烤烟成熟期烟叶木质素合成关键酶基因的转录分析
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摘要
木质素对烟叶抗性形成具有重要意义,但含量过高会降低卷烟产品品质。为系统探究木质素在烟叶成熟期合成积累的关键点及调控基因的变化规律,本研究以K326为试验材料,测定烤烟中部叶成熟期叶片硬度、组织结构、木质素含量及相关合成酶基因的表达模式。结果表明,成熟期是烤烟木质素合成积累的重要时期,在整个成熟期后烟叶木质素含量和硬度呈现先增加后降低的变化规律,在打顶后30天,木质素含量和烟叶硬度达到最大值;烟叶成熟期木质素合成酶相关基因的q RT-PCR表达分析表明:PAL、C4H、4CL1、CCR1、CCo AOMT5、COMT1、CAD2、PAO1和F5H1基因表达规律与木质素变化规律较为一致,呈现先增加后降低的变化趋势,且在打顶后30天,表达量达到峰值。其中PAL、C4H、CCR1、CAD2和PAO1基因对烟叶木质素合成积累的贡献较大(p<0.01,r=0.93, 0.90, 0.91, 0.90, 0.89;P=0.24, 0.19, 0.16, 0.17, 0.19),是烟叶木质素合成积累过程中的关键调控基因。
Lignin plays an important role in the formation of tobacco leaf resistance, but excessive content of lignin affect quality of cigarette products. In order to systematically explore the key points for lignin synthesis and accumulation in tobacco leaves at maturity stage as well as the change of regulatory genes, this study analyzes leaf hardness, tissue structure, lignin content and expression patterns of lignin synthase-related genes in middle leaves of flue-cured tobacco at maturity stage using K326 as experimental material. Results showed that maturity stage is an important period for the accumulation of lignin in flue-cured tobacco; the lignin content and hardness of tobacco leaves increased first and then decreased after entering maturity period and then reached the maximum 30 days after topping. Results of quantitative RT-PCR analysis of lignin synthase-related genes in mature tobacco leaves indicated that the expression patterns of PAL, C4 H,4CL1, CCR1, CCoAOMT5, COMT1, CAD, PAO1 and F5H1 were consistent with that of lignin changes, i.e. showing a trend of increasing first and then decreasing, and reaching its peak 30 days after topping. Among them, PAL, C4H, CCR1, CAD2 and PAO1 genes contributed more to lignin synthesis and accumulation in tobacco leaves(p<0.01, r=0.93, 0.90, 0.91, 0.90, 0.89; P=0.24, 0.19, 0.16, 0.17, 0.19), which can be regarded as key regulatory genes for lignin synthesis and accumulation in middle leaves of flue-cured tobacco.
Lignin plays an important role in the formation of tobacco leaf resistance, but excessive content of lignin affect quality of cigarette products. In order to systematically explore the key points for lignin synthesis and accumulation in tobacco leaves at maturity stage as well as the change of regulatory genes, this study analyzes leaf hardness, tissue structure, lignin content and expression patterns of lignin synthase-related genes in middle leaves of flue-cured tobacco at maturity stage using K326 as experimental material. Results showed that maturity stage is an important period for the accumulation of lignin in flue-cured tobacco; the lignin content and hardness of tobacco leaves increased first and then decreased after entering maturity period and then reached the maximum 30 days after topping. Results of quantitative RT-PCR analysis of lignin synthase-related genes in mature tobacco leaves indicated that the expression patterns of PAL, C4 H,4CL1, CCR1, CCoAOMT5, COMT1, CAD, PAO1 and F5H1 were consistent with that of lignin changes, i.e. showing a trend of increasing first and then decreasing, and reaching its peak 30 days after topping. Among them, PAL, C4H, CCR1, CAD2 and PAO1 genes contributed more to lignin synthesis and accumulation in tobacco leaves(p<0.01, r=0.93, 0.90, 0.91, 0.90, 0.89; P=0.24, 0.19, 0.16, 0.17, 0.19), which can be regarded as key regulatory genes for lignin synthesis and accumulation in middle leaves of flue-cured tobacco.
