松科植物木质素合成相关基因研究进展
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摘要
植物木质素是影响纸浆材品质的关键因素,而松科植物作为重要的纸浆材,对其木质素合成相关调控基因进行系统了解,可为完善木质素生物合成模型提供一定的理论依据。笔者通过对国内外已有研究进行分析,总结植物木质素结构、木质素合成途径、木质素合成相关酶基因及松科植物木质素合成基因的研究进展。归纳出松科植物火炬松(Pinus taeda)、马尾松(P.massoniana)、辐射松(P.radiata)、挪威云杉(Picea abie)等木质素合成酶基因研究主要集中在苯丙氨酸解氨酶(phenylalanineammonialyase,PAL)、4-香豆酸辅酶A连接酶(4-coumarate Co A ligase,4CL)、肉桂酸4-羟化酶(cinnamic 4-hydroxygenase,C4H)、香豆酸3-羟化酶(Coumarate 3-hydroxylase,C3H)、咖啡酰辅酶A-O-甲基转移酶(caffeoyl coenzyme A-O-methyltransferase,CCo AOMT)、肉桂酰基辅酶A还原酶(Cinnamoyl Co A reductase,CCR)和肉桂醇脱氢酶(Cinnamyl alcohol dehydrogenase,CAD)等一些常见基因上,其他莽草酸羟基肉桂酰转移酶(shikimate hydroxycinnamoyl transferase,HCT)、阿魏酸5-羟化酶(ferulo 5-hydroxylase,F5H)、O-甲基转移酶(O-methyltransferase,OMT)及咖啡酸-O-甲基转移酶(caffeic acid O-methyltransferase,COMT)等基因研究较欠缺,此外,还有部分基因未涉及。木质素合成是研究松科植物纸浆材的重要切入点,对现有松科植物木质素合成过程中存在问题进行精确的估计,对进一步深入研究松科植物木质素合成机制和优质纸浆材选育具有重要意义。
Plant lignin is an essential factor of affecting pulp quality. Pinaceae plants is an important pulp material which means the overall understanding of patterns and controlling factors of lignin synthesis related genes is key for improving the modeling of lignin synthesis. According the current research results,the lignin structure,the lignin synthesis pathway and the related enzymes gene of lignin synthesis in Pinaceae are summarized. Some related genes of lignin synthesis in Pinus taeda,P. massoniana,P. radiata,Picea abie and so on have been researched. The lignin syntheses genes of Pinaceae researches were mainly concentrated in common genes,such as phenylalanine ammonolyase( PAL),4-coumarate:coenzyme A ligase( 4 CL),cinnamic 4-hydroxygenase( C4 H),coumarate 3-hydroxylase( C3 H),caffeoyl coenzyme AO-methyltransferase( CCo AOMT),cinnamoyl Co A reductase( CCR) and cinnamyl alcohol dehydrogenase( CAD),however,some genes that lack current research,including shikimate hydroxycinnamoyl transferase( HCT),ferulate 5-hydroxylase( F5 H),O-methyltransferase( OMT),and caffeic acid O-methyltransferase( COMT). Moreover,some genes were not involved in the study. Lignin synthesis is the essential process of exploring pulp quality of Pinaceae. A better understanding of existing problems of the research on lignin synthesis could contribute to further research on the mechanism of lignin synthesis and help to select high quality pulp genotype in Pinaceae plants.
