrode SD,Feldmann KA(1995).Leaf epicuticular waxes of the eceriferum mutants in Arabidopsis.Plant Physiol,108:369-377
Jetter R,Sch?ffer S(2001).Chemical composition of the Prunus laurocerasus leaf surface.Dynamic changes of the epicuticular wax film during leaf development.Plant Physiol,126(4):1725Cititower1737
Kim H,Lee SB,Kim HJ,Min MK,Hwang I,Suh MC(2012).Characterization of glycosylphosphatidylinositol-anchored lipid transfer protein 2(LTPG2)and overlapping function between LTPG/LTPG1 and LTPG2 in cuticular wax export or accumulation in Arabidopsis thaliana.Plant Cell Physiol,53:1391-1403
Koornneef M,Hanhart CJ,Thiel F(1989).A genetic and phenotypic description of Eceriferum(cer)mutants in Arabidopsis thaliana.Heredity,80(2):118-122
Kunst L,Samuels AL(2003).Biosynthesis and secretion of plant cuticular wax.Prog Lipid Res,42(1):51-80
Lee SB,Go YS,Bae HJ,Park JH,Cho SH,Cho HJ,Lee DS,Park OK,Hwang I,Suh MC(2009).Disruption of glycosylphosphatidylinositolanchored lipid transfer protein gene altered cuticular lipid composition,increased plastoglobules,and enhanced susceptibility to infection by the fungal pathogen Alternaria brassicicola.Plant Physiol,150:42-54
Leliaert F,Verbruggen H,Zechman FW(2011).Into the deep:new discoveries at the base of the green plant phylogeny.Bioessays,33:683-692
Li F,Wu X,Lam P,Bird D,Zheng H,Samuels L,Jetter R,Kunst L(2008).Identification of the wax ester synthase/acyl-coenzyme A:diacylglycerol acyltransferase WSD1 required for stem wax ester biosynthesis in Arabidopsis.Plant Physiol,148:97-107
Li LZ,Ma J,Xiang JH,Chen XB(2011).Composition differences of epicuticular and intracuticular wax layers and the relationship between cuticle and plant stress tolerance.Plant Physiol J,47(7):680-684(in Chinese with English abstract)[李灵之,马杰,向建华,陈信波(2011).植物角质层内外蜡质的差异及其与抗逆性的关系.植物生理学报,47(7):680-684]
Li-Beisson Y,Pollard M,Sauveplane V,Pinot F,Ohlrogge J,Beisson F(2009).Nanoridges that characterize the surface morphology of flowers require the synthesis of cutin polyester.Proc Natl Acad Sci USA,106:22008-22013
Littlejohn GR,Mansfield GC,Parker D,Lind R,Perfect S,Seymour M,Smirnoff N,Love J,Moger J(2015).In vivo chemical and structural analysis of plant cuticular waxes using stimulated raman scattering microscopy.Plant Physiol,168(1):18-28
Mc Farlane HE,Shin JJ,Bird DA,Samuels AL(2010).Arabidopsis ABCG transporters,which are required for export of diverse cuticular lipids,dimerize in different combinations.Plant Cell,22:3066-3075
Mc Nevin JP,Woodward W,Hannoufa A,Feldmann KA,Lemieux B(1993).Isolation and characterization of eceriferum(cer)mutants induced by T-DNA insertions in Arabidopsis thaliana.Genome,36(3):610-618
Metz JG,Pollard MR,Anderson L,Hayes TR,Lassner MW(2000).Purification of a jojoba embryo fatty acyl-coenzyme A reductase and expression of its c DNA in high erucic acid rapeseed.Plant Physiol,122:635-644
Millar AA,Clemens S,Zachgo S,Giblin ME,Taylor DC,Kunst L(1999).CUT1,an Arabidopsis gene required for cuticular wax biosynthesis and pollen fertility,encodes a very-long-chain fatty acid condensing enzyme.Plant Cell,11:825-838
Ohlrogge J,Browse J(1995).Lipid biosynthesis.Plant Cell,7:957-970
