植物油脂合成调控与遗传改良研究进展
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摘要
植物种子油脂储存的主要形式是三酰甘油。在植物生长发育中,脂肪酸和三酰甘油具有重要功能。分析了植物种子油脂合成代谢过程调控研究进展,总结了改良植物种子油脂策略:(1)调节碳源分配,抑制碳源流入蛋白质合成路径的关键酶编码基因的表达、增强碳源流入脂肪酸合成路径的关键酶编码基因的表达;(2)干预油脂合成,促进脂肪酸生物合成,提高三酰甘油组装过程的关键酶编码基因的表达水平;(3)提高种子油脂的品质:改变植物脂肪酸组成,调节脂肪酸脱氢酶基因的表达,引入外源超长链多不饱和脂肪酸合成途径;(4)调控油脂合成代谢途径的转录因子表达,提高种子油脂含量。还讨论了植物油脂合成的三酰甘油前体转运机制及合成途径多基因共同调节合成途径等。
Triglycerides is the main storage form of oil in plant seed. Both fatty acids and triglycerides possessed important functions in the process of plant growth and development. This paper analyzed the research progress in oil regulation and synthesis metabolism process of plant seed; and summarized the strategies for the improvement of plant seed oil:(1) to regulate carbon distribution by inhibiting the expression of genes encoding key enzymes,allocating carbon source into the protein synthesis pathway; and enhancing the expression of key genes encoding key enzymes,leading carbon source into the synthesis pathway of fatty acid;(2) to intervene lipid synthesis by promoting biosynthesis fatty acids,and improving the expression level of key genes encoding enzymes in TAG assembly process;(3) to improve seed oil quality by altering plant fatty acid composition,and regulating the gene expression of fatty acid desaturase; and introducing exogenous synthesis pathway of long chain polyunsaturated fatty acid;(4) to regulate the expression of transcription factors for lipid synthesis metabolism to increase seed oil content. Besides,the paper also discussed the mechanism of TAG precursors synthesis and the co-regulation of multiple genes in the synthesis of plant seed oil.
Triglycerides is the main storage form of oil in plant seed. Both fatty acids and triglycerides possessed important functions in the process of plant growth and development. This paper analyzed the research progress in oil regulation and synthesis metabolism process of plant seed; and summarized the strategies for the improvement of plant seed oil:(1) to regulate carbon distribution by inhibiting the expression of genes encoding key enzymes,allocating carbon source into the protein synthesis pathway; and enhancing the expression of key genes encoding key enzymes,leading carbon source into the synthesis pathway of fatty acid;(2) to intervene lipid synthesis by promoting biosynthesis fatty acids,and improving the expression level of key genes encoding enzymes in TAG assembly process;(3) to improve seed oil quality by altering plant fatty acid composition,and regulating the gene expression of fatty acid desaturase; and introducing exogenous synthesis pathway of long chain polyunsaturated fatty acid;(4) to regulate the expression of transcription factors for lipid synthesis metabolism to increase seed oil content. Besides,the paper also discussed the mechanism of TAG precursors synthesis and the co-regulation of multiple genes in the synthesis of plant seed oil.
引文
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[24]Lotan T,Ohto M,Yee K M,et al..Arabidopsis LEAFY COTYLEDON1 is sufficient to induce embryo development in vegetative cells[J].Cell,1998,93(7):1195-1205.
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[27]Focks N,Benning C.Wrinkled1:A novel,low seed oil mutant of Arabidopsis with efficiency in the seed-specific regulation of carbohydrate metabolism[J].Plant Physiol.,1998,118(1):91-101.
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[30]Mcglew K,Shaw V,Zhang M,et al..An annotated database of Arabidopsis mutants of acyl lipid metabolism[J].Plant Cell Rep.,2015,34(4):519-532.
[31]Marillia E F,Micallef B J,Micallef M,et al..Biochemical and physiological studies of Arabidopsis thaliana transgenic lines with repressed expression of the mitochondrial pyruvate dehydrogenase kinase[J].J.Exp.Bot.,2003,54(381):259-270.
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[33]Pidkowitch M S,Nguyen H T,Heilmann I,et al..Modulating seed-ketoacyl-acyl carrier protein synthase II level converts the composition of a temperate seed oil to that of a palm-like tropical oil[J].Proc.Natl.Acad.Sci.USA,2007,104(11):4742-4747.
