微藻油脂合成代谢途径及调控机制的研究进展
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摘要
随着经济的快速发展,各国对石油的需求仍有增无减,但是石油作为不可再生能源不利于可持续发展。相比之下,生物柴油应运而生且已发展到了第三代,而微藻作为第三代生物柴油的主角因为具有生长速度快,不占用耕地等优势,已逐渐成为具有极大发展潜力的能源原料,所以利用微藻生产生物柴油的技术近年来也成为研究的重点。本综述针对微藻容易进行基因改造的优势,综述了国内外对微藻油脂代谢通路的研究现状及进展,讨论了目前的基因改造方法对通路中关键酶产生的影响,以求找到能有效并稳定增强微藻油脂代谢途径的方法,为以后的研究提供理论指导。
With the rapid development of economy, the demand for petroleum in various countries is still on the rise, but as non-renewable energy, it is not good for sustainable development. In contrast, biodiesel emerges at the right moment and has developed to the third generation. Microalgae served as a leading role in the third generation of biodiesel has gradually become the energy raw materials with great development potential because of its advantages such as fast growth and non-occupation of arable land, etc. Therefore, it has become a research focus to utilize microalgae to product biodiesel in recent years. Since microalgae could be genetically modified easily, this paper reviewed the current situation and research advances of microalgae lipid metabolic pathway at home and abroad. The effect of gene modification methods at present on key enzyme in the pathway were discussed in order to find the effective and stable method for enhancing microalgae lipid metabolic pathway and to lay a theoretical foundation for further researches.
With the rapid development of economy, the demand for petroleum in various countries is still on the rise, but as non-renewable energy, it is not good for sustainable development. In contrast, biodiesel emerges at the right moment and has developed to the third generation. Microalgae served as a leading role in the third generation of biodiesel has gradually become the energy raw materials with great development potential because of its advantages such as fast growth and non-occupation of arable land, etc. Therefore, it has become a research focus to utilize microalgae to product biodiesel in recent years. Since microalgae could be genetically modified easily, this paper reviewed the current situation and research advances of microalgae lipid metabolic pathway at home and abroad. The effect of gene modification methods at present on key enzyme in the pathway were discussed in order to find the effective and stable method for enhancing microalgae lipid metabolic pathway and to lay a theoretical foundation for further researches.
引文
Adrio J.L.,2017,Oleaginous yeasts:promising platforms for the production of oleochemicals and biofuels,Biotechnol.Bioeng.,114(9):1915-1920
Bagnato C.,Prados M.B.,Franchini G.R.,Scaglia N.,Miranda S.E.,and Beligni M.V.,2017,Analysis of triglyceride synthesis unveils a green algal soluble diacylglycerol acyltransferase and provides clues to potential enzymatic components of the chloroplast pathway,BMC Genomics,18(1):223
Bharathiraja B.,Jayamuthunagai J.,Chakravarthy M.,Kumar R.R.,Yogendran D.,and Praveenkumar R.,2015,Algae:Promising future feedstock for biofuels,In:Singh B.,Bauddh K.,and Bux F.(eds.),Algae and environmental sustainability,Springer-Verlag,Berlin,Germany,pp.1-8
Brembu T.,Mühlroth A.,Alipanah L.,and Bones A.M.,2017,The effects of phosphorus limitation on carbon metabolism in diatoms,Philos.Trans.R.Soc.Lond.B.Biol.Sci.,372(1728):20160406
