基因工程对林木生长表型与细胞壁组分及构造的影响研究
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摘要
通过基因工程技术培养出木质素含量低、纤维素含量高和糖转化效率高以及优质的木材,对于将其定向应用于制浆造纸、生物炼制、木质建筑及装饰材料方面具有重要的研究意义。文章详细阐明了木质素和纤维素基因调控技术对转基因林木生长表型、细胞壁化学组分含量及其微区分布、组织细胞形态及细胞壁超微构造影响的研究进展,并对转基因林木今后的重点发展方向进行了展望,以期为我国定向培育优质速生人工林提供理论依据。
Transgenic trees with low lignin content, high cellulose content, high conversion efficiency of sugar,and excellent wood properties were achieved by genetic engineering, which have important research value for the directional application in pulping and paper making, biomass energy refining and wood building and decorative materials. In this paper, the effects of lignin and cellulose gene regulation on growth phenotype, cell wall chemical component and distribution, tissue cells morphology and ultrastructure of cell wall in transgenic trees were reviewed in detail. And the prospects for future research and development direction of transgenic forest are put forward. The purpose of this article is to provide theoretical guidance for the directional cultivation of high quality and fast growing plantations in China.
Transgenic trees with low lignin content, high cellulose content, high conversion efficiency of sugar,and excellent wood properties were achieved by genetic engineering, which have important research value for the directional application in pulping and paper making, biomass energy refining and wood building and decorative materials. In this paper, the effects of lignin and cellulose gene regulation on growth phenotype, cell wall chemical component and distribution, tissue cells morphology and ultrastructure of cell wall in transgenic trees were reviewed in detail. And the prospects for future research and development direction of transgenic forest are put forward. The purpose of this article is to provide theoretical guidance for the directional cultivation of high quality and fast growing plantations in China.
引文
[1]刘一星,赵广杰.木质资源材料学[M].北京:中国林业出版社,2004.
[2]Sticklen M.Plant genetic engineering to improve biomass characteristics for biofuels[J].Current Opinion in Biotechnology,2006,17(3):315-319.
[3]Rubin E M.Genomics of cellulosic biofuels[J].Nature,2008,454(7206):841-845.
[4]Chen F,Dixon R A.Lignin modification improves fermentable sugar yields for biofuel production[J].Nature Biotechnology,2007,25(7):759-761.
[5]周贤武,高玉磊,苏明垒,等.基因工程改良木材性质研究进展[J].林业科学,2018,54(3):152-160.
[6]田晓明,谭晓风.转基因杨树的研究进展及展望[J].湖南林业科技,2009,36(2):71-73.
[7]孔华,郭安平,郭运玲,等.木质素生物合成及转基因调控研究进展[J].热带农业工程,2009,33(5):47-53.
[8]高艳,陈光辉,陈秀娟,等.植物细胞壁纤维素生物合成的调控[J].生物技术通报,2014(1):1-7.
[9]Verma S R,Dwivedi U N.Lignin genetic engineering for improvement of wood quality:Applications in paper and textile industries,fodder and bioenergy production[J].South African Journal of Botany,2014,91(3):107-125.
[10]赵华燕,魏建华,宋艳茹.木质素生物合成及其基因工程研究进展[J].植物生理与分子生物学学报,2004,30(4):361-370.
[11]Baucher M,Halpin C,Petit-Conil M,et al.Lignin:genetic engineering and impact on pulping[J].Critical Reviews in Biochemistry and Molecular Biology,2003,38(4):305-350.
[12]刘苍伟,苏明垒,王玉荣,等.木质素基因调控对木材细胞壁化学组分与微观结构影响的研究进展[J].西北农林科技大学学报(自然科学版),2018,46(2):39-46.
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[14]Somerville C.Cellulose synthesis in higher plants[J].Annual review of cell and developmental biology,2006,22(1):53-78.
[15]Chen S,Ehrhardt D W,Somerville C R.Mutations of cellulose synthase(CESA1)phosphorylation sites modulate anisotropic cell expansion and bidirectional mobility of cellulose synthase[J].Proceedings of the National Academy of Sciences,2010,107(40):17188-17193.
[16]Carroll A,Mansoori N,Li S,et al.Complexes with mixed primary and secondary cellulose synthases are functional in arabidopsis plants[J].Plant Physiology,2012,160(2):726-737.
