柳叶蜡梅叶和花转录组测序及香气合成差异基因的筛选和表达
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摘要
柳叶蜡梅(Chimonanthus salicifolius)是蜡梅科蜡梅属植物,在气味上主要表现为叶揉碎后极芳香,其花香气清淡。为阐明柳叶蜡梅不同部位香气差异的关键调控基因及分子调控途径,本研究选取同时期叶和花作为生物学材料开展转录组测序、拼接序列的功能注释与分类,并使用qRT-PCR技术分析香气合成相关基因的表达。利用转录组测序和生物信息学分析,从柳叶蜡梅叶和花中共获得153 059个Unigene,注释到非冗余蛋白库(Non-Redundant Protein Sequence Database, NR)、Swiss-Prot蛋白质序列数据库(Manually Annotated and Reviewed Protein Sequence Database, Swiss-Prot)、京都基因与基因组百科全书(Kyoto Encyclopedia of Genes and Genomes, KEGG)、蛋白质家族的集合数据库(Protein Family, Pfam)、真核生物蛋白质同源簇数据库(euKaryotic Ortholog Groups, KOG)和基因本体数据库(Gene Ontology, GO)中的Unigene分别有56 964、40 746、22 173、47 858、46 196和37 029个。通过差异表达分析发现在叶和花中共有5 517个具有差异的表达基因,其中2 104个基因上调,3 413个基因下调,分别获得314个GO注释和33个显著富集代谢通路,包括7个与香气合成相关的显著富集代谢通路,共涉及33个差异酶,其中有7个差异酶基因在叶中表达高于花,而其余26个酶基因表达量低于花,在转录组水平上筛选出柳叶蜡梅叶及花香气表观性状差异的基因及调控途径。通过q RT-PCR验证,发现丙二烯氧化物环化酶基因(allene oxide cyclase, AOC)、酪氨酸转氨酶基因(tyrosine aminotransferase, TAT)、香叶基香叶基还原酶基因(geranylgeranyl reductase, CHLP)、1-脱氧-木糖-5-磷酸合酶基因(1-deoxy-D-xylulose-5-phosphate synthase,DXS)、萜品醇合成酶基因((-)-α-terpineol synthase, TES)、柠檬烯合成酶基因((R)-limonene synthase, LMS)及赤霉素3-β-双氧酶基因(gibberellin 3-beta-dioxygenase, GA3OX)在叶中表达水平高,对叶中香气成分的形成贡献度较大,可促进香气成分生物量的增加。本研究结果可为柳叶蜡梅分子生物学研究提供基因组数据库资源。
Chimonanthus salicifolius belongs to Chimonanthus of Calycanthaceae family, the leaves of Ch.salicifolius are shattered with a very fragrant smell, while its flowers smell rather lighter. In order to investigate the key regulation gene and molecular regulation pathway related aroma components in leaves and flowers of Ch. salicifolius, the transcriptome sequencing, sequencing data assembly, and functional annotation and classification of the sequences were conducted by using the leaves and flowers of Ch. salicifolius as materials in this study, and the aroma synthesis related differentially expressed genes were analyzed by qRT-PCR. By using transcriptome sequencing and bioinformatics analysis, 153 059 Unigenes were obtained by De novo assembly in leaves and flowers. Among them, 56 964, 40 746, 22 173, 47 858, 46 196 and 37 029 unigenes were assigned into NR(Non-Redundant Protein Sequence Database), Swiss-Prot(Manually Annotated and Reviewed Protein Sequence Database), KEGG(Kyoto Encyclopedia of Genes and Genomes), Pfam(Protein Family), KOG(euKaryotic Ortholog Groups) and GO(Gene Ontology), respectively. Though the differentially expressed analysis of 2 materials, 5 517 differentially expressed genes were annotated, of which2 104 genes up-regulated and 3 413 genes down-regulated. After GO and KEGG enrichment analysis, 314 GO annotations and 33 significantly enrichment metabolic pathways were obtained, including 7 aroma synthesis related significant enrichment of metabolic pathways involved 33 different enzymes, seven different enzyme genes expression in leaves were higher than that in flowers, while the remaining 26 enzyme genes expression were lower than that in flowers. Differentially expressed genes and regulation pathways of leaves and flowers of Ch. salicifolius were screened by transcriptional analysis, which showed that the genes AOC(allene oxide cyclase), TAT(tyrosine aminotransferase), CHLP(geranylgeranyl reductase), DXS(1-deoxy-D-xylulose-5-phosphate synthase), TES((-)-α-terpineol synthase), LMS((R)-limonene synthase) and GA3 OX(gibberellin 3-beta-dioxygenase) high expression in leaves of Ch. salicifolius, and they may have a greater contribution to the formation of aroma components in leaves, promoting the synthetic biomass of aroma components. This study results will provide the valuable genome data sources in molecular biology of Ch. salicifolius.
