植物角质层结构组成、生物学功能及分离方法研究进展
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摘要
角质层包括角质和蜡质,是覆盖在花、叶和非木质茎等植物地上部分器官的一层疏水性脂类物质,具有限制植物体内水分过度散失、抵御紫外辐射、防止病虫害入侵等功能,在植物生长发育和适应生物与非生物逆境过程中起到重要保护作用。角质层结构以及化学组成是影响其生物学功能的主要原因,具有十分重要的研究价值。综述了植物角质层的结构组成、生物学功能及其常用分离方法,并对已有研究中存在的问题进行了探讨,以期为今后角质层生理作用及其生物学功能方面研究提供理论参考。
The cuticle membrane consists of cutin and wax. It is a layer of hydrophobic lipid material covering the upper parts of plant,like flower,leaf and non-woody stem organs. It has the functions of limiting water plant excessive loss,resisting ultraviolet radiation,preventing pest invasion,and plays protective roles in the process of plant growth and development,and adaptation to biotic and abiotic stresses. It is very important to study cuticle structure and chemical composition,which are the main reasons influencing its biological function. The paper reviewed the structure composition,biological function and commonly used methods for its separation; discussed problems existing in research,hoping to provide theoretical references for studying its physiological and biological functions in the future.
The cuticle membrane consists of cutin and wax. It is a layer of hydrophobic lipid material covering the upper parts of plant,like flower,leaf and non-woody stem organs. It has the functions of limiting water plant excessive loss,resisting ultraviolet radiation,preventing pest invasion,and plays protective roles in the process of plant growth and development,and adaptation to biotic and abiotic stresses. It is very important to study cuticle structure and chemical composition,which are the main reasons influencing its biological function. The paper reviewed the structure composition,biological function and commonly used methods for its separation; discussed problems existing in research,hoping to provide theoretical references for studying its physiological and biological functions in the future.
引文
[1]Kolattukudy P.Polyesters in higher plants[J].Adv.Biochem.Engin.Biotechnol.,2001,71:1-49.
[2]朱墨.表面活性剂对赤霉素和铁元素在植物表面吸收作用的研究[D].郑州:郑州大学,硕士学位论文,2014.Zhu M.Research on the effect of surfactants on permeation of gibberellin and iron through plant cuticular[D].Zhengzhou:Zhengzhou University,Master Dissertation,2014.
[3]Nawrath C.Unraveling the complex network of cuticular structure and function[J].Curr.Opin.Plant Biol.,2006,9(3):281-287.
[4]Leide J,Hildebrandt U,Vogg G,et al..The positional sterile(ps)mutation affects cuticular transpiration and wax biosynthesis of tomato fruits[J].J.Plant Physiol.,2011,168(9):871-877.
[5]魏雪琴,张长峰,庞杰,等.植物角质层生物学特性及水分渗透性研究进展[J].植物学报,2013,48(5):573-587.Wei X Q,Zhang C F,Pang J,et al..Research progress in the chemistry and structure of plant cuticles related to water permeability[J].Bull.Bot.,2013,48(5):573-587.
[6]张明.碳水化合物及脂肪在黑麦草吸收多环芳烃中的作用[D].杭州:浙江大学,博士学位论文,2009.Zhang M.Effects of carbohydrate and fat on polycyclic aromatic hydrocarbons uptake by ryegrass[D].Hangzhou:Zhejiang University,Doctor Dissertation,2009.
[7]Riederer M,Müller C.Biology of the plant cuticle[J].Syst.Bot.,2006,33(3):613.
[8]Gra9a J,Schreiber L,Rodrigues J,et al..Glycerol and glyceryl esters ofω-hydroxyacids in cutins[J].Phytochemistry,2002,61(2):205-215.
[9]Schreiber L.Review of sorption and diffusion of lipophilic molecules in cuticular waxes and the effects of accelerators on solute mobilities[J].Exp.Bot.,2006,57(11):2515-2523.