引文
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[2]杨辉,沈火林,朱鑫,等.防御酶活性、木质素和总酚含量与辣椒抗黄瓜花叶病毒的关系[J].中国农学通报,2006(05):369-373.YANG Hui,SHEN Huolin,ZHU Xin,et al.A Study on the Relationship between the Resistance to CMV of Hot Pepper and Related Defendant Enzyme Activity,the Content of Lignins,and Total Phenols[J].Chinese Agricultural Science Bulletin,2006(05):369-373.
[3]齐曼·尤努斯.麦类根输导组织坚韧度与抗旱性的关系[J].干旱区研究,1996(03):46-49.
[4]于建军,魏登辉,田斌强,等.响应面法优化漆酶降解梗丝木质素的工艺[J].烟草科技,2017,50(03):57-64.YU Jianjun,WEI Denghui,TIAN Binqiang,et al.Optimization of lignin degradation by laccase in cut stem with response surface method[J].Tobacco Science&Technology,2017,50(3):57-64.
[5]Barros J,Serrani-Yarce J C,Chen F,et al.Role of bifunctional ammonia-lyase in grass cell wall biosynthesis[J].Nat Plants,2016,2(6):16050.
[6]Renault H,De Marothy M,Jonasson G,et al.Gene Duplication Leads to Altered Membrane Topology of a Cytochrome P450Enzyme in Seed Plants[J].Molecular Biology&Evolution,2017,34(8).
[7]Li Y,Kim J I,Pysh L,et al.Four isoforms of Arabidopsis thaliana4-coumarate:CoA ligase(4CL)have overlapping yet distinct roles in phenylpropanoid metabolism[J].Plant Physiology,2015,169(4):2409.
[8]Ruben Vanholme,Kris Morreel,Chiarina Darrah,Paula Oyarce,John H.Grabber,John Ralph,Wout Boerjan.Metabolic engineering of novel lignin in biomass crops[J].New Phytologist,2012,196(4).
[9]Boerjan W,Ralph J,Baucher M.Lignin biosynthesis[J].Annual Review of Plant Biology,2003,54(1):519-546.
[10]Sewalt V J H,Ni W,Jung H G,et al.Lignin Impact on Fiber Degradation:?Increased Enzymatic Digestibility of Genetically Engineered Tobacco(\r,Nicotiana tabacum\r,)Stems Reduced in Lignin Content[J].Journal of Agricultural and Food Chemistry,1997,45(5):1977-1983.
[11]Kitin,Peter,Voelker,Steven L,Meinzer,Frederick C,Beeckman,Hans,Strauss,Steven H,Lachenbruch,Barbara.Tyloses and Phenolic Deposits in Xylem Vessels Impede Water Transport in Low-Lignin Transgenic Poplars:A Study by Cryo-Fluorescence Microscopy1[W][OA][J].Plant Physiology,2010,154(2).
[12]孟靖,王成,赵曼琳,等.反义4CL基因调节烟草木质素生物合成[J].分子植物育种,2017,15(11):4383-4388.Jing M,Cheng W,Manlin Z,et al.Lignin Biosynthesis of Tobacco(Nicotiana tabacum L.)Regulated by the Antisense 4CLGene[J].Molecular Plant Breeding,2017,15(11):4383-4388.
[13]周培禄,刘光亮,王树声,等.低温胁迫下烟苗多酚代谢及其抗氧化能力分析[J].中国烟草科学,2018,39(05):33-39.Peilu Z,Guangliang L,Shusheng W,et al.Analysis of Polyphenol Metabolism and Antioxidant Capacity of Tobacco Seedlings under Cold Stress[J].Chinese Tobacco Science,2018,39(05):33-39.
[14]李雪平,彭镇华,高志民,等.抑制COMT基因表达对转基因烟草木质素合成的影响[J].分子植物育种,2012,10(06):689-692.The Effects of Depressing Expression of COMT on Lignin Synthesis of Transgenic Tobacco[J].Molecular Plant Breeding,2012,10(06):689-692.
[15]蒋博文,魏硕,王涛,等.采后烟叶软化与淀粉、糖代谢及相关基因表达的研究[J].中国农业科技导报,2018(03):30-38.JIANG Bowen,WEI Suo,WANG Tao,et al.Study on Softening and Metabolism of Starch and Sugar and Related Gene Expression in Postharvest Tobacco Leaves[J].Journal of Agricultural Science&Technology,2018(03):30-38.