[34]Salas J J,Ohlrogge J B.Characterization of substrate specicity of plant Fat A and Fat B acyl-ACP thioesterases[J].Arch.Biochem.Biophys.,2002,403(1):25-34.
[35]Kumar S,Dhingra A,Daniell H.Stable transformation of the cotton plastid genome and maternal inheritance of transgenes[J].Plant Mol.Biol.,2004,56(2):203-216.
[36]Madoka Y,Tomizawa K I,Miozoi J,et al..Chloroplast transformation with modied acc D operon increases acetyl-Co A carboxylase and causes extension of leaf longevity and increase in seed yield in tobacco[J].Plant Cell Physiol.,2002,43(12):1518-1525.
[37]Shockey J,Regmi A,Cotton K,et al..Identification of Arabidopsis GPAT9(At5g60620)as an essential gene involved in triacylglycerol biosynthesis[J].Plant Physiol.,2016,170(1):163-179.
[38]Hernández M L,Whitehead L,He Z S,et al..A cytosolic acyltransferase contributes to triacylglycerol synthesis in sucrose-rescued Arabidopsis seed oil catabolism mutants[J].Plant Physiol.,2012,160(1):215-225.
[39]Dahlqvist A,Stahl U,Lenman M,et al..Phospholipid:Diacylglycerol acyltransferase:An enzyme that catalyzes the acyl-Co A-independent formation of triacylglycerol in yeast and plants[J].Proc.Natl.Acad.Sci.USA,2000,97(12):6487-6492.
[40]Stahl U,Carlsson A S,Lenman M,et al..Cloning and functional characterization of a phospholipid:Diacylglycerol acyltransferase from Arabidopsis[J].Plant Physiol.,2004,135(3):1324-1335.
[41]Hovav R,Faigenboim-Doron A,Kadmon N,et al..A transcriptome profile for developing seed of polyploid cotton[J].Plant Genome,2015,8(1):1-15.
[42]Wang L H,Yu S,Tong C B,et al..Genome sequencing of the high oil crop sesame provides insight into oil biosynthesis[J].Genome Biol.,2014,15(2):1-13.
[43]Bates P D,Ohlrogge J B,Pollard M.Incorporation of newly synthesized fatty acids into cytosolic glycerolipids in pea leaves occurs via acyl editing[J].J.Biol.Chem.,2007,282(43):31206-31216.
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[2]Libeisson Y,Shorrosh B,Beisson F,et al..Acyl-lipid metabolism[J].Arabidopsis Book,2010,8(8):e0133.
[3]Simcox P D,Reid E E,Canvin D T,et al..Enzymes of the glycolytic and pentose phosphate pathways in proplastids from the developing endosperm of Ricinus communis L.[J].Plant Physiol.,1977,59(6):1128-1132.
[4]王镜岩,朱圣庚,徐长法.生物化学[M].北京:高等教育出版社,2004,265.Wang J Y,Zhu S G,Xu C F.Biochemistry[M].Beijing:Higher Education Press,2004,265.
[5]陈锦清,郎春秀,胡张华,等.反义PEP基因调控油菜籽粒蛋白质/油脂含量比率的研究[J].农业生物技术学报,1999,7(4):316-320.Chen J Q,Lang C X,Hu Z H,et al..Antisense PEP gene regulates to ratio of protein and lipid content in Brassica napus seeds[J].J.Agric.Biotechnol.,1999,7(4):316-320.
[6]Li X R,Wang L,Ruan Y L.Developmental and molecular physiological evidence for the role of phosphoenolpyruvate carboxylase in rapid cotton fibre elongation[J].J.Exp.Bot.,2010,61(1):287-295.
[7]Xu Z P,Li J W,Guo X P,et al..Metabolic engineering of cottonseed oil biosynthesis pathway via RNA interference[J].Sci.Rep.,2016,6:1-14.
[8]Williams M,Randall D D.Pyruvate dehydrogenase complex from chloroplasts of Pisum sativum L.[J].Plant Physiol.,1979,64(6):1099-1103.
[9]Yui R,Iketani S,Mikami T,et al..Antisense inhibition of mitochondrial pyruvate dehydrogenase E1a subunit in anther tapetum causes male sterility[J].Plant J.,2003,34(1):57-66.
