类黄酮对油菜脂肪酸积累的影响及其分子机制
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摘要
油菜是世界上最重要的油料作物之一,在我国长江流域有较大的种植面积。菜籽油作为主要食用油之一,在人类生命活动中具有十分重要的作用,再加上全球菜籽油的需求量不断增长,提高油菜籽产油量已成为油菜育种的重要目标。类黄酮作为自然界分布最广泛的次级代谢产物之一,在植物的生长发育过程中扮演着十分重要的角色。类黄酮不仅能够提高植物逆境耐性,还可以调节植物各种生理活动,但是,目前关于类黄酮对种子含油量的影响及种皮中色素积累的进化机制尚不清晰。因此,本文主要综述了种子类黄酮的合成调控、分布、功能及抑制脂肪酸合成的分子机制,以了解类黄酮影响脂肪酸积累的分子机制,为今后通过调控碳源分配来提高种子含油量的研究提供理论基础。
Oilseed rape(Brassica napus L.) is one of the most important oil crops worldwide,and has a large planting area in Yangtze River basin.As one of the four major edible oils,rapeseed oil plays a fundamental role in human life activities.In addition,with the increasing global demand for rapeseed oil,raising oilseed rape production becomes an important direction of rapeseed breeding.As one of the most widely distributed secondary metabolites in nature,flavonoids play a critical part in plant growth and development.Flavonoids can not only improve stress tolerance,but also regulate various physiological activities of plants.However,the role of flavonoids in determining seed oil contents and the evolutionary mechanism of how pigments accumulate in seed coats remain elusive.In this review,we highlighted the metabolism,distribution,and function of flavonoids and its inhibiting effects on fatty acid accumulation in rapeseed.Understanding the molecular mechanism of flavonoids affecting fatty acid accumulation can provide a theoretical basis for increasing oil content of seeds by regulating the allocation of carbon sources in the future.
Oilseed rape(Brassica napus L.) is one of the most important oil crops worldwide,and has a large planting area in Yangtze River basin.As one of the four major edible oils,rapeseed oil plays a fundamental role in human life activities.In addition,with the increasing global demand for rapeseed oil,raising oilseed rape production becomes an important direction of rapeseed breeding.As one of the most widely distributed secondary metabolites in nature,flavonoids play a critical part in plant growth and development.Flavonoids can not only improve stress tolerance,but also regulate various physiological activities of plants.However,the role of flavonoids in determining seed oil contents and the evolutionary mechanism of how pigments accumulate in seed coats remain elusive.In this review,we highlighted the metabolism,distribution,and function of flavonoids and its inhibiting effects on fatty acid accumulation in rapeseed.Understanding the molecular mechanism of flavonoids affecting fatty acid accumulation can provide a theoretical basis for increasing oil content of seeds by regulating the allocation of carbon sources in the future.
引文
[1]国家粮油信息中心.2016年1-12月中国进口食用植物油数据.http://www.grainmarket.com.cn.China National Grain and Oils Information Center.Data of imported edible oil from January to December in 2016.http://www.grainmarket.com.cn.
[2]关周博,田建华,董育红.我国油菜发展的现状、面临的问题以及应对策略.陕西农业科学,2016,62(3):99-101.GUAN Z B,TIAN J H,DONG Y H.Current situation,problems and countermeasures of rapeseed development in China.Shaanxi Journal of Agricultural Sciences,2016,62(3):99-101.(in Chinese)
[3]LIAN J P,LU X C,YIN N W,et al.Silencing of Bn TT1family genes affects seed flavonoid biosynthesis and alters seed fatty acid composition in Brassica napus.Plant Science,2017,254:32-47.
[4]LI N N,XU C C,LI-BEISSON Y H,et al.Fatty acid and lipid transport in plant cells.Trends in Plant Science,2016,21(2):145-158.
[5]BAUD S,BOUTIN J P,MIQUEL M,et al.An integrated overview of seed development in Arabidopsis thaliana ecotype WS.Plant Physiology and Biochemistry,2002,40(2):151-160.
[6]O’NEILL C M,GILL S,HOBBS D,et al.Natural variation for seed oil composition in Arabidopsis thaliana.Phytochemistry,2003,64(6):1077-1090.
[7]KONISHI T,SHINOHARA K,YAMADA K,et al.AcetylCo A carboxylase in higher plants:Most plants other than Gramineae have both the prokaryotic and the eukaryotic forms of this enzyme.Plant and Cell Physiology,1996,37(2):117-122.