[17]Arioli T,Peng L,Betzner A S,et al.Molecular analysis of cellulose biosynthesis in Arabidopsis[J].Science,1998,279(5351):717-720.
[18]Wang L,Guo K,Li Y,et al.Expression profiling and integrative analysis of the CESA/CSL superfamily in rice[J].BMC Plant Biology,2010,10(1):282-297.
[19]Djerbi S,Lindskog M,Arvestad L,et al.The genome sequence of black cottonwood(Populus trichocarpa)reveals 18 conserved cellulose synthase(CesA)genes[J].Planta,2005,221(5):739-746.
[20]Kim W C,Ko J H,Kim J Y,et al.MYB46 directly regulates the gene expression of secondary wall associated cellulose synthases in Arabidopsis[J].Plant Journal,2013,73(1):26-36.
[21]Huang D,Wang S,Zhang B,et al.A gibberellin-mediated DELLA-NAC signaling cascade regulates cellulose synthesis in rice[J].Plant Cell,2015,27(6):1681-1696.
[22]Singh B,Cheek H D,Haigler C H.A synthetic auxin(NAA)suppresses secondary wall cellulose synthesis and enhances elongation in cultured cotton fiber[J].Plant Cell,2009,28(7):1023-1032.
[23]邰付菊,李学宝.植物纤维素生物合成及其相关酶类[J].细胞生物学杂志,2004,26(5):490-494.
[24]Xie L,Yang C,Wang X.Brassinosteroids can regulate cellulose biosynthesis by controlling the expression of CESA genes in Arabidopsis[J].Journal of Experimental Botany,2011,62(13):4495-4506.
[25]Zhou X,Ren S,Lu M,Zhao S,et al.Preliminary study of cell wall structure and its mechanical properties of C3H and HCT RNAi transgenic poplar sapling[J].scientific reports,2018(8):105-108.
[26]彭霄鹏.杨树木质素合成基因C3H与HCT表达下调对细胞壁组成与结构的影响[D].北京:北京林业大学,2015.
[27]Hu W J,Harding S A,Jrhau L,et al.Repression of lignin biosynthesis promotes cellulose accumulation and growth in transgenic trees.[J].Nature Biotechnology,1999,17(8):808-812.
[28]Tian X M,Xie J,Zhao Y L,et al.Sense-,antisense-and RNAi-4CL1 regulate soluble phenolic acids,cell wall components and growth in transgenic Populus tomentosa Carr.[J].Plant Physiology&Biochemistry Ppb,2013,65(6):111-119.
[29]Zhang G,Zhang Y,Xu J,et al.The CCoAOMT1 gene from jute(Corchorus capsularis L.)is involved in lignin biosynthesis in Arabidopsis thaliana[J].Gene,2014,546(2):398-402.
[30]Liu Y,Xu F,Gou J,et al.Importance of two consecutive methionines at the N-terminus of a cellulose synthase(PtdCesA8A)for normal wood cellulose synthesis in aspen[J].Tree Physiology,2012,32(11):1403-1412.
[31]魏凯莉.MYB216调控杨树纤维素合成的机制研究[D].北京:中国林业科学研究院,2016.
[32]Jung Y H,Cho H J,Lee J S,et al.Evaluation of a transgenic poplar as a potential biomass crop for biofuel production[J].Bioresource Technology,2013,129(2):639-641.
[33]Payyavula R S,Tschaplinski T J,Jawdy S S,et al.Metabolic p?rofiling reveals altered sugar and secondary metabolism in response to UGPase overexpression in Populus[J].BMC Plant Biology,2014,14(1):1-14.
[34]Wagner A,Ralph J,Akiyama T,et al.Exploring lignification in conifers by silencing hydroxycinnamoyl-CoA:shikimate hydroxycinnamoyl transferase in Pinus radiata.[J].Proceedings of the National Academy of Sciences of the United States of America,2007,104(28):11856-11861.
[35]?zparpucu M,Gierlinger N,Burgert I,et al.The effect of altered lignin composition on mechanical properties of Cinnamyl Alcohol Dehydrogenase(CAD)deficient poplars[J].Planta,2018,247(4):887-897.