[10]曾琼,刘德春,刘勇.植物角质层蜡质的化学组成研究综述[J].生态学报,2013,33(17):5133-5140.Zeng Q,Liu D C,Liu Y.The overview and prospect of chemical composition of plant cuticular wax[J].Acta Ecol.Sin.,2013,33(17):5133-5140.
[11]柴凌燕,董易凡,李士伟,等.植物角质膜及其渗透性与抗旱性研究进展[D].郑州:郑州大学,硕士学位论文,2010.Chai L Y,Dong Y F,Li S W,et al..Research progress in the penetrability and drought resistance of plant cuticular[D].Zhengzhou:Zhengzhou University,Master Dissertation,2010.
[12]Schreiber L,Skrabs M,Hartmann K,et al..Effect of humidity on cuticular water permeability of isolated cuticular membranes and leaf disks[J].Planta,2001,214:274-282.
[13]Sch9nherr J,Luber M.Cuticular penetration of potassium salt:Effects of humidity,anions,and temperature[J].Plant Soil,2001,236:117-122.
[14]王冰.植物角质膜渗透性的研究以及高效表面活性剂的筛选[D].郑州:郑州大学,硕士学位论文,2011.Wang B.Research on the permeability of cuticle and screening for efficient surfactant[D].Zhengzhou:Zhengzhou University,Master Dissertation,2011.
[15]Samuels L,Kunst L,Jetter R.Sealing plant surfaces:Cuticular wax formation by epidermal cells[J].Annu.Rev.Plant Biol.,2008,59(1):683-707.
[16]Li Y H,Beisson F,Koo A J K,et al..Identification of acyltransferases required for cutin biosynthesis and production of cutin with suberin-like monomers[J].Proc.Natl.Acad.Sci.USA,2007,104(46):18339-18344.
[17]Yeats T H,Buda G J,Wang Z,et al..The fruit cuticles of wild tomato species exhibit architectural and chemical diversity,providing a new model for studying the evolution of cuticle function[J].Plant J.,2012,69(4):655-666.
[18]Kunst L,Samuels A.Biosynthesis and secretion of plant cuticular wax[J].Prog.Lipid Res.,2003,42(1):51-80.
[19]Kunst L,Samuels L.Plant cuticles shine:Advances in wax biosynthesis and export[J].Curr.Opin.Plant Biol.,2009,12(6):721-727.
[20]Postbeittenmiller D.Biochemistry and molecular biology of wax production in plants[J].Annu.Rev.Plant Biol.,1996,47(47):405-430.
[21]Daniel J,Sirakova T,Kolattukudy P.An acyl-Co A synthetase in Mycobacterium tuberculosis involved in triacylglycerol accumulation during dormancy[J].PLo S ONE,2014,9(12):e114877.
[22]Lv S,Song T,Kosma D,et al..Arabidopsis CER8 encodes long-chain acyl-Co A synthetase 1(LACS1)that has overlapping functions with LACS2 in plant wax and cutin synthesis[J].Plant J.,2009,59(4):553-564.
[23]Lee S,Jung S,Go Y,et al..Two arabidopsis 3-ketoacyl Co Asynthase genes,KCS20 and KCS2/DAISY,are functionally redundant in cuticular wax and root suberin biosynthesis,but differentially controlled by osmotic stress[J].Plant J.,2009,60(3):462-475.
[24]Wen M,Jetter R.Composition of secondary alcohols,ketones,alkanediols,and ketols in Arabidopsis thaliana cuticular waxes[J].J.Exp.Bot.,2009,60(6):1811.
[25]Domínguez E,Heredia-Guerrero J,Heredia A.The biophysical design of plant cuticles:An overview[J].New Phytol.,2011,189(4):938-949.
[26]Oliveira A,Meirelles S,Salatino A.Epicuticular waxes from caatinga and cerrado species and their efficiency against water loss[J].Anais Acad.Brasileira Ciências,2003,75(4):431-439.