[8]OHLROGGE J,BROWSE J.Lipid biosynthesis.Plant Cell,1995,7(7):957-970.
[9]HOBBS D,LU C F,HILLS M J.Cloning of a c DNAencoding diacylglycerol acyltransferase from Arabidopsis thaliana and its functional expression.FEBS Letters,1999,452(3):145-149.
[10]ROUTABOUL J M,BENNING C,BECHTOLD N,et al.The TAG1 locus of Arabidopsis encodes for a diacylglycerol acyltransferase.Plant Physiology and Biochemistry,1999,37(11):831-840.
[11]ZOU J T,WEI Y D,JAKO C,et al.The Arabidopsis thaliana TAG1 mutant has a mutation in a diacylglycerol acyltransferase gene.The Plant Journal,1999,19(6):645-653.
[12]BAUD S,LEPINIEC L.Regulation of de novo fatty acid synthesis in maturing oilseeds of Arabidopsis.Plant Physiology and Biochemistry,2009,47(6):448-455.
[13]IBA K,GIBSON S,NISHIUCHI T,et al.A gene encoding a chloroplast omega-3 fatty acid desaturase complements alterations in fatty acid desaturation and chloroplast copy number of the fad7 mutant of Arabidopsis thaliana.Journal of Biological Chemistry,1993,268(32):24099-24105.
[14]MCCONN M,HUGLY S,BROWSE J,et al.A mutation at the fad8 locus of Arabidopsis identifies a second chloroplastω-3 desaturase.Plant Physiology,1994,106(4):1609-1614.
[15]LI J Y,OULEE T M,RABA R,et al.Arabidopsis flavonoid mutants are hypersensitive to UV-B irradiation.The Plant Cell,1993,5(2):171-179.
[16]TREUTTER D.Significance of flavonoids in plant resistance and enhancement of their biosynthesis.Plant Biology,2010,7(6):581-591.
[17]DEBEAUJON I,LéONKLOOSTERZIEL K M,KOORNNEEFM.Influence of the testa on seed dormancy,germination,and longevity in Arabidopsis.Plant Physiology,2000,122(2):403-414.
[18]MOLA J,GROTEWOLD E,KOESA R.How genes paint flowers and seeds.Trends in Plant Science,1998,3(6):212-217.
[19]BUER C S,MUDAY G K.The transparent testa4 mutation prevents flavonoid synthesis and alters auxin transport and the response of Arabidopsis roots to gravity and light.The Plant Cell,2004,16(5):1191-1205.
[20]ROUTABOUL J M,KERHOAS L,DEBEAUJON I,et al.Flavonoid diversity and biosynthesis in seed of Arabidopsis thaliana.Planta,2006,224(1):96-107.
[21]LEPINIEC L,DEBEAUJON I,ROUTABOUL J M,et al.Genetics and biochemistry of seed flavonoids.Annual Review of Plant Biology,2006,57:405-430.
[22]AUGER B,MARNET N,GAUTIER V,et al.A detailed survey of seed coat flavonoids in developing seeds of Brassica napus L.Journal of Agricultural and Food Chemistry,2010,58(10):6246-6256.
[23]YU C Y.Molecular mechanism of manipulating seed coat coloration in oilseed Brassica species.Journal of Applied Genetics,2013,54(2):135-145.
[24]WANG F L,HE J W,SHI J H,et al.Embryonal control of yellow seed coat locus ECY1 is related to alanine and phenylalanine metabolism in the seed embryo of Brassica napus.G3:Genes,Genomes,Genetics,2016,6(4):1073-1081.
[25]CHEN M X,MAODZEKA A,ZHOU L H,et al.Removal of DELLA repression promotes leaf senescence in Arabidopsis.Plant Science,2014,219/220:26-34.
[26]WANG Z,CHEN M X,CHEN T L,et al.TRANSPARENTTESTA2 regulates embryonic fatty acid biosynthesis by targeting FUSCA3 during the early developmental stage of Arabidopsis seeds.The Plant Journal,2014,77(5):757-769.
[27]CHEN M X,WANG Z,ZHU Y N,et al.The effect of transparent TESTA2 on seed fatty acid biosynthesis and tolerance to environmental stresses during young seedling establishment in Arabidopsis.Plant Physiology,2012,160:1023-1036.
[28]LI C X,ZHANG B,CHEN B,et al.Site-specific phosphorylation of TRANSPARENT TESTA GLABRA1mediates carbon partitioning in Arabidopsis seeds.Nature Communications,2018,9:571.