[36]苏明垒.HCT和MYB216转基因银腺杨木材化学组分与微观构造研究[D].北京:中国林业科学研究院,2018.
[37]Tang X,Zhuang Y,Qi G,et al.Poplar PdMYB221 is involved in the direct and indirect regulation of secondary wall biosynthesis during wood formation[J].Scientific Reports,2015,5:12240.
[38]Schrick K,Fujioka S,Takatsuto S,et al.A link between sterol biosynthesis,the cell wall,and cellulose in Arabidopsis[J].Plant Journal,2004,38(2):227-243.
[39]叶胜龙.毛白杨MYB055转录因子在次生壁合成中的调控机制研究[D].重庆:西南大学,2015.
[40]Silvia F,Montserrat C,Antonio E,et al.Altered lignin biosynthesis improves cellulosic bioethanol production in transgenic maize plants down-regulated for Cinnamyl Alcohol Dehydrogenase[J].Molecular Plant,2012,5(4):817-830.
[41]刘苍伟,苏明垒,周贤武,等.FTIR及CLSM对转基因杨木细胞壁木质素含量及微区分布研究[J].光谱学与光谱分析,2017,37(11):3404-3408.
[42]LepléJ C,Dauwe R,Morreel K,et al.Downregulation of cinnamoylcoenzyme A reductase in poplar:multiple-level phenotyping reveals effects on cell wall polymer metabolism and structure[J].Plant Cell,2007,19(11):3669-3691.
[43]Roudier F,Fernandez A G,Fujita M,et al.COBRA,an Arabidopsis extracellular glycosyl-phosphatidyl inositol-anchored protein,specifically controls highly anisotropic expansion through its involvement in cellulose microfibril orientation[J].Plant Cell,2005,17(6):1749-1763.
[44]Pattathil S,Saffold T,Gallegogiraldo L,et al.Changes in cell wall carbohydrate extractability are correlated with reduced recalcitrance of HCT downregulated alfalfa biomass[J].Industrial Biotechnology,2012,8(4):217-221.
[45]Ruel K,Berriosierra J,Derikvand M M,et al.Impact of CCR1silencing on the assembly of lignified secondary walls in Arabidopsis thaliana[J].New Phytologist,2009,184(1):99-113.
[46]Nolin F.Caffeoyl coenzyme A O-methyltransferase down-regulation is associated with modifications in lignin and cell-wall architecture in flax secondary xylem[J].Plant Physiology&Biochemistry,2009,47(1):9-19.
[47]苏明垒,刘苍伟,王玉荣,等.C3H-RNAi转基因杨木木质部纤维形态特征研究[J].中国造纸,2017,36(12):21-26.
[48]刘苍伟.C3H-RNAi转基因杨木组织细胞形态特征及细胞壁木质素变化规律研究[D].北京:中国林业科学研究院,2017.
[49]Tan H T,Shirley N J,Singh R R,et al.Powerful regulatory systems and post-transcriptional gene silencing resist increases in cellulose content in cell walls of barley[J].BMC Plant Biology,2015,15(1):1-16.
[2]Sticklen M.Plant genetic engineering to improve biomass characteristics for biofuels[J].Current Opinion in Biotechnology,2006,17(3):315-319.
[3]Rubin E M.Genomics of cellulosic biofuels[J].Nature,2008,454(7206):841-845.
[4]Chen F,Dixon R A.Lignin modification improves fermentable sugar yields for biofuel production[J].Nature Biotechnology,2007,25(7):759-761.
[5]周贤武,高玉磊,苏明垒,等.基因工程改良木材性质研究进展[J].林业科学,2018,54(3):152-160.
[6]田晓明,谭晓风.转基因杨树的研究进展及展望[J].湖南林业科技,2009,36(2):71-73.
[7]孔华,郭安平,郭运玲,等.木质素生物合成及转基因调控研究进展[J].热带农业工程,2009,33(5):47-53.
[8]高艳,陈光辉,陈秀娟,等.植物细胞壁纤维素生物合成的调控[J].生物技术通报,2014(1):1-7.
[9]Verma S R,Dwivedi U N.Lignin genetic engineering for improvement of wood quality:Applications in paper and textile industries,fodder and bioenergy production[J].South African Journal of Botany,2014,91(3):107-125.