Chen J.E.,and Smith A.G.,2012,A look at diacylglycerol acyltransferases(DGATs)in algae,J.Biotechnol.,162(1):28-39
Dahlqvist A.,Stahl U.,Lenman M.,Banas A.,Lee M.,Sandager L.,Ronne H.,and Stymne S.,2000,Phospholipid:diacylglycerol acyltransferase:an enzyme that catalyzes the acyl-coa-independent formation of triacylglycerol in yeast and plants,Proc.Natl.Acad.Sci.USA,97(12):6487-6492
Deng X.D.,Gu B.,Li Y.J.,Hu X.W.,Guo J.C.,and Fei X.W.,2012,The roles of acyl-coa:diacylglycerol acyltransferase 2genes in the biosynthesis of triacylglycerols by the green algae Chlamydomonas reinhardtii,Mol.Plant,5(4):945-947
Dunahay T.G.,Jarvis E.E.,Dais S.S.,and Roessler P.G.,1996,Manipulation of microalgal lipid production using genetic engineering,Appl.Biochem.Biotechnol.,57-58(1):223-231
Goodwin S.,Mcpherson J.D.,and Mccombie W.R.,2016,Coming of age:ten years of next-generation sequencing technologies,Nat.Rev.Genet.,17(6):333-351
Gorain P.C.,Bagchi S.K.,and Mallick N.,2013,Effects of calcium,magnesium and sodium chloride in enhancing lipid accumulation in two green microalgae,Environ.Technol.,34(13-16):1887-1894
Hern觔N.A.,Zapata-Morales A.L.,Ochoa Alfaro A.E.,and Soria-Guerra R.E.,2016,Identification of gene transcripts involved in lipid biosynthesis in Chlamydomonas reinhardtii under nitrogen,iron and sulfur deprivation,World J.Microbiol.Biotechnol.,32(4):1-10
Hu Q.,Sommerfeld M.,Jarvis E.,Ghirardi M.,Posewitz M.,Seibert,M.,and Darzins A.,2008,Microalgal triacylglycerols as feedstocks for biofuel production:perspectives and advances,Plant J.,54(4):621-639
Hung C.H.,Kanehara K.,and Nakamura Y.,2016,Isolation and characterization of a mutant defective in triacylglycerol accumulation in nitrogen-starved Chlamydomonas reinhardtii,Biochim.Biophys.Acta,1861(9):1282-1293
Jaeger L.D.,Verbeek R.E.,Draaisma R.B.,Martens D.E.,Springer J.,Eggink G.,and Wijffels R.H.,2014,Superior triacylglycerol(TAG)accumulation in starchless mutants of scenedesmus obliquus:(i)mutant generation and characterization,Biotechnol.Biofuels.,7(1):69
Kan G.F.,Shi C.G.,Wang X.F.,Xie Q.J.,Wang M.,Wang X.L.,and Miao J.L.,2012,Acclimatory responses to high-salt stress in Chlamydomonas(Chlorophyta,Chlorophyceae)from Antarctica,Acta Oceanol.Sin.,31(1):116-124
Kim H.U.,Lee K.R.,Go Y.S.,Jung J.H.,Suh M.C.,and Kim J.B.,2011,Endoplasmic reticulum-located pdat1-2 from castor bean enhances hydroxy fatty acid accumulation in transgenic plants,Plant Cell Physiol.,52(6):983-993
Klaus D.,Ohlrogge J.B.,Neuhaus H.E.,and D觟rmann P.,2004,Increased fatty acid production in potato by engineering of acetyl-coa carboxylase,Planta,219(3):389-396
Klok A.J.,Martens D.E.,Wijffels R.H.,and Lamers P.P.,2013,Simultaneous growth and neutral lipid accumulation in microalgae,Bioresour.Technol.,134:233-243
Lehner R.,and Kuksis A.,1993,Triacylglycerol synthesis by an sn-1,2(2,3)-diacylglycerol transacylase from rat intestinal microsomes,J.Biol.Chem.,268(12):8781-8186
Li Y.,Han D.,Hu G.,Sommerfeld M.,and Hu Q.,2010,Inhibition of starch synthesis results in overproduction of lipids in Chlamydomonas reinhardtii,Biotechnol.Bioeng.,107(2):258-268
Li-Beisson Y.,Beisson F.,and Riekhof W.,2015,Metabolism of acyl-lipids in chlamydomonas reinhardtii,Plant J.,82(3):504