[29]SHI L,KATAVIC V,YU Y Y,et al.Arabidopsis glabra2mutant seeds deficient in mucilage biosynthesis produce more oil.The Plant Journal,2012,69(1):37-46.
[30]MASHIGUCHI K,TANAKA K,SAKAI T,et al.The main auxin biosynthesis pathway in Arabidopsis.Proceedings of the National Academy of Sciences of the USA,2011,108(45):18512-18517.
[31]BROWN D E,RASHOTTE A M,MURPHY A S,et al.Flavonoids act as negative regulators of auxin transport in vivo in Arabidopsis.Plant Physiology,2001,126:524-535.
[32]WEIJERS D,SAUER M,MEURETTE O,et al.Maintenance of embryonic auxin distribution for apical-basal patterning by PIN-FORMED-dependent auxin transport in Arabidopsis.The Plant Cell,2005,17(9):2517-2526.
[33]PEER W A,BANDYOPADHYAY A,BLAKESLEE J J,et al.Variation in expression and protein localization of the PIN family of auxin efflux facilitator proteins in flavonoid mutants with altered auxin transport in Arabidopsis thaliana.The Plant Cell,2004,16(7):1898-1911.
[34]XUAN L J,HUSSAIN N,WANG Z,et al.Comparison of vitality between seedlings germinated from black-coated and yellow-coated seeds of a turnip rape(Brassica rape L.)subjected to Na Cl and Cd Cl2stresses.Plant Growth Regulation,2015,76(1):61-70.
[2]关周博,田建华,董育红.我国油菜发展的现状、面临的问题以及应对策略.陕西农业科学,2016,62(3):99-101.GUAN Z B,TIAN J H,DONG Y H.Current situation,problems and countermeasures of rapeseed development in China.Shaanxi Journal of Agricultural Sciences,2016,62(3):99-101.(in Chinese)
[3]LIAN J P,LU X C,YIN N W,et al.Silencing of Bn TT1family genes affects seed flavonoid biosynthesis and alters seed fatty acid composition in Brassica napus.Plant Science,2017,254:32-47.
[4]LI N N,XU C C,LI-BEISSON Y H,et al.Fatty acid and lipid transport in plant cells.Trends in Plant Science,2016,21(2):145-158.
[5]BAUD S,BOUTIN J P,MIQUEL M,et al.An integrated overview of seed development in Arabidopsis thaliana ecotype WS.Plant Physiology and Biochemistry,2002,40(2):151-160.
[6]O’NEILL C M,GILL S,HOBBS D,et al.Natural variation for seed oil composition in Arabidopsis thaliana.Phytochemistry,2003,64(6):1077-1090.
[7]KONISHI T,SHINOHARA K,YAMADA K,et al.AcetylCo A carboxylase in higher plants:Most plants other than Gramineae have both the prokaryotic and the eukaryotic forms of this enzyme.Plant and Cell Physiology,1996,37(2):117-122.
[8]OHLROGGE J,BROWSE J.Lipid biosynthesis.Plant Cell,1995,7(7):957-970.
[9]HOBBS D,LU C F,HILLS M J.Cloning of a c DNAencoding diacylglycerol acyltransferase from Arabidopsis thaliana and its functional expression.FEBS Letters,1999,452(3):145-149.
[10]ROUTABOUL J M,BENNING C,BECHTOLD N,et al.The TAG1 locus of Arabidopsis encodes for a diacylglycerol acyltransferase.Plant Physiology and Biochemistry,1999,37(11):831-840.
[11]ZOU J T,WEI Y D,JAKO C,et al.The Arabidopsis thaliana TAG1 mutant has a mutation in a diacylglycerol acyltransferase gene.The Plant Journal,1999,19(6):645-653.
[12]BAUD S,LEPINIEC L.Regulation of de novo fatty acid synthesis in maturing oilseeds of Arabidopsis.Plant Physiology and Biochemistry,2009,47(6):448-455.
[13]IBA K,GIBSON S,NISHIUCHI T,et al.A gene encoding a chloroplast omega-3 fatty acid desaturase complements alterations in fatty acid desaturation and chloroplast copy number of the fad7 mutant of Arabidopsis thaliana.Journal of Biological Chemistry,1993,268(32):24099-24105.
[14]MCCONN M,HUGLY S,BROWSE J,et al.A mutation at the fad8 locus of Arabidopsis identifies a second chloroplastω-3 desaturase.Plant Physiology,1994,106(4):1609-1614.