[27]许发喜,刘翠芳,邹杰,等.植物角质层对非生物逆境胁迫响应研究进展[J].中国生物工程杂志,2010,30(8):126-130.Xu F X,Liu C F,Zou J,et al..Research progress in plant cuticle responses to abiotic stresses[J].China Biotechnol.,2010,30(8):126-130.
[28]Cameron K D,Teece M A,Smart L B.Increased accumulation of cuticular wax and expression of lipid transfer protein in response to periodic drying events in leaves of tree tobacco[J].Plant Physiol.,2006,140:176-183.
[29]Kosma D K,Bourdenx B,Bernard A,et al..The impact of water deficiency on leaf cuticle lipids of Arabidopsis[J].Plant Physiol.,2009,151(4):1918-1929.
[30]Islam M,Du H,Ning J,et al..Characterization of glossy1-homologous genes in rice involved in leaf wax accumulation and drought resistance[J].Plant Mol.Biol.,2009,70(4):443-456.
[31]Chen X,Goodwin S,Boroff V,et al..Cloning and characterization of the WAX2 gene of arabidopsis involved in cuticle membrane and wax production[J].Plant Cell,2003,15(5):1170-1185.
[32]Liu R,Chen L,Jia Z,et al..Transcription factor At MYB44regulates induced expression of the ETHYLENE INSENSITIVE2gene in Arabidopsis responding to a harpin protein[J].Mol.Plant Microbe Interact.,2011,24(3):377-389.
[33]Kurdyukov S,Faust A,Nawrath C,et al..The epidermisspecific extracellular bodyguard controls cuticle development and morphogenesis in Arabidopsis[J].Plant Cell,2006,18(2):321-339.
[34]Schnurr J,Shockey J,Browse J.The acyl-Co A synthetase encoded by LACS2 is essential for normal cuticle development in Arabidopsis[J].Plant Cell,2004,16(3):629-642.
[35]Long L,Patelh P,Coryw C,et al..The maize epicuticular wax layer provides UV protection[J].Funct.Plant Biol.,2003,30(1):75-81.
[36]Wang H,Hao J,Chen X,et al..Overexpression of rice WRKY89 enhances ultraviolet B tolerance and disease resistance in rice plants[J].Plant Mol.Biol.,2007,65(6):799-815.
[37]Holmes M,Keiller D.Effects of pubescence and waxes on the reflectance of leaves in the ultraviolet and photosynthetic wavebands:A comparison of a range of species[J].Plant Cell Environ.,2002,25(1):85-93.
[38]Skorska E,Szwarc W.Influence of UV-B radiation on young triticale plants with different wax cover[J].Biol.Plant.,2007,51(1):189-192.
[39]景岚,王丽芳,康俊,等.向日葵品种叶片组织结构与抗锈病的关系[J].植物保护,2009,35(2):81-84.Jing L,Wang L F,Kang J,et al..Leaf tissue structure of sunflower cultivars in relation to their resistance to rust disease[J].Plant Protection,2009,35(2):81-84.
[40]Castillo L,Díaz M,Gonzálezcoloma A,et al..Clytostoma callistegioides(Bignoniaceae)wax extract with activity on aphid settling[J].Phytochemistry,2010,71(17-18):2052-2057.
[41]Sherwood R,Vance C.Resistance to fungal penetration in gramineae[J].Phytopathology,1980,70(4):273-279.
[42]刘勇,陈巨莲,程登发.不同小麦品种(系)叶片表面蜡质对两种麦蚜取食的影响[J].应用生态学报,2007,18(8):1785-1788.Liu Y,Chen J L,Cheng D F.Effects of wheat leaf surface waxes on the feeding of two wheat aphid species[J].Chin.J.Appl.Ecol.,2007,18(8):1785-1788.
[43]Brennan E,Weinbaum S.Effect of epicuticular wax on adhesion of psyllids to glaucous juvenile and glossy adult leaves of Eucalyptus globulus,Labillardière[J].Aust.J.Entomol.,2001,40(3):270-277.