[10]赵华燕,魏建华,宋艳茹.木质素生物合成及其基因工程研究进展[J].植物生理与分子生物学学报,2004,30(4):361-370.
[11]Baucher M,Halpin C,Petit-Conil M,et al.Lignin:genetic engineering and impact on pulping[J].Critical Reviews in Biochemistry and Molecular Biology,2003,38(4):305-350.
[12]刘苍伟,苏明垒,王玉荣,等.木质素基因调控对木材细胞壁化学组分与微观结构影响的研究进展[J].西北农林科技大学学报(自然科学版),2018,46(2):39-46.
[13]高原,陈信波,张志扬.木质素生物合成途径及其基因调控的研究进展[J].生物技术通报,2007(2):47-51.
[14]Somerville C.Cellulose synthesis in higher plants[J].Annual review of cell and developmental biology,2006,22(1):53-78.
[15]Chen S,Ehrhardt D W,Somerville C R.Mutations of cellulose synthase(CESA1)phosphorylation sites modulate anisotropic cell expansion and bidirectional mobility of cellulose synthase[J].Proceedings of the National Academy of Sciences,2010,107(40):17188-17193.
[16]Carroll A,Mansoori N,Li S,et al.Complexes with mixed primary and secondary cellulose synthases are functional in arabidopsis plants[J].Plant Physiology,2012,160(2):726-737.
[17]Arioli T,Peng L,Betzner A S,et al.Molecular analysis of cellulose biosynthesis in Arabidopsis[J].Science,1998,279(5351):717-720.
[18]Wang L,Guo K,Li Y,et al.Expression profiling and integrative analysis of the CESA/CSL superfamily in rice[J].BMC Plant Biology,2010,10(1):282-297.
[19]Djerbi S,Lindskog M,Arvestad L,et al.The genome sequence of black cottonwood(Populus trichocarpa)reveals 18 conserved cellulose synthase(CesA)genes[J].Planta,2005,221(5):739-746.
[20]Kim W C,Ko J H,Kim J Y,et al.MYB46 directly regulates the gene expression of secondary wall associated cellulose synthases in Arabidopsis[J].Plant Journal,2013,73(1):26-36.
[21]Huang D,Wang S,Zhang B,et al.A gibberellin-mediated DELLA-NAC signaling cascade regulates cellulose synthesis in rice[J].Plant Cell,2015,27(6):1681-1696.
[22]Singh B,Cheek H D,Haigler C H.A synthetic auxin(NAA)suppresses secondary wall cellulose synthesis and enhances elongation in cultured cotton fiber[J].Plant Cell,2009,28(7):1023-1032.
[23]邰付菊,李学宝.植物纤维素生物合成及其相关酶类[J].细胞生物学杂志,2004,26(5):490-494.
[24]Xie L,Yang C,Wang X.Brassinosteroids can regulate cellulose biosynthesis by controlling the expression of CESA genes in Arabidopsis[J].Journal of Experimental Botany,2011,62(13):4495-4506.
[25]Zhou X,Ren S,Lu M,Zhao S,et al.Preliminary study of cell wall structure and its mechanical properties of C3H and HCT RNAi transgenic poplar sapling[J].scientific reports,2018(8):105-108.
[26]彭霄鹏.杨树木质素合成基因C3H与HCT表达下调对细胞壁组成与结构的影响[D].北京:北京林业大学,2015.
[27]Hu W J,Harding S A,Jrhau L,et al.Repression of lignin biosynthesis promotes cellulose accumulation and growth in transgenic trees.[J].Nature Biotechnology,1999,17(8):808-812.
[28]Tian X M,Xie J,Zhao Y L,et al.Sense-,antisense-and RNAi-4CL1 regulate soluble phenolic acids,cell wall components and growth in transgenic Populus tomentosa Carr.[J].Plant Physiology&Biochemistry Ppb,2013,65(6):111-119.
[29]Zhang G,Zhang Y,Xu J,et al.The CCoAOMT1 gene from jute(Corchorus capsularis L.)is involved in lignin biosynthesis in Arabidopsis thaliana[J].Gene,2014,546(2):398-402.
[30]Liu Y,Xu F,Gou J,et al.Importance of two consecutive methionines at the N-terminus of a cellulose synthase(PtdCesA8A)for normal wood cellulose synthesis in aspen[J].Tree Physiology,2012,32(11):1403-1412.