Liu B.,and Benning C.,2013,Lipid metabolism in microalgae distinguishes itself,Curr.Opin.Biotechnol.,24(2):300-309
Lv H.,Qu G.,Qi X.,Lu L.,Tian C.,and Ma Y.,2013,Transcriptome analysis of Chlamydomonas reinhardtii during the process of lipid accumulation,Genomics,101(4):229-237
Maisonneuve S.,Bessoule J.J.,Lessire R.,Delseny M.,and Roscoe T.J.,2010,Expression of rapeseed microsomal lysophosphatidic acid acyltransferase isozymes enhances seed oil content in arabidopsis,Plant Physiol.,152(2):670-684
Michelon W.,Da S.M.,Mezzari M.P.,Pirolli M.,Prandini J.M.,and Soares H.M.,2015,Effects of nitrogen and phosphorus on biochemical composition of microalgae polyculture harvested from phycoremediation of piggery wastewater digestate,Appl.Biochem.Biotechnol.,178(7):1-13
Miller R.,Wu G.X.,Deshpande R.R.,Vieler A.,G覿rtner K.,Li X.B.,Moellering E.R.,Z覿uner S.,Cornish A.J.,Liu B.,Bullard B.,Sears B.B.,Kuo M.H.,Hegg E.L.,Shachar-Hill Y.,Shiu S.H.,and Benning C.,2010,Changes in transcript abundance in Chlamydomonas reinhardtii following nitrogen deprivation predict diversion of metabolism,Plant Physiol.,154(4):1737-1752
Msanne J.,Xu D.,Kond a A.R.,Casasmollano J.A.,Awada T.,Cahoon E.B.,Cerutti H.,2012,Metabolic and gene expression changes triggered by nitrogen deprivation in the pho toautotrophically grown microalgae Chlamydomonas reinhardtii and coccomyxa sp.c-169,Phytochemistry.,75:50-59
Mujtaba G.,Choi W.,Lee C.G.,and Lee K.,2012,Lipid production by chlorella vulgaris after a shift from nutrient-rich to nitrogen starvation conditions,Bioresour.Technol.,123(123):279-283
Ota S.,Oshima K.,Yamazaki T.,Kim S.,Yu Z.,Yoshihara M.,Takeda K.,Takeshita T.,Hirata A.,Bi觢ováK.,Zachleder V.,Hattori M.,and Kawano S.,2016,Highly efficient lipid production in the green alga parachlorella kessleri:draft genome and transcriptome endorsed by whole-cell 3d ultrastructure,Biotechnol.Biofuels.,9(1):13
Pan X.,Siloto R.M.P.,Wickramarathna A.D.,Mietkiewska E.,and Weselake R.J.,2013,Identification of a pair of phospholipid:diacylglycerol acyltransferases from developing flax(Linum usitatissimum L.)seed catalyzing the selective production of trilinolenin,J.Biol.Chem.,288(33):24173-24188
Pandit P.R.,Fulekar M.H.,and Karuna M.S.,2017,Effect of salinity stress on growth,lipid productivity,fatty acid composition,and biodiesel properties in acutodesmus obliquus and chlorella vulgaris,Environ.Sci.Pollut.Res.Int.,24(15):13437-13451
Radakovits R.,Jinkerson R.E.,Darzins A.,and Posewitz M.C.,2010,Genetic engineering of algae for enhanced biofuel production,Eukaryot.Cell.,9(4):486-501
Rani S.H.,Krishna T.H.,Saha S.,Negi A.S.,and Rajasekharan R.,2010,Defective in cuticular ridges(DCR)of Arabidopsis thaliana,a gene associated with surface cutin formation,encodes a soluble diacylglycerol acyltransferase,J.Biol.Chem.,285(49):38337-38347
Riekhof W.R.,Sears B.B.,and Benning C.,2005,Annotation of genes involved in glycerolipid biosynthesis in Chlamydomonas reinhardtii:discovery of the betaine lipid synthase bta1cr,Eukaryot.Cell,4(2):242-252
Roopnarain A.,Gray V.M.,and Sym S.D.,2014,Phosphorus limitation and starvation effects on cell growth and lipid accumulation in isochrysis galbana u4 for biodiesel production,Bioresour.Technol.,156(2):408-411