[44]李云桂.典型有机污染物在植物角质层上的吸附行为与跨膜过程[D].杭州:浙江大学,博士学位论文,2011.Li Y G.Sorption behavior and penetration process of typical organic pollutants onto plant cuticles[D].Hangzhou:Zhejiang University,Doctor Dissertation,2011.
[45]昝利.城市环境条件下植物角质膜渗透特征研究[D].西安:西安建筑科技大学,硕士学位论文,2015.Zan L.The study of water permeability of plant under the urban environment[D].Xi'an:Xi'an University of Architecture and Technology,Master Dissertation,2015.
[46]Heredia A.Biophysical and biochemical characteristics of cutin,a plant barrier biopolymer[J].Biochim.Biophys.Acta,2003,1620(1-3):1-7.
[47]Knoche M,Bukovac M.Effect of triton X-100 concentration on NAA penetration through the isolated tomato fruit cuticular membrane[J].Crop.Prot.,2004,23(2):141-146.
[2]朱墨.表面活性剂对赤霉素和铁元素在植物表面吸收作用的研究[D].郑州:郑州大学,硕士学位论文,2014.Zhu M.Research on the effect of surfactants on permeation of gibberellin and iron through plant cuticular[D].Zhengzhou:Zhengzhou University,Master Dissertation,2014.
[3]Nawrath C.Unraveling the complex network of cuticular structure and function[J].Curr.Opin.Plant Biol.,2006,9(3):281-287.
[4]Leide J,Hildebrandt U,Vogg G,et al..The positional sterile(ps)mutation affects cuticular transpiration and wax biosynthesis of tomato fruits[J].J.Plant Physiol.,2011,168(9):871-877.
[5]魏雪琴,张长峰,庞杰,等.植物角质层生物学特性及水分渗透性研究进展[J].植物学报,2013,48(5):573-587.Wei X Q,Zhang C F,Pang J,et al..Research progress in the chemistry and structure of plant cuticles related to water permeability[J].Bull.Bot.,2013,48(5):573-587.
[6]张明.碳水化合物及脂肪在黑麦草吸收多环芳烃中的作用[D].杭州:浙江大学,博士学位论文,2009.Zhang M.Effects of carbohydrate and fat on polycyclic aromatic hydrocarbons uptake by ryegrass[D].Hangzhou:Zhejiang University,Doctor Dissertation,2009.
[7]Riederer M,Müller C.Biology of the plant cuticle[J].Syst.Bot.,2006,33(3):613.
[8]Gra9a J,Schreiber L,Rodrigues J,et al..Glycerol and glyceryl esters ofω-hydroxyacids in cutins[J].Phytochemistry,2002,61(2):205-215.
[9]Schreiber L.Review of sorption and diffusion of lipophilic molecules in cuticular waxes and the effects of accelerators on solute mobilities[J].Exp.Bot.,2006,57(11):2515-2523.
[10]曾琼,刘德春,刘勇.植物角质层蜡质的化学组成研究综述[J].生态学报,2013,33(17):5133-5140.Zeng Q,Liu D C,Liu Y.The overview and prospect of chemical composition of plant cuticular wax[J].Acta Ecol.Sin.,2013,33(17):5133-5140.
[11]柴凌燕,董易凡,李士伟,等.植物角质膜及其渗透性与抗旱性研究进展[D].郑州:郑州大学,硕士学位论文,2010.Chai L Y,Dong Y F,Li S W,et al..Research progress in the penetrability and drought resistance of plant cuticular[D].Zhengzhou:Zhengzhou University,Master Dissertation,2010.
[12]Schreiber L,Skrabs M,Hartmann K,et al..Effect of humidity on cuticular water permeability of isolated cuticular membranes and leaf disks[J].Planta,2001,214:274-282.
[13]Sch9nherr J,Luber M.Cuticular penetration of potassium salt:Effects of humidity,anions,and temperature[J].Plant Soil,2001,236:117-122.