[10]Gengenbach B G,Somers D A,Wyse D L,et al..Methods and an acetyl-Co A carboxylase gene for conferring herbicide tolerance and an alteration in oil content of plants[P].USA:US6069298.
[11]Cui Y P,Liu Z J,Zhao Y P,et al..Overexpression of heteromeric Gh ACCase subunits enhanced oil accumulation in upland cotton[J].Plant Mol.Bio.Rep.,2017,35(2):287-297.
[12]Voellcer T A,Hayes T R,Cranmer A C,et al..Genetic engineering of a quantitative trait:Metabolic and genetic parameters influencing the accumulation of laurate in rapeseed[J].Plant J.,1996,9(2):229-241.
[13]Roesler K,Shintani D,Savage L,et al..Targeting of the Arabidopsis homomeric acetyl-coenzyme A carboxylase to plastids of rapeseeds[J].Plant Physiol.,1997,113(1):75-81.
[14]Stoll C,Lühs W,Zarhloul M K,et al..Knockout of KASIII regulation changes fatty acid composition in canola(Brassica napus L.)[J].Eur.J.Lipid Sci.Technol.,2006,108(4):277-286.
[15]Wei Q,Li J,Zhang L,et al..Cloning and characterization of aβ-ketoacyl-acyl carrier protein synthaseⅡfrom Jatropha curcas[J].J.Plant Physiol.,2012,169(8):816-824.
[16]Bourgis F,Kilaru A,Cao X,et al..Comparative transcriptome and metabolite analysis of oil palm and date palm mesocarp that differ dramatically in carbon partitioning[J].Proc.Natl.Acad.Sci.USA,2011,108(30):12527-12532.
[17]Chi X Y,Yang Q L,Pan L J,et al..Cloning and expression analysis ofβ-ketoacyl-ACP reductase,β-hydroxyacyl-ACP dehydrase and enoyl-ACP reductase from Arachis hypogaea L.[J].Genomics App.Biol.,2010,2(1):1-9.
[18]刘丽,王玉美,赵彦朋,等.棉花脂肪酸合成酶基因Gh KAR和Gh ENR表达载体构建及其功能初探[J].棉花学报,2016,28(6):527-537.Liu L,Wang Y M,Zhao Y P,et al..Construction of expression vectors and a preliminarily functional analysis of fatty acid synthetase genes of Gh KAR and Gh ENR in upland cotton[J].Cotton Sci.,2016,28(6):527-537.
[19]Jain R K,Coffey M,Lai K,et al..Enhancement of seed oil content by expression of glycerol-3-phosphate acyltransferase genes[J].Biochem.Soc.Trans.,2000,28(6):958-961.
[20]Jako C,Kumar A,Wei Y D,et al..Seed-specific overexpression of an Arabidopsis c DNA encoding a diacylglycerel acyltransferase enhances seed oil content and seed weight[J].Plant Physiol.,2001,126(2):861-874.
[21]刘正杰,张园,王玉美,等.陆地棉Gh DGAT1基因干涉载体构建与遗传转化[J].中国农业大学学报,2013,18(5):1-8.Liu Z J,Zhang Y,Wang Y M,et al..Construction and transformation of RNAi vector of gene Gh DGAT1 in upland cotton[J].J.Chin.Agric.Univ.,2013,18(5):1-8.
[22]Lardizabal K,Effertz R,Levering C,et al..Expression of Umbelopsis ramanniana DGAT2A in seed increases oil in soybean[J].Plant physiol.,2008,148(1):89-96.
[23]Oakes J,Brackenridge D,Colletti R,et al..Expression of fungal diacylglycerol acyltransferase2 genes to increase kernel oil in maize[J].Plant Physiol.,2011,155(3):1146-1157.
[24]Lotan T,Ohto M,Yee K M,et al..Arabidopsis LEAFY COTYLEDON1 is sufficient to induce embryo development in vegetative cells[J].Cell,1998,93(7):1195-1205.
[25]Angeles-Núnez J G,Tiessen A.Mutation of the transcription factor LEAFY COTYLEDON2 alters the chemical composition of Arabidopsis seeds,decreasing oil and protein content,while maintaining high levels of starch and sucrose in mature seeds[J].J.Plant Physiol.,2011,168(16):1891-1900.