Saha S.,Enugutti B.,Rajakumari S.,and Rajasekharan R.,2006,Cytosolic triacylglycerol biosynthetic pathway in oilseeds molecular cloning and expression of peanut cytosolic diacylglycerol acyltransferase,Plant Physiol.,141(4):1533-1543
Sasaki Y.,and Nagano Y.,2004,Plant acetyl-coa carboxylase:structure,biosynthesis,regulation,and gene manipulation for plant breeding,Biosci.Biotechnol.Biochem.,68(6):1175
Sato A.,Matsumura R.,Hoshino N.,Tsuzuki M.,and Sato N.,2014,Responsibility of regulatory gene expression and repressed protein synthesis for triacylglycerol accumulation on sulfur-starvation in Chlamydomonas reinhardtii,Front.Plant Sci.,5:444
Schuler L.M.,Schulze P.S.C.,Pereira H.,Barreira L.,Leon R.,and Varela J.,2017,Trends and strategies to enhance triacylglycerols and high-value compounds in microalgae,Algal Res.,25:263-273
Sharma K.K.,Schuhmann H.,and Schenk P.M.,2012,High lipid induction in microalgae for biodiesel production,Energies.,5(5):1532-1553
Stahl U.,Carlsson A.S.,Lenman M.,Dahlqvist A.,Huang B.,Banas W.,Banas A.,and Stymne S.,2004,Cloning and functional characterization of a phospholipid:diacylglycerol acyltransferase from Arabidopsis,Plant Physiol.,135(3):1324-1335
Stobart K.,Mancha M.,Lenman M.,Dahlqvist A.,and Stymne S.,1997,Triacylglycerols are synthesised and utilized by transacylation reactions in microsomal preparations of develop ing safflower(Carthamus tinctorius L.)seeds,Planta,203(1):58-66
Sun D.,Zhu J.Q.,Lei F.,Xin Z.,Chow Y.,and Liu J.H.,2013,De novo transcriptome profiling uncovers a drastic downregulation of photosynthesis upon nitrogen deprivation in the nonmodel green alga botryosphaerella sudeticus,BMCGenomics,14(1):715
van Erp H.,Bates P.D.,Burgal J.,Shockey J.,and Browse J.,2011,Castor phospholipid:diacylglycerol acyltransferase facilitates efficient metabolism of hydroxy fatty acids in transgenic Arabidopsis,Plant Physiol.,155(2):683-693
Vitova M.,Bisova K.,Kawano S.,and Zachleder V.,2015,Accumulation of energy reserves in algae:from cell cycles to biotechnological applications,Biotechnol.Adv.,33(6):1204-1218
Wendel A.A.,Lewin T.M.,and Coleman R.A.,2009,Glycerol-3-phosphate acyltransferases:rate limiting enzymes of triacylglycerol biosynthesis,Biochim Biophys Acta.,1791(6):501-506
Xu J.Y.,Zheng Z.F.,Zou J.T.,Peterson R.L.,and Shelp B.J.,2009,A membrane-bound glycerol-3-phosphate acyltransferase from thalassiosira pseudonana regulates acyl composition of glycerolipids,Botany-botanique,87(6):544-551
Yoon K.,Han D.,Li Y.,Sommerfeld M.,and Hu Q.,2012,Phospholipid:diacylglycerol acyltransferase is a multifunctional enzyme involved in membrane lipid turnover and degradation while synthesizing triacylglycerol in the unicellular green microalga Chlamydomonas reinhardtii,Plant Cell,24(9):3708-3724
Zhang H.,Damude H.G.,and Yadav N.S.,2012,Three diacylglycerol acyltransferases contribute to oil biosynthesis and normal growth in Yarrowia lipolytica,Yeast,29(1):25-38
Bagnato C.,Prados M.B.,Franchini G.R.,Scaglia N.,Miranda S.E.,and Beligni M.V.,2017,Analysis of triglyceride synthesis unveils a green algal soluble diacylglycerol acyltransferase and provides clues to potential enzymatic components of the chloroplast pathway,BMC Genomics,18(1):223