[14]王冰.植物角质膜渗透性的研究以及高效表面活性剂的筛选[D].郑州:郑州大学,硕士学位论文,2011.Wang B.Research on the permeability of cuticle and screening for efficient surfactant[D].Zhengzhou:Zhengzhou University,Master Dissertation,2011.
[15]Samuels L,Kunst L,Jetter R.Sealing plant surfaces:Cuticular wax formation by epidermal cells[J].Annu.Rev.Plant Biol.,2008,59(1):683-707.
[16]Li Y H,Beisson F,Koo A J K,et al..Identification of acyltransferases required for cutin biosynthesis and production of cutin with suberin-like monomers[J].Proc.Natl.Acad.Sci.USA,2007,104(46):18339-18344.
[17]Yeats T H,Buda G J,Wang Z,et al..The fruit cuticles of wild tomato species exhibit architectural and chemical diversity,providing a new model for studying the evolution of cuticle function[J].Plant J.,2012,69(4):655-666.
[18]Kunst L,Samuels A.Biosynthesis and secretion of plant cuticular wax[J].Prog.Lipid Res.,2003,42(1):51-80.
[19]Kunst L,Samuels L.Plant cuticles shine:Advances in wax biosynthesis and export[J].Curr.Opin.Plant Biol.,2009,12(6):721-727.
[20]Postbeittenmiller D.Biochemistry and molecular biology of wax production in plants[J].Annu.Rev.Plant Biol.,1996,47(47):405-430.
[21]Daniel J,Sirakova T,Kolattukudy P.An acyl-Co A synthetase in Mycobacterium tuberculosis involved in triacylglycerol accumulation during dormancy[J].PLo S ONE,2014,9(12):e114877.
[22]Lv S,Song T,Kosma D,et al..Arabidopsis CER8 encodes long-chain acyl-Co A synthetase 1(LACS1)that has overlapping functions with LACS2 in plant wax and cutin synthesis[J].Plant J.,2009,59(4):553-564.
[23]Lee S,Jung S,Go Y,et al..Two arabidopsis 3-ketoacyl Co Asynthase genes,KCS20 and KCS2/DAISY,are functionally redundant in cuticular wax and root suberin biosynthesis,but differentially controlled by osmotic stress[J].Plant J.,2009,60(3):462-475.
[24]Wen M,Jetter R.Composition of secondary alcohols,ketones,alkanediols,and ketols in Arabidopsis thaliana cuticular waxes[J].J.Exp.Bot.,2009,60(6):1811.
[25]Domínguez E,Heredia-Guerrero J,Heredia A.The biophysical design of plant cuticles:An overview[J].New Phytol.,2011,189(4):938-949.
[26]Oliveira A,Meirelles S,Salatino A.Epicuticular waxes from caatinga and cerrado species and their efficiency against water loss[J].Anais Acad.Brasileira Ciências,2003,75(4):431-439.
[27]许发喜,刘翠芳,邹杰,等.植物角质层对非生物逆境胁迫响应研究进展[J].中国生物工程杂志,2010,30(8):126-130.Xu F X,Liu C F,Zou J,et al..Research progress in plant cuticle responses to abiotic stresses[J].China Biotechnol.,2010,30(8):126-130.
[28]Cameron K D,Teece M A,Smart L B.Increased accumulation of cuticular wax and expression of lipid transfer protein in response to periodic drying events in leaves of tree tobacco[J].Plant Physiol.,2006,140:176-183.
[29]Kosma D K,Bourdenx B,Bernard A,et al..The impact of water deficiency on leaf cuticle lipids of Arabidopsis[J].Plant Physiol.,2009,151(4):1918-1929.
[30]Islam M,Du H,Ning J,et al..Characterization of glossy1-homologous genes in rice involved in leaf wax accumulation and drought resistance[J].Plant Mol.Biol.,2009,70(4):443-456.