[16]宋朝鹏,魏硕,刘相甫,等.开片状况对上部烟叶烘烤过程中失水特性的影响[J].西北农林科技大学学报(自然科学版),2017,45(07):8-14.SONG Zhaopeng,WEI Suo,LIU Xiangpu,et al.Effects of different leaf openness condition on dehydration characteristics of tobacco upper leaves during curing[J].Journal of Northwest A&FUniversity,2017,45(07):8-14.
[17]朱金峰,李广良,黄海棠.不同成熟度烟叶烘烤过程中叶片组织结构和形态的变化[J].江西农业学报,2011,23(07):53-56.ZHU Jinfeng,LI Guangliang,HUANG Haitang.Changes in Tissue Structure and Morphology of Tobacco Leaves during Curing of Tobacco Leaves with Different Maturities[J].Acta Agriculturae Jiangxi,2011,23(07):53-56.
[18]丁福章,李继新,袁有波,等.烟草不同组织总RNA的提取方法初探[J].中国农学通报,2007,23(12):98-101.DING Fuzhang,LI Jixin,YUAN Youbo,et al.Effective Isolation of Total RNA from Tobacco Tissues[J].Chinese Agricultural Science Bulletin,2007,23(12):98-101.
[19]蒋博文,王涛,宋朝鹏,等.采后烟叶碳代谢的动态变化分析[J].中国烟草科学,2018,39(01):32-41.JIANG Bowen,WANG Tao,SONG Zhaopeng,et al.Dynamic Analysis of Carbon Metabolism in Tobacco Leaves after Harvest[J].Chinese Tobacco Science,2018,39(01):32-41.
[20]John M Humphreys,Clint Chapple.Rewriting the lignin roadmap[J].Current Opinion in Plant Biology,2002,5(3).
[21]Smita Rastogi,Upendra Nath Dwivedi.Manipulation of lignin in plants with special reference to O-methyltransferase[J].Plant Science,2007,174(3).
[22]杨慧芹,王莎莎,金云峰,等.生长温度对不同生育期烟草多酚物质代谢的影响[J].基因组学与应用生物学,2015,34(09):1957-1974.Yang H,Wang S,Jin Y,et al.Effects of Different Growth Temperature on Polyphenols Metabolism in Tobacco(Nicotiana tobacum L.)Leaves at Different Growth Stages[J].Genomics&Applied Biology,2015,34(09):1957-1974.
[23]Boerjan W,Ralph J,Baucher M.Lignin biosynthesis[J].Annual Review of Plant Biology,2003,54(1):519-546.
[24]Ralph J,Lundquist K,Brunow G,et al.Lignins:Natural polymers from oxidative coupling of 4-hydroxyphenyl-propanoids[J].Phytochemistry Reviews,2004,3(1-2):29-60.
[25]徐宇强,胡轶,付凤玲,等.干旱胁迫下玉米自交系叶片木质素含量变化及其与耐旱性的关系[J].玉米科学,2007(05):72-75.XU Yuqiang,HU Yi,FU Fengling,et al.Changes of Lignin Content in Leaf of Maize Inbred Lines Under Drought Stress and Its Relationship with Drought Tolerance[J].Journal of Maize Sciences,2007(05):72-75.
[26]余朝阁,黄欣阳,李天来,等.钙对化学诱抗剂诱导番茄叶片木质素合成的影响[J].植物营养与肥料学报,2013,19(06):1445-1449.YU Chaoge,HUANG Xinyang,LI Tianlai,et al.Effect of calcium on lignin synthesis induced by chemical elicitors[J].Journal of Plant Nutrition&Fertilizer,2013,19(06):1445-1449.
[27]宋朝鹏.烤烟烟叶成熟和烘烤过程中色素变化特征及其机理研究[D].南京:南京农业大学,2010.SONG Zhaopeng.Pigment changes and their mechanisms of fluecured tobacco leaf during maturing and curing phases[D].Nanjing:Nanjing Agricultural University,2010.
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