Pascal S,Bernard A,Sorel M,Pervent M,Vile D,Haslam RP,Napier JA,Lessire R,Domergue F,Joubès J(2013).The Arabidopsis cer26 mutant,like the cer2 mutant,is specifically affected in the very long chain fatty acid elongation process.Plant J,73:733-746
Pighin JA,Zheng H,Balakshin LJ,Goodman IP,Western TL,Jetter R,Kunst L,Samuels AL(2004).Plant cuticular lipid export requires an ABC transporter.Science,306:702-704
Preuss D,Lemieux B,Yen G,Davis RW(1993).A conditional sterile mutation eliminates surface components from Arabidopsis pollen and disrupts cell signaling during fertilization.Genes Dev,7:974-985
Pulsifer IP,Kluge S,Rowland O(2012).Arabidopsis long-chain acylCo A synthetase 1(LACS1),LACS2,and LACS3 facilitate fatty acid uptake in yeast.Plant Physiol Bioch,51:31-39
Rowland O,Zheng H,Hepworth SR,Lam P,Jetter R,Kunst L(2006).CER4 encodes an alcohol-forming fatty acyl-Coenzyme A reductase involved in cuticular wax production in Arabidopsis.Plant Physiol,142:866-877
Schneider-Belhaddad F,Kolattukudy P(2000).Solubilization,partial purification,and characterization of a fatty aldehyde decarbonylase from a higher plant,Pisum sativum.Arch Biochem Biophys,377:341-349
Schreiber L,Skrabs M,Hartmann KD,Diamantopoulos P,Simanova E,Santrucek J(2001).Effect of humidity on cuticular water permeability of isolated cuticular membranes and leaf disks.Planta,214:274-282
Song C,Wang J,Guo YW,Ni Y(2011).Response of plant epicuticular wax to environmental factors.Plant Physiol J,47(10):951-956(in Chinese with English abstract)[宋超,王婧,郭彦文,倪郁(2011).植物表皮蜡质对环境胁迫的响应.植物生理学报,47(10):951-956]
Sun W(2007).Studying the functions of biosynthesis and secretion genes of plant culticular wax(Ph D thesis).Ji’nan:Shandong Normal University(in Chinese with English abstract)[孙伟(2007).蜡质合成与转运相关基因的功能研究(博士论文).济南:山东师范大学]
Xu SJ,Jiang PA,Wang ZW,Wang Y(2009).Crystal structures and chemical composition of leaf surface wax depositions on the desert moss Syntrichia caninervis.Biochem Syst Ecol,37:723-730
Yang W,Pollard M,Li-Beisson Y,Beisson F,Feig M,Ohlrogge J(2010).A distinct type of glycerol-3-phosphate acyltransferase with sn-2 preference and phosphatase activity producing2-monoacylglycerol.Proc Natl Acad Sci USA,107:12040-12045
Yeats TH,Martin LB,Viart HM,Isaacson T,He Y,Zhao L,Matas AJ,Buda GJ,Domozych DS,Clausen MH,et al(2012).The identification of cutin synthase:formation of the plant polyester cutin.Nat Chem Biol,8:609-611
Yeats TH,Rose JKC(2008).The biochemistry and biology of extracellular plant lipid-transfer proteins(LTPs).Protein Sci,17:191-198
Yeats TH,Rose JKC(2013).The formation and function of plant cuticles.Plant Physiol,163(1):5-20
Zeng Q,Liu DC,Liu Y(2013).The overview and prospect of chemical composition of plant cuticular wax.Acta Ecolog Sin,33(17):5133-5140(in Chinese with English abstract)[曾琼,刘德春,刘勇(2013).植物角质层蜡质的化学组成研究综述.生态学报,33(17):5133-5140]
Zheng H,Rowland O,Kunst L(2005).Disruptions of the Arabidopsis enoyl-Co A reductase gene reveal an essential role for very-longchain fatty acid synthesis in cell expansion during plant morphogenesis.Plant Cell,17:1467-1481