Bharathiraja B.,Jayamuthunagai J.,Chakravarthy M.,Kumar R.R.,Yogendran D.,and Praveenkumar R.,2015,Algae:Promising future feedstock for biofuels,In:Singh B.,Bauddh K.,and Bux F.(eds.),Algae and environmental sustainability,Springer-Verlag,Berlin,Germany,pp.1-8
Brembu T.,Mühlroth A.,Alipanah L.,and Bones A.M.,2017,The effects of phosphorus limitation on carbon metabolism in diatoms,Philos.Trans.R.Soc.Lond.B.Biol.Sci.,372(1728):20160406
Chen J.E.,and Smith A.G.,2012,A look at diacylglycerol acyltransferases(DGATs)in algae,J.Biotechnol.,162(1):28-39
Dahlqvist A.,Stahl U.,Lenman M.,Banas A.,Lee M.,Sandager L.,Ronne H.,and Stymne S.,2000,Phospholipid:diacylglycerol acyltransferase:an enzyme that catalyzes the acyl-coa-independent formation of triacylglycerol in yeast and plants,Proc.Natl.Acad.Sci.USA,97(12):6487-6492
Deng X.D.,Gu B.,Li Y.J.,Hu X.W.,Guo J.C.,and Fei X.W.,2012,The roles of acyl-coa:diacylglycerol acyltransferase 2genes in the biosynthesis of triacylglycerols by the green algae Chlamydomonas reinhardtii,Mol.Plant,5(4):945-947
Dunahay T.G.,Jarvis E.E.,Dais S.S.,and Roessler P.G.,1996,Manipulation of microalgal lipid production using genetic engineering,Appl.Biochem.Biotechnol.,57-58(1):223-231
Goodwin S.,Mcpherson J.D.,and Mccombie W.R.,2016,Coming of age:ten years of next-generation sequencing technologies,Nat.Rev.Genet.,17(6):333-351
Gorain P.C.,Bagchi S.K.,and Mallick N.,2013,Effects of calcium,magnesium and sodium chloride in enhancing lipid accumulation in two green microalgae,Environ.Technol.,34(13-16):1887-1894
Hern觔N.A.,Zapata-Morales A.L.,Ochoa Alfaro A.E.,and Soria-Guerra R.E.,2016,Identification of gene transcripts involved in lipid biosynthesis in Chlamydomonas reinhardtii under nitrogen,iron and sulfur deprivation,World J.Microbiol.Biotechnol.,32(4):1-10
Hu Q.,Sommerfeld M.,Jarvis E.,Ghirardi M.,Posewitz M.,Seibert,M.,and Darzins A.,2008,Microalgal triacylglycerols as feedstocks for biofuel production:perspectives and advances,Plant J.,54(4):621-639
Hung C.H.,Kanehara K.,and Nakamura Y.,2016,Isolation and characterization of a mutant defective in triacylglycerol accumulation in nitrogen-starved Chlamydomonas reinhardtii,Biochim.Biophys.Acta,1861(9):1282-1293
Jaeger L.D.,Verbeek R.E.,Draaisma R.B.,Martens D.E.,Springer J.,Eggink G.,and Wijffels R.H.,2014,Superior triacylglycerol(TAG)accumulation in starchless mutants of scenedesmus obliquus:(i)mutant generation and characterization,Biotechnol.Biofuels.,7(1):69
Kan G.F.,Shi C.G.,Wang X.F.,Xie Q.J.,Wang M.,Wang X.L.,and Miao J.L.,2012,Acclimatory responses to high-salt stress in Chlamydomonas(Chlorophyta,Chlorophyceae)from Antarctica,Acta Oceanol.Sin.,31(1):116-124
Kim H.U.,Lee K.R.,Go Y.S.,Jung J.H.,Suh M.C.,and Kim J.B.,2011,Endoplasmic reticulum-located pdat1-2 from castor bean enhances hydroxy fatty acid accumulation in transgenic plants,Plant Cell Physiol.,52(6):983-993
Klaus D.,Ohlrogge J.B.,Neuhaus H.E.,and D觟rmann P.,2004,Increased fatty acid production in potato by engineering of acetyl-coa carboxylase,Planta,219(3):389-396
Klok A.J.,Martens D.E.,Wijffels R.H.,and Lamers P.P.,2013,Simultaneous growth and neutral lipid accumulation in microalgae,Bioresour.Technol.,134:233-243
Lehner R.,and Kuksis A.,1993,Triacylglycerol synthesis by an sn-1,2(2,3)-diacylglycerol transacylase from rat intestinal microsomes,J.Biol.Chem.,268(12):8781-8186