[31]Chen X,Goodwin S,Boroff V,et al..Cloning and characterization of the WAX2 gene of arabidopsis involved in cuticle membrane and wax production[J].Plant Cell,2003,15(5):1170-1185.
[32]Liu R,Chen L,Jia Z,et al..Transcription factor At MYB44regulates induced expression of the ETHYLENE INSENSITIVE2gene in Arabidopsis responding to a harpin protein[J].Mol.Plant Microbe Interact.,2011,24(3):377-389.
[33]Kurdyukov S,Faust A,Nawrath C,et al..The epidermisspecific extracellular bodyguard controls cuticle development and morphogenesis in Arabidopsis[J].Plant Cell,2006,18(2):321-339.
[34]Schnurr J,Shockey J,Browse J.The acyl-Co A synthetase encoded by LACS2 is essential for normal cuticle development in Arabidopsis[J].Plant Cell,2004,16(3):629-642.
[35]Long L,Patelh P,Coryw C,et al..The maize epicuticular wax layer provides UV protection[J].Funct.Plant Biol.,2003,30(1):75-81.
[36]Wang H,Hao J,Chen X,et al..Overexpression of rice WRKY89 enhances ultraviolet B tolerance and disease resistance in rice plants[J].Plant Mol.Biol.,2007,65(6):799-815.
[37]Holmes M,Keiller D.Effects of pubescence and waxes on the reflectance of leaves in the ultraviolet and photosynthetic wavebands:A comparison of a range of species[J].Plant Cell Environ.,2002,25(1):85-93.
[38]Skorska E,Szwarc W.Influence of UV-B radiation on young triticale plants with different wax cover[J].Biol.Plant.,2007,51(1):189-192.
[39]景岚,王丽芳,康俊,等.向日葵品种叶片组织结构与抗锈病的关系[J].植物保护,2009,35(2):81-84.Jing L,Wang L F,Kang J,et al..Leaf tissue structure of sunflower cultivars in relation to their resistance to rust disease[J].Plant Protection,2009,35(2):81-84.
[40]Castillo L,Díaz M,Gonzálezcoloma A,et al..Clytostoma callistegioides(Bignoniaceae)wax extract with activity on aphid settling[J].Phytochemistry,2010,71(17-18):2052-2057.
[41]Sherwood R,Vance C.Resistance to fungal penetration in gramineae[J].Phytopathology,1980,70(4):273-279.
[42]刘勇,陈巨莲,程登发.不同小麦品种(系)叶片表面蜡质对两种麦蚜取食的影响[J].应用生态学报,2007,18(8):1785-1788.Liu Y,Chen J L,Cheng D F.Effects of wheat leaf surface waxes on the feeding of two wheat aphid species[J].Chin.J.Appl.Ecol.,2007,18(8):1785-1788.
[43]Brennan E,Weinbaum S.Effect of epicuticular wax on adhesion of psyllids to glaucous juvenile and glossy adult leaves of Eucalyptus globulus,Labillardière[J].Aust.J.Entomol.,2001,40(3):270-277.
[44]李云桂.典型有机污染物在植物角质层上的吸附行为与跨膜过程[D].杭州:浙江大学,博士学位论文,2011.Li Y G.Sorption behavior and penetration process of typical organic pollutants onto plant cuticles[D].Hangzhou:Zhejiang University,Doctor Dissertation,2011.
[45]昝利.城市环境条件下植物角质膜渗透特征研究[D].西安:西安建筑科技大学,硕士学位论文,2015.Zan L.The study of water permeability of plant under the urban environment[D].Xi'an:Xi'an University of Architecture and Technology,Master Dissertation,2015.
[46]Heredia A.Biophysical and biochemical characteristics of cutin,a plant barrier biopolymer[J].Biochim.Biophys.Acta,2003,1620(1-3):1-7.
[47]Knoche M,Bukovac M.Effect of triton X-100 concentration on NAA penetration through the isolated tomato fruit cuticular membrane[J].Crop.Prot.,2004,23(2):141-146.