Barthlott W,Neinhuis C,Cutler D,Ditsch F,Meusel I,Theisen I,Wilhelmi H(1998).Classification and term inology of plant epicuticular waxes.Bot J Linn Soc,126(3):237-260
Beaudoin F,Wu X,Li F,Haslam RP,Markham JE,Zheng H,Napier JA,Kunst L(2009).Functional characterization of the Arabidopsisβ-ketoacyl-coenzyme A reductase candidates of the fatty acid elongase.Plant Physiol,150:1174-1191
Bernard A,Domergue F,Pascal S,Jetter R,Renne C,Faure JD,Haslam RP,Napier JA,Lessire R,Joubès J(2012).Reconstitution of plant alkane biosynthesis in yeast demonstrates that Arabidopsis ECERIFERUM1 and ECERIFERUM3 are core components of a very-long-chain alkane synthesis complex.Plant Cell,24:3106-3118
Bessire M,Borel S,Fabre G,Carra?a L,Efremova N,Yephremov A,Cao Y,Jetter R,Jacquat AC,Métraux JP,et al(2011).A member of the PLEIOTROPIC DRUG RESISTANCE family of ATPbinding cassette transporters is required for the formation of a functional cuticle in Arabidopsis.Plant Cell,23:1958-1970
Bird D,Beisson F,Brigham A,Shin J,Greer S,Jetter R,Kunst L,Wu X,Yephremov A,Samuels L(2007).Characterization of Arabidopsis ABCG11/WBC11,an ATP binding cassette(ABC)transporter that is required for cuticular lipid secretion.Plant J,52:485-498
Bohnert HJ,Nelson DE,Jensen RJ(1995).Adaptations to environmental stresses.Plant Cell,7(7):1099-1111
Bourdenx B,Bernard A,Domergue F,Pascal S,Léger A,Roby D,Pervent M,Vile D,Haslam RP,Napier JA,et al(2011).Overexpression of Arabidopsis ECERIFERUM1 promotes wax verylong-chain alkane biosynthesis and influences plant response to biotic and abiotic stresses.Plant Physiol,156:29-44
Cheesbrough TM,Kolattukudy PE(1984).Alkane biosynthesis by decarbonylation of aldehydes catalyzed by a particulate preparation from Pisum sativum.Proc Natl Acad Sci USA,81:6613-6617
Debono A,Yeats TH,Rose JKC,Bird D,Jetter R,Kunst L,Samuels L(2009).Arabidopsis LTPG is a glycosylphosphatidylinositol-anchored lipid transfer protein required for export of lipids to the plant surface.Plant Cell,21:1230-1238
Fiebig A,Mayfield JA,Miley NL,Chau S,Fischer RL,Preuss D(2000).Alterations in CER6,a gene identical to CUT1,differentially affect longchain lipid content on the surface of pollen and stems.Plant Cell,12:2001-2008
Girard AL,Mounet F,Lemaire-Chamley M,Gaillard C,Elmorjani K,Vivancos J,Runavot JL,Quemener B,Petit J,Germain V,et al(2012).Tomato GDSL1 is required for cutin deposition in the fruit cuticle.Plant Cell,24:3119-3134
Greer S,Wen M,Bird D,Wu X,Samuels L,Kunst L,Jetter R(2007).The cytochrome P450 enzyme CYP96A15 is the midchain alkane hydroxylase responsible for formation of secondary alcohols and ketones in stem cuticular wax of Arabidopsis.Plant Physiol,145:653-667
Haslam TM,Ma?as-Fernández A,Zhao L,Kunst L(2012).Arabidopsis ECERIFERUM2 is a component of the fatty acid elongation machinery required for fatty acid extension to exceptional lengths.Plant Physiol,160:1164-1174
Isaacson T,Kosma DK,Matas AJ,Buda GJ,He Y,Yu B,Pravitasari A,Batteas JD,Stark RE,Jenks MA,et al(2009).Cutin deficiency in the tomato fruit cuticle consistently affects resistance to microbial infection and biomechanical properties,but not transpirational water loss.Plant J,60:363-377