[31]魏凯莉.MYB216调控杨树纤维素合成的机制研究[D].北京:中国林业科学研究院,2016.
[32]Jung Y H,Cho H J,Lee J S,et al.Evaluation of a transgenic poplar as a potential biomass crop for biofuel production[J].Bioresource Technology,2013,129(2):639-641.
[33]Payyavula R S,Tschaplinski T J,Jawdy S S,et al.Metabolic p?rofiling reveals altered sugar and secondary metabolism in response to UGPase overexpression in Populus[J].BMC Plant Biology,2014,14(1):1-14.
[34]Wagner A,Ralph J,Akiyama T,et al.Exploring lignification in conifers by silencing hydroxycinnamoyl-CoA:shikimate hydroxycinnamoyl transferase in Pinus radiata.[J].Proceedings of the National Academy of Sciences of the United States of America,2007,104(28):11856-11861.
[35]?zparpucu M,Gierlinger N,Burgert I,et al.The effect of altered lignin composition on mechanical properties of Cinnamyl Alcohol Dehydrogenase(CAD)deficient poplars[J].Planta,2018,247(4):887-897.
[36]苏明垒.HCT和MYB216转基因银腺杨木材化学组分与微观构造研究[D].北京:中国林业科学研究院,2018.
[37]Tang X,Zhuang Y,Qi G,et al.Poplar PdMYB221 is involved in the direct and indirect regulation of secondary wall biosynthesis during wood formation[J].Scientific Reports,2015,5:12240.
[38]Schrick K,Fujioka S,Takatsuto S,et al.A link between sterol biosynthesis,the cell wall,and cellulose in Arabidopsis[J].Plant Journal,2004,38(2):227-243.
[39]叶胜龙.毛白杨MYB055转录因子在次生壁合成中的调控机制研究[D].重庆:西南大学,2015.
[40]Silvia F,Montserrat C,Antonio E,et al.Altered lignin biosynthesis improves cellulosic bioethanol production in transgenic maize plants down-regulated for Cinnamyl Alcohol Dehydrogenase[J].Molecular Plant,2012,5(4):817-830.
[41]刘苍伟,苏明垒,周贤武,等.FTIR及CLSM对转基因杨木细胞壁木质素含量及微区分布研究[J].光谱学与光谱分析,2017,37(11):3404-3408.
[42]LepléJ C,Dauwe R,Morreel K,et al.Downregulation of cinnamoylcoenzyme A reductase in poplar:multiple-level phenotyping reveals effects on cell wall polymer metabolism and structure[J].Plant Cell,2007,19(11):3669-3691.
[43]Roudier F,Fernandez A G,Fujita M,et al.COBRA,an Arabidopsis extracellular glycosyl-phosphatidyl inositol-anchored protein,specifically controls highly anisotropic expansion through its involvement in cellulose microfibril orientation[J].Plant Cell,2005,17(6):1749-1763.
[44]Pattathil S,Saffold T,Gallegogiraldo L,et al.Changes in cell wall carbohydrate extractability are correlated with reduced recalcitrance of HCT downregulated alfalfa biomass[J].Industrial Biotechnology,2012,8(4):217-221.
[45]Ruel K,Berriosierra J,Derikvand M M,et al.Impact of CCR1silencing on the assembly of lignified secondary walls in Arabidopsis thaliana[J].New Phytologist,2009,184(1):99-113.
[46]Nolin F.Caffeoyl coenzyme A O-methyltransferase down-regulation is associated with modifications in lignin and cell-wall architecture in flax secondary xylem[J].Plant Physiology&Biochemistry,2009,47(1):9-19.
[47]苏明垒,刘苍伟,王玉荣,等.C3H-RNAi转基因杨木木质部纤维形态特征研究[J].中国造纸,2017,36(12):21-26.
[48]刘苍伟.C3H-RNAi转基因杨木组织细胞形态特征及细胞壁木质素变化规律研究[D].北京:中国林业科学研究院,2017.
[49]Tan H T,Shirley N J,Singh R R,et al.Powerful regulatory systems and post-transcriptional gene silencing resist increases in cellulose content in cell walls of barley[J].BMC Plant Biology,2015,15(1):1-16.