[26]Shen B,Allen W B,Zheng P,et al..Expression of Zm LEC1and Zm WRI1 increases seed oil production in maize[J].Plant Physiol.,2010,153(3):980-987.
[27]Focks N,Benning C.Wrinkled1:A novel,low seed oil mutant of Arabidopsis with efficiency in the seed-specific regulation of carbohydrate metabolism[J].Plant Physiol.,1998,118(1):91-101.
[28]Yanagisawa S,Izui K.Molecular cloning of two DNA-binding proteins of maize that are structurally different but interact with the same sequence motif[J].J.Biol.Chem.,1993,268(21):16028-16036.
[29]Wang H W,Zhang B,Hao Y J,et al..The soybean Dof-type transcription factor genes,Gm Dof4 and Gm Dof11,enhance lipid content in the seeds of transgenic Arabidopsis plants[J].Plant J.,2007,52(4):716-729.
[30]Mcglew K,Shaw V,Zhang M,et al..An annotated database of Arabidopsis mutants of acyl lipid metabolism[J].Plant Cell Rep.,2015,34(4):519-532.
[31]Marillia E F,Micallef B J,Micallef M,et al..Biochemical and physiological studies of Arabidopsis thaliana transgenic lines with repressed expression of the mitochondrial pyruvate dehydrogenase kinase[J].J.Exp.Bot.,2003,54(381):259-270.
[32]Vigeolas H,Waldeck P,Zank T,et al..Increasing seed oil content in oil-seed rape(Brassica napus L.)by over-expression of a yeast glycerol-3-phosphate dehydrogenase under the control of a seed-specic promoter[J].Plant Biotechnol.J.,2007,5(3):431-441.
[33]Pidkowitch M S,Nguyen H T,Heilmann I,et al..Modulating seed-ketoacyl-acyl carrier protein synthase II level converts the composition of a temperate seed oil to that of a palm-like tropical oil[J].Proc.Natl.Acad.Sci.USA,2007,104(11):4742-4747.
[34]Salas J J,Ohlrogge J B.Characterization of substrate specicity of plant Fat A and Fat B acyl-ACP thioesterases[J].Arch.Biochem.Biophys.,2002,403(1):25-34.
[35]Kumar S,Dhingra A,Daniell H.Stable transformation of the cotton plastid genome and maternal inheritance of transgenes[J].Plant Mol.Biol.,2004,56(2):203-216.
[36]Madoka Y,Tomizawa K I,Miozoi J,et al..Chloroplast transformation with modied acc D operon increases acetyl-Co A carboxylase and causes extension of leaf longevity and increase in seed yield in tobacco[J].Plant Cell Physiol.,2002,43(12):1518-1525.
[37]Shockey J,Regmi A,Cotton K,et al..Identification of Arabidopsis GPAT9(At5g60620)as an essential gene involved in triacylglycerol biosynthesis[J].Plant Physiol.,2016,170(1):163-179.
[38]Hernández M L,Whitehead L,He Z S,et al..A cytosolic acyltransferase contributes to triacylglycerol synthesis in sucrose-rescued Arabidopsis seed oil catabolism mutants[J].Plant Physiol.,2012,160(1):215-225.
[39]Dahlqvist A,Stahl U,Lenman M,et al..Phospholipid:Diacylglycerol acyltransferase:An enzyme that catalyzes the acyl-Co A-independent formation of triacylglycerol in yeast and plants[J].Proc.Natl.Acad.Sci.USA,2000,97(12):6487-6492.
[40]Stahl U,Carlsson A S,Lenman M,et al..Cloning and functional characterization of a phospholipid:Diacylglycerol acyltransferase from Arabidopsis[J].Plant Physiol.,2004,135(3):1324-1335.
[41]Hovav R,Faigenboim-Doron A,Kadmon N,et al..A transcriptome profile for developing seed of polyploid cotton[J].Plant Genome,2015,8(1):1-15.
[42]Wang L H,Yu S,Tong C B,et al..Genome sequencing of the high oil crop sesame provides insight into oil biosynthesis[J].Genome Biol.,2014,15(2):1-13.
[43]Bates P D,Ohlrogge J B,Pollard M.Incorporation of newly synthesized fatty acids into cytosolic glycerolipids in pea leaves occurs via acyl editing[J].J.Biol.Chem.,2007,282(43):31206-31216.
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