Jenks MA,Tuttle HA,Eigenbrode SD,Feldmann KA(1995).Leaf epicuticular waxes of the eceriferum mutants in Arabidopsis.Plant Physiol,108:369-377
Jetter R,Sch?ffer S(2001).Chemical composition of the Prunus laurocerasus leaf surface.Dynamic changes of the epicuticular wax film during leaf development.Plant Physiol,126(4):1725Cititower1737
Kim H,Lee SB,Kim HJ,Min MK,Hwang I,Suh MC(2012).Characterization of glycosylphosphatidylinositol-anchored lipid transfer protein 2(LTPG2)and overlapping function between LTPG/LTPG1 and LTPG2 in cuticular wax export or accumulation in Arabidopsis thaliana.Plant Cell Physiol,53:1391-1403
Koornneef M,Hanhart CJ,Thiel F(1989).A genetic and phenotypic description of Eceriferum(cer)mutants in Arabidopsis thaliana.Heredity,80(2):118-122
Kunst L,Samuels AL(2003).Biosynthesis and secretion of plant cuticular wax.Prog Lipid Res,42(1):51-80
Lee SB,Go YS,Bae HJ,Park JH,Cho SH,Cho HJ,Lee DS,Park OK,Hwang I,Suh MC(2009).Disruption of glycosylphosphatidylinositolanchored lipid transfer protein gene altered cuticular lipid composition,increased plastoglobules,and enhanced susceptibility to infection by the fungal pathogen Alternaria brassicicola.Plant Physiol,150:42-54
Leliaert F,Verbruggen H,Zechman FW(2011).Into the deep:new discoveries at the base of the green plant phylogeny.Bioessays,33:683-692
Li F,Wu X,Lam P,Bird D,Zheng H,Samuels L,Jetter R,Kunst L(2008).Identification of the wax ester synthase/acyl-coenzyme A:diacylglycerol acyltransferase WSD1 required for stem wax ester biosynthesis in Arabidopsis.Plant Physiol,148:97-107
Li LZ,Ma J,Xiang JH,Chen XB(2011).Composition differences of epicuticular and intracuticular wax layers and the relationship between cuticle and plant stress tolerance.Plant Physiol J,47(7):680-684(in Chinese with English abstract)[李灵之,马杰,向建华,陈信波(2011).植物角质层内外蜡质的差异及其与抗逆性的关系.植物生理学报,47(7):680-684]
Li-Beisson Y,Pollard M,Sauveplane V,Pinot F,Ohlrogge J,Beisson F(2009).Nanoridges that characterize the surface morphology of flowers require the synthesis of cutin polyester.Proc Natl Acad Sci USA,106:22008-22013
Littlejohn GR,Mansfield GC,Parker D,Lind R,Perfect S,Seymour M,Smirnoff N,Love J,Moger J(2015).In vivo chemical and structural analysis of plant cuticular waxes using stimulated raman scattering microscopy.Plant Physiol,168(1):18-28
Mc Farlane HE,Shin JJ,Bird DA,Samuels AL(2010).Arabidopsis ABCG transporters,which are required for export of diverse cuticular lipids,dimerize in different combinations.Plant Cell,22:3066-3075
Mc Nevin JP,Woodward W,Hannoufa A,Feldmann KA,Lemieux B(1993).Isolation and characterization of eceriferum(cer)mutants induced by T-DNA insertions in Arabidopsis thaliana.Genome,36(3):610-618
Metz JG,Pollard MR,Anderson L,Hayes TR,Lassner MW(2000).Purification of a jojoba embryo fatty acyl-coenzyme A reductase and expression of its c DNA in high erucic acid rapeseed.Plant Physiol,122:635-644
Millar AA,Clemens S,Zachgo S,Giblin ME,Taylor DC,Kunst L(1999).CUT1,an Arabidopsis gene required for cuticular wax biosynthesis and pollen fertility,encodes a very-long-chain fatty acid condensing enzyme.Plant Cell,11:825-838
Ohlrogge J,Browse J(1995).Lipid biosynthesis.Plant Cell,7:957-970