[15]LI J Y,OULEE T M,RABA R,et al.Arabidopsis flavonoid mutants are hypersensitive to UV-B irradiation.The Plant Cell,1993,5(2):171-179.
[16]TREUTTER D.Significance of flavonoids in plant resistance and enhancement of their biosynthesis.Plant Biology,2010,7(6):581-591.
[17]DEBEAUJON I,LéONKLOOSTERZIEL K M,KOORNNEEFM.Influence of the testa on seed dormancy,germination,and longevity in Arabidopsis.Plant Physiology,2000,122(2):403-414.
[18]MOLA J,GROTEWOLD E,KOESA R.How genes paint flowers and seeds.Trends in Plant Science,1998,3(6):212-217.
[19]BUER C S,MUDAY G K.The transparent testa4 mutation prevents flavonoid synthesis and alters auxin transport and the response of Arabidopsis roots to gravity and light.The Plant Cell,2004,16(5):1191-1205.
[20]ROUTABOUL J M,KERHOAS L,DEBEAUJON I,et al.Flavonoid diversity and biosynthesis in seed of Arabidopsis thaliana.Planta,2006,224(1):96-107.
[21]LEPINIEC L,DEBEAUJON I,ROUTABOUL J M,et al.Genetics and biochemistry of seed flavonoids.Annual Review of Plant Biology,2006,57:405-430.
[22]AUGER B,MARNET N,GAUTIER V,et al.A detailed survey of seed coat flavonoids in developing seeds of Brassica napus L.Journal of Agricultural and Food Chemistry,2010,58(10):6246-6256.
[23]YU C Y.Molecular mechanism of manipulating seed coat coloration in oilseed Brassica species.Journal of Applied Genetics,2013,54(2):135-145.
[24]WANG F L,HE J W,SHI J H,et al.Embryonal control of yellow seed coat locus ECY1 is related to alanine and phenylalanine metabolism in the seed embryo of Brassica napus.G3:Genes,Genomes,Genetics,2016,6(4):1073-1081.
[25]CHEN M X,MAODZEKA A,ZHOU L H,et al.Removal of DELLA repression promotes leaf senescence in Arabidopsis.Plant Science,2014,219/220:26-34.
[26]WANG Z,CHEN M X,CHEN T L,et al.TRANSPARENTTESTA2 regulates embryonic fatty acid biosynthesis by targeting FUSCA3 during the early developmental stage of Arabidopsis seeds.The Plant Journal,2014,77(5):757-769.
[27]CHEN M X,WANG Z,ZHU Y N,et al.The effect of transparent TESTA2 on seed fatty acid biosynthesis and tolerance to environmental stresses during young seedling establishment in Arabidopsis.Plant Physiology,2012,160:1023-1036.
[28]LI C X,ZHANG B,CHEN B,et al.Site-specific phosphorylation of TRANSPARENT TESTA GLABRA1mediates carbon partitioning in Arabidopsis seeds.Nature Communications,2018,9:571.
[29]SHI L,KATAVIC V,YU Y Y,et al.Arabidopsis glabra2mutant seeds deficient in mucilage biosynthesis produce more oil.The Plant Journal,2012,69(1):37-46.
[30]MASHIGUCHI K,TANAKA K,SAKAI T,et al.The main auxin biosynthesis pathway in Arabidopsis.Proceedings of the National Academy of Sciences of the USA,2011,108(45):18512-18517.
[31]BROWN D E,RASHOTTE A M,MURPHY A S,et al.Flavonoids act as negative regulators of auxin transport in vivo in Arabidopsis.Plant Physiology,2001,126:524-535.
[32]WEIJERS D,SAUER M,MEURETTE O,et al.Maintenance of embryonic auxin distribution for apical-basal patterning by PIN-FORMED-dependent auxin transport in Arabidopsis.The Plant Cell,2005,17(9):2517-2526.
[33]PEER W A,BANDYOPADHYAY A,BLAKESLEE J J,et al.Variation in expression and protein localization of the PIN family of auxin efflux facilitator proteins in flavonoid mutants with altered auxin transport in Arabidopsis thaliana.The Plant Cell,2004,16(7):1898-1911.
[34]XUAN L J,HUSSAIN N,WANG Z,et al.Comparison of vitality between seedlings germinated from black-coated and yellow-coated seeds of a turnip rape(Brassica rape L.)subjected to Na Cl and Cd Cl2stresses.Plant Growth Regulation,2015,76(1):61-70.