Li Y.,Han D.,Hu G.,Sommerfeld M.,and Hu Q.,2010,Inhibition of starch synthesis results in overproduction of lipids in Chlamydomonas reinhardtii,Biotechnol.Bioeng.,107(2):258-268
Li-Beisson Y.,Beisson F.,and Riekhof W.,2015,Metabolism of acyl-lipids in chlamydomonas reinhardtii,Plant J.,82(3):504
Liu B.,and Benning C.,2013,Lipid metabolism in microalgae distinguishes itself,Curr.Opin.Biotechnol.,24(2):300-309
Lv H.,Qu G.,Qi X.,Lu L.,Tian C.,and Ma Y.,2013,Transcriptome analysis of Chlamydomonas reinhardtii during the process of lipid accumulation,Genomics,101(4):229-237
Maisonneuve S.,Bessoule J.J.,Lessire R.,Delseny M.,and Roscoe T.J.,2010,Expression of rapeseed microsomal lysophosphatidic acid acyltransferase isozymes enhances seed oil content in arabidopsis,Plant Physiol.,152(2):670-684
Michelon W.,Da S.M.,Mezzari M.P.,Pirolli M.,Prandini J.M.,and Soares H.M.,2015,Effects of nitrogen and phosphorus on biochemical composition of microalgae polyculture harvested from phycoremediation of piggery wastewater digestate,Appl.Biochem.Biotechnol.,178(7):1-13
Miller R.,Wu G.X.,Deshpande R.R.,Vieler A.,G覿rtner K.,Li X.B.,Moellering E.R.,Z覿uner S.,Cornish A.J.,Liu B.,Bullard B.,Sears B.B.,Kuo M.H.,Hegg E.L.,Shachar-Hill Y.,Shiu S.H.,and Benning C.,2010,Changes in transcript abundance in Chlamydomonas reinhardtii following nitrogen deprivation predict diversion of metabolism,Plant Physiol.,154(4):1737-1752
Msanne J.,Xu D.,Kond a A.R.,Casasmollano J.A.,Awada T.,Cahoon E.B.,Cerutti H.,2012,Metabolic and gene expression changes triggered by nitrogen deprivation in the pho toautotrophically grown microalgae Chlamydomonas reinhardtii and coccomyxa sp.c-169,Phytochemistry.,75:50-59
Mujtaba G.,Choi W.,Lee C.G.,and Lee K.,2012,Lipid production by chlorella vulgaris after a shift from nutrient-rich to nitrogen starvation conditions,Bioresour.Technol.,123(123):279-283
Ota S.,Oshima K.,Yamazaki T.,Kim S.,Yu Z.,Yoshihara M.,Takeda K.,Takeshita T.,Hirata A.,Bi觢ováK.,Zachleder V.,Hattori M.,and Kawano S.,2016,Highly efficient lipid production in the green alga parachlorella kessleri:draft genome and transcriptome endorsed by whole-cell 3d ultrastructure,Biotechnol.Biofuels.,9(1):13
Pan X.,Siloto R.M.P.,Wickramarathna A.D.,Mietkiewska E.,and Weselake R.J.,2013,Identification of a pair of phospholipid:diacylglycerol acyltransferases from developing flax(Linum usitatissimum L.)seed catalyzing the selective production of trilinolenin,J.Biol.Chem.,288(33):24173-24188
Pandit P.R.,Fulekar M.H.,and Karuna M.S.,2017,Effect of salinity stress on growth,lipid productivity,fatty acid composition,and biodiesel properties in acutodesmus obliquus and chlorella vulgaris,Environ.Sci.Pollut.Res.Int.,24(15):13437-13451
Radakovits R.,Jinkerson R.E.,Darzins A.,and Posewitz M.C.,2010,Genetic engineering of algae for enhanced biofuel production,Eukaryot.Cell.,9(4):486-501
Rani S.H.,Krishna T.H.,Saha S.,Negi A.S.,and Rajasekharan R.,2010,Defective in cuticular ridges(DCR)of Arabidopsis thaliana,a gene associated with surface cutin formation,encodes a soluble diacylglycerol acyltransferase,J.Biol.Chem.,285(49):38337-38347