Plant lignin is an essential factor of affecting pulp quality. Pinaceae plants is an important pulp material which means the overall understanding of patterns and controlling factors of lignin synthesis related genes is key for improving the modeling of lignin synthesis. According the current research results,the lignin structure,the lignin synthesis pathway and the related enzymes gene of lignin synthesis in Pinaceae are summarized. Some related genes of lignin synthesis in Pinus taeda,P. massoniana,P. radiata,Picea abie and so on have been researched. The lignin syntheses genes of Pinaceae researches were mainly concentrated in common genes,such as phenylalanine ammonolyase( PAL),4-coumarate:coenzyme A ligase( 4 CL),cinnamic 4-hydroxygenase( C4 H),coumarate 3-hydroxylase( C3 H),caffeoyl coenzyme AO-methyltransferase( CCo AOMT),cinnamoyl Co A reductase( CCR) and cinnamyl alcohol dehydrogenase( CAD),however,some genes that lack current research,including shikimate hydroxycinnamoyl transferase( HCT),ferulate 5-hydroxylase( F5 H),O-methyltransferase( OMT),and caffeic acid O-methyltransferase( COMT). Moreover,some genes were not involved in the study. Lignin synthesis is the essential process of exploring pulp quality of Pinaceae. A better understanding of existing problems of the research on lignin synthesis could contribute to further research on the mechanism of lignin synthesis and help to select high quality pulp genotype in Pinaceae plants.
引文
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[2]DAVIN L B,LEWIS N G.Phenylpropanoid metabolism:biosynthesis of monolignols,lignans,neolignans,lignins and suberins[J].Rec Adv Phytochem,1992,26:325-375.DOI:10.1007/978-1-4615-3430-3_11.
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[5]BAURHOO B,RUIZ-FERIA C A,ZHAO X.Purified lignin:nutritional and health impacts on farm animals:a review[J].Animal Feed Science and Technology,2008,144(3-4):175-184.DOI:10.1016/j.anifeedsci.2007.10.016.
[6]BHUIYAN N H,SELVARAJ G,WEI Y,et al.Role of lignification in plant defense[J].Plant Signaling&Behavior,2009,4:158-159.DOI:10.4161/psb.4.2.7688.
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[8]魏建华,宋艳茹.木质素生物合成途径及调控的研究进展[J].植物学报,2001,43(8):771-779.WEI J H,SUN Y R.Recent advances in study of lignin biosynthesis and manipulation[J].Journal of Integrative Plant Biology,2001,43(8)771-779.
[9]ONYSKO K A.Biological bleaching of chemical pulps:a review[J].Biotechnology Advances,1993,11(2):179-198.DOI:10.1016/0734-9750(93)90040-T.
[10]BOERJAN W,RALPH J,BAUCHER M.Lignin biosynthesis[J].Annual Review of Plant Biology,2003,54:519-546.DOI:10.1146/annurev.arplant.54.031902.134938.
[11]BOUDET A M,KAJITA S,GRIMA-PETTENATI J,et al.Lignins and lignocellulosics:a better control of synthesis for new and improved uses[J].Trends in Plant Science,2003,8(12):576-581.DOI:10.1016/j.tplants.2003.10.001.
[12]WENG J K,CHAPPLE C T.The origin and evolution of lignin biosynthesis[J].New Phytologist,2010,187(2):273-285.DOI:10.1111/j.1469-8137.2010.03327.x.
[13]PLOMION C,LEPROVOST G,STOKES A.Wood formation in trees[J].Plant Physiology,2001,127:1513-1523.DOI:10.1104/pp.010816.
[14]HATAKEYAMA H,MATSUMURA H,HATAKEYAMA T.Glass transition and thermal degradation of rigid polyurethane foams derived from castor oil-molasses polyols[J].Journal of Thermal A-nalysis and Calorimetry,2013,111(2):1545-1552.DOI:10.1007/s10973-012-2501-5.
[15]HOLLADAY J E,BOZELL J J,WHITE J F,et al.Top value added candidates from biomass,volume II:results of screening for potential candidates from biorefinery lignin[J].Biomass Fuels,2007,(2):263-275.DOI:10.2172/921839.
[16]CHEN Y R,SARKANEN S.X-Ray powder diffraction analyses of kraft lignin-based thermoplastic polymer blends[M].Oxford,UK:Blackwell Publishing Ltd,2009:301-315.