Pascal S,Bernard A,Sorel M,Pervent M,Vile D,Haslam RP,Napier JA,Lessire R,Domergue F,Joubès J(2013).The Arabidopsis cer26 mutant,like the cer2 mutant,is specifically affected in the very long chain fatty acid elongation process.Plant J,73:733-746
Pighin JA,Zheng H,Balakshin LJ,Goodman IP,Western TL,Jetter R,Kunst L,Samuels AL(2004).Plant cuticular lipid export requires an ABC transporter.Science,306:702-704
Preuss D,Lemieux B,Yen G,Davis RW(1993).A conditional sterile mutation eliminates surface components from Arabidopsis pollen and disrupts cell signaling during fertilization.Genes Dev,7:974-985
Pulsifer IP,Kluge S,Rowland O(2012).Arabidopsis long-chain acylCo A synthetase 1(LACS1),LACS2,and LACS3 facilitate fatty acid uptake in yeast.Plant Physiol Bioch,51:31-39
Rowland O,Zheng H,Hepworth SR,Lam P,Jetter R,Kunst L(2006).CER4 encodes an alcohol-forming fatty acyl-Coenzyme A reductase involved in cuticular wax production in Arabidopsis.Plant Physiol,142:866-877
Schneider-Belhaddad F,Kolattukudy P(2000).Solubilization,partial purification,and characterization of a fatty aldehyde decarbonylase from a higher plant,Pisum sativum.Arch Biochem Biophys,377:341-349
Schreiber L,Skrabs M,Hartmann KD,Diamantopoulos P,Simanova E,Santrucek J(2001).Effect of humidity on cuticular water permeability of isolated cuticular membranes and leaf disks.Planta,214:274-282
Song C,Wang J,Guo YW,Ni Y(2011).Response of plant epicuticular wax to environmental factors.Plant Physiol J,47(10):951-956(in Chinese with English abstract)[宋超,王婧,郭彦文,倪郁(2011).植物表皮蜡质对环境胁迫的响应.植物生理学报,47(10):951-956]
Sun W(2007).Studying the functions of biosynthesis and secretion genes of plant culticular wax(Ph D thesis).Ji’nan:Shandong Normal University(in Chinese with English abstract)[孙伟(2007).蜡质合成与转运相关基因的功能研究(博士论文).济南:山东师范大学]
Xu SJ,Jiang PA,Wang ZW,Wang Y(2009).Crystal structures and chemical composition of leaf surface wax depositions on the desert moss Syntrichia caninervis.Biochem Syst Ecol,37:723-730
Yang W,Pollard M,Li-Beisson Y,Beisson F,Feig M,Ohlrogge J(2010).A distinct type of glycerol-3-phosphate acyltransferase with sn-2 preference and phosphatase activity producing2-monoacylglycerol.Proc Natl Acad Sci USA,107:12040-12045
Yeats TH,Martin LB,Viart HM,Isaacson T,He Y,Zhao L,Matas AJ,Buda GJ,Domozych DS,Clausen MH,et al(2012).The identification of cutin synthase:formation of the plant polyester cutin.Nat Chem Biol,8:609-611
Yeats TH,Rose JKC(2008).The biochemistry and biology of extracellular plant lipid-transfer proteins(LTPs).Protein Sci,17:191-198
Yeats TH,Rose JKC(2013).The formation and function of plant cuticles.Plant Physiol,163(1):5-20
Zeng Q,Liu DC,Liu Y(2013).The overview and prospect of chemical composition of plant cuticular wax.Acta Ecolog Sin,33(17):5133-5140(in Chinese with English abstract)[曾琼,刘德春,刘勇(2013).植物角质层蜡质的化学组成研究综述.生态学报,33(17):5133-5140]
Zheng H,Rowland O,Kunst L(2005).Disruptions of the Arabidopsis enoyl-Co A reductase gene reveal an essential role for very-longchain fatty acid synthesis in cell expansion during plant morphogenesis.Plant Cell,17:1467-1481
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