Riekhof W.R.,Sears B.B.,and Benning C.,2005,Annotation of genes involved in glycerolipid biosynthesis in Chlamydomonas reinhardtii:discovery of the betaine lipid synthase bta1cr,Eukaryot.Cell,4(2):242-252
Roopnarain A.,Gray V.M.,and Sym S.D.,2014,Phosphorus limitation and starvation effects on cell growth and lipid accumulation in isochrysis galbana u4 for biodiesel production,Bioresour.Technol.,156(2):408-411
Saha S.,Enugutti B.,Rajakumari S.,and Rajasekharan R.,2006,Cytosolic triacylglycerol biosynthetic pathway in oilseeds molecular cloning and expression of peanut cytosolic diacylglycerol acyltransferase,Plant Physiol.,141(4):1533-1543
Sasaki Y.,and Nagano Y.,2004,Plant acetyl-coa carboxylase:structure,biosynthesis,regulation,and gene manipulation for plant breeding,Biosci.Biotechnol.Biochem.,68(6):1175
Sato A.,Matsumura R.,Hoshino N.,Tsuzuki M.,and Sato N.,2014,Responsibility of regulatory gene expression and repressed protein synthesis for triacylglycerol accumulation on sulfur-starvation in Chlamydomonas reinhardtii,Front.Plant Sci.,5:444
Schuler L.M.,Schulze P.S.C.,Pereira H.,Barreira L.,Leon R.,and Varela J.,2017,Trends and strategies to enhance triacylglycerols and high-value compounds in microalgae,Algal Res.,25:263-273
Sharma K.K.,Schuhmann H.,and Schenk P.M.,2012,High lipid induction in microalgae for biodiesel production,Energies.,5(5):1532-1553
Stahl U.,Carlsson A.S.,Lenman M.,Dahlqvist A.,Huang B.,Banas W.,Banas A.,and Stymne S.,2004,Cloning and functional characterization of a phospholipid:diacylglycerol acyltransferase from Arabidopsis,Plant Physiol.,135(3):1324-1335
Stobart K.,Mancha M.,Lenman M.,Dahlqvist A.,and Stymne S.,1997,Triacylglycerols are synthesised and utilized by transacylation reactions in microsomal preparations of develop ing safflower(Carthamus tinctorius L.)seeds,Planta,203(1):58-66
Sun D.,Zhu J.Q.,Lei F.,Xin Z.,Chow Y.,and Liu J.H.,2013,De novo transcriptome profiling uncovers a drastic downregulation of photosynthesis upon nitrogen deprivation in the nonmodel green alga botryosphaerella sudeticus,BMCGenomics,14(1):715
van Erp H.,Bates P.D.,Burgal J.,Shockey J.,and Browse J.,2011,Castor phospholipid:diacylglycerol acyltransferase facilitates efficient metabolism of hydroxy fatty acids in transgenic Arabidopsis,Plant Physiol.,155(2):683-693
Vitova M.,Bisova K.,Kawano S.,and Zachleder V.,2015,Accumulation of energy reserves in algae:from cell cycles to biotechnological applications,Biotechnol.Adv.,33(6):1204-1218
Wendel A.A.,Lewin T.M.,and Coleman R.A.,2009,Glycerol-3-phosphate acyltransferases:rate limiting enzymes of triacylglycerol biosynthesis,Biochim Biophys Acta.,1791(6):501-506
Xu J.Y.,Zheng Z.F.,Zou J.T.,Peterson R.L.,and Shelp B.J.,2009,A membrane-bound glycerol-3-phosphate acyltransferase from thalassiosira pseudonana regulates acyl composition of glycerolipids,Botany-botanique,87(6):544-551
Yoon K.,Han D.,Li Y.,Sommerfeld M.,and Hu Q.,2012,Phospholipid:diacylglycerol acyltransferase is a multifunctional enzyme involved in membrane lipid turnover and degradation while synthesizing triacylglycerol in the unicellular green microalga Chlamydomonas reinhardtii,Plant Cell,24(9):3708-3724
Zhang H.,Damude H.G.,and Yadav N.S.,2012,Three diacylglycerol acyltransferases contribute to oil biosynthesis and normal growth in Yarrowia lipolytica,Yeast,29(1):25-38