[17]SCHORR D,DIOUF PN,STEVANOVIC T.Evaluation of industrial lignins for biocomposites production[J].Industrial Crops and Products,2014,52(1):65-73.DOI:10.1016/j.indcrop.2013.10.014.
[18]LAPIERRE C,POLLET B,PETIT-CORIL M,et al.Structural alteration of lignin in transgenic poplars with depressed cinnamoyl alcohol dehydrogenase or caffeic acid O-methyltransferase activity have an oppsite impact on the efficiency of industrial kraft pulping[J].Plant Physiology,1999,119:153-164.DOI:10.1104/pp.119.1.153.
[19]BUGOS R C,CHIANG V L C,CAMPBELL W H.c DNA clonging,sequence analysis and seasonal expression of lignin-bispecific caffeic acid/5-hydroxyferulic acid O-methyltransferase of aspen[J].Plant Molecular Biology,1991,17(6):1203-1215.DOI:10.1007/BF00028736.
[20]DOORSSELAERE J V,BAUCHER M,CHOGNOT E,et al.Anovel lignin in poplar trees with a reduced caffeic acid/5-hydroxyferulicacid O-methyltransferase activity[J].Plant Journal,1995,8(6):855-864.DOI:10.1046/j.1365-313X.1995.8060855.x.
[21]HUMPHREYS J M,CHAPPLE C.Rewriting the lignin roadmap[J].Current Opinion in Plant Biology,2002,5(3):224-229.DOI:10.1016/S1369-5266(02)00257-1.
[22]RASTOGI S,DWIVEDI U N.Manipulation of lignin in plants with special reference to O-methyltransferase[J].Plant Science.2008,174(3):264-277.DOI:10.1016/j.plantsci.2007.11.014.
[23]GRIMA-PETTENATI J,GOFFNER D.Lignin genetic engineering revisited[J].Plant Science,1999,145:51-65.DOI:10.1016/S0168-9452(99)00051-5.
[24]RALPH J,LUNDQUIST K,BRUNOW G,et al.Lignins:natural polymers from oxidative coupling of 4-hydroxyphenyl-propanoids[J].Phytochemistry Reviews,2004,3(1):29-60.DOI:10.1023/B:PHYT.0000047809.65444.a4.
[25]VANHOLME R,RALPH J,AKIYAMA T,et al.Engineering traditional monolignols out of lignin by concomitant up-regulation of F5H1 and down-regulation of COMT in Arabidopsis[J].Plant Journal for Cell&Molecular Biology,2010,64:885-897.DOI:10.1111/j.1365-313X.2010.04353.x.
[26]VANHOLME R,CESARINO I,RATAJ K,et al.Caffeoyl shikimate esterase(CSE)is an enzyme in the lignin biosynthetic pathway in Arabidopsis[J].Science,2013,341(6150):1103-1106.DOI:10.1126/science.1241602.
[27]VARGAS L,CESARINO I,VANHOLME R,et al.Improving total saccharification yield of Arabidopsis plants by vesselspecific complementation of caffeoyl shikimate esterase(cse)mutants[J].Biotechnology for Biofuels,2016,9:139-155.DOI:10.1186/s13068-016-0551-9.
[28]DIXON R A,PAIVA N L.Stress-induced phenylpropanoid metabolism[J].The Plant Cell,1995,7:1085-1097.DOI:10.1105/tpc.7.7.1085.
[29]RALPH J,MACKAY J J,HATFIELD R D,et al.Abnormal lignin in a loblolly pine mutant[J].Science,1997,227(5325):235-239.DOI:10.1126/science.277.5323.235.
[30]MACKAY J J,OMALLEY D M,PRESNELL T,et al.Inheritance,gene expression and lignin characterization in a mutant pine deficient in cinnamoyl alcohol dehydrogenase[J].Proc Natl Acad Sci USA,1997,94(15):8255-8260.
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