Chimonanthus salicifolius belongs to Chimonanthus of Calycanthaceae family, the leaves of Ch.salicifolius are shattered with a very fragrant smell, while its flowers smell rather lighter. In order to investigate the key regulation gene and molecular regulation pathway related aroma components in leaves and flowers of Ch. salicifolius, the transcriptome sequencing, sequencing data assembly, and functional annotation and classification of the sequences were conducted by using the leaves and flowers of Ch. salicifolius as materials in this study, and the aroma synthesis related differentially expressed genes were analyzed by qRT-PCR. By using transcriptome sequencing and bioinformatics analysis, 153 059 Unigenes were obtained by De novo assembly in leaves and flowers. Among them, 56 964, 40 746, 22 173, 47 858, 46 196 and 37 029 unigenes were assigned into NR(Non-Redundant Protein Sequence Database), Swiss-Prot(Manually Annotated and Reviewed Protein Sequence Database), KEGG(Kyoto Encyclopedia of Genes and Genomes), Pfam(Protein Family), KOG(euKaryotic Ortholog Groups) and GO(Gene Ontology), respectively. Though the differentially expressed analysis of 2 materials, 5 517 differentially expressed genes were annotated, of which2 104 genes up-regulated and 3 413 genes down-regulated. After GO and KEGG enrichment analysis, 314 GO annotations and 33 significantly enrichment metabolic pathways were obtained, including 7 aroma synthesis related significant enrichment of metabolic pathways involved 33 different enzymes, seven different enzyme genes expression in leaves were higher than that in flowers, while the remaining 26 enzyme genes expression were lower than that in flowers. Differentially expressed genes and regulation pathways of leaves and flowers of Ch. salicifolius were screened by transcriptional analysis, which showed that the genes AOC(allene oxide cyclase), TAT(tyrosine aminotransferase), CHLP(geranylgeranyl reductase), DXS(1-deoxy-D-xylulose-5-phosphate synthase), TES((-)-α-terpineol synthase), LMS((R)-limonene synthase) and GA3 OX(gibberellin 3-beta-dioxygenase) high expression in leaves of Ch. salicifolius, and they may have a greater contribution to the formation of aroma components in leaves, promoting the synthetic biomass of aroma components. This study results will provide the valuable genome data sources in molecular biology of Ch. salicifolius.
引文
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刘晶晶,王富民,刘国峰,等.2014.茶树萜类香气物质代谢谱与相关基因表达谱时空变化的关系[J].园艺学报,41(10):2094-2106.(Liu J J,Wang F M,Liu G F,et al.2014.Correlation between spatiotemporal profiles of volatile terpenoids and relevantterpenoid synthase gene expression in Camellia sinensis[J].Acta Horticulturae Sinica,41(10):2094-2106.)
刘静轩,许海峰,王得云,等.2017.两个耐贮性不同的红肉苹果株系果实硬度与香气成分及相关酶活性与基因表达差异分析[J].园艺学报,44(2):330-342.(Liu J X,Xu H F,Wang D Y,et al.2017.Changes of firmness,aroma,cell wall-modifying enzyme activities and analysis of related-gene expression in 2 red flesh apple strains during fruit storage[J].Acta Horticulturae Sinica,44(2):330-342.)
刘玲,余义勋.2008.植物香气的合成和代谢的分子生物学研究进展[J].中国生物工程杂志,28(s1):289-294.(Liu L,Yu Y X.2008.Advances in molecule biology of synthesis and metabolism of plant fragrance compounds[J].China Biotechnology,28(s1):289-294.)
吕婧.2014.烟草重要基因篇:5.烟草香气物质合成代谢相关基因[J].中国烟草科学,35(5):109-111.(Lv J.2014.Important gene of tobacco:5.The related genes for the synthesis of tobacco aroma compounds[J].Chinese Tobacco Science,35(5):109-111.)
欧阳婷,麦曦,夏红英,等.2010.柳叶蜡梅叶挥发油化学成分GC-MS分析[J].南昌大学学报,34(1):78-81.(Ou YT,Mai X,Xia H Y,et al.2010.Analysis of chemical constituents of volatile oil from Chimonathus salicifolius S.Y.Hu by GC-MS[J].Journal of Nanchang University,34(1):78-81.)
潘心禾,史小娟,蒋燕锋,等.2011.叶片中含极芳香物质的蜡梅属植物遗传多样性与亲缘关系[J].中国中药杂志,36(12):1553-1557.(Pan X H,Shi X J,Jiang Y F,et al.2011.Genetic diversity and genetic relationships of spacies containing extremely aromatic compounds in leaves of Chimonanthus[J].China Journal of Chinese Materia Medica,36(12):1553-1557.)
桑晓华,谷巍,巢建国,等.2017.茅苍术内参基因筛选及其在活性成分生物合成研究中的应用[J].植物生理学报,53(9):109-115.(Sang X H,Gu W,Chao J G,et al.2017.Selection of reference genes of Atractylodes lancea and its application in biosynthesis of active ingredients[J].Plant Physiology Journal,53(9):109-115.)
沙君雪,迟宝岩,张金波,等.2018.大猿叶甲转录组测序及生物学信息分析[J].农业生物技术学报,26(06):72-80.(Sha J X,Chi B Y,Zhang J B,et al.2018.Transcriptome sequencing and bio-information analysis of Colaphellus bowringi[J].Journal of Agricultural Biotechnology,26(7):1093-1106.)
史小娟,潘心禾,张新凤,等.2011.柳叶蜡梅叶挥发性成分的提取及GC-MS分析[J].中国实验方剂学杂志,17(09):129-132.(Shi X J,Pan X H,Zhang X F,et al.2011.Extract and analysis of volatile components in Chimonanthus salicifolius by GC-MS[J].Chinese Journal of Experimental Traditional Medical Formulae,17(09):129-132.)
孙君.2014.茉莉花香气合成相关酶基因克隆与表达分析[D].福建农林大学,导师:叶乃兴,pp.51-53.(Sun J.2014.Cloning and expression analysis of genes related to floral scent formation from Jasminum sambac[D].Fujian Agriculture and Forestry University,Supervisor:Ye N X.pp.51-53.)
孙明,吕晋慧,张启翔.2007.花香基因工程研究进展[J].安徽农业科学,35(11):3169-3171.(Sun M,Lv J H,Zhang Q X.2007.Research advance in floral fragrance genetic engineering[J].Journal of Anhui Agricultural Science,35(11):3169-3171.)
王继源,冯娇,侯旭东,等.2017.不同果袋对‘阳光玫瑰’葡萄香气组分及合成相关基因表达的影响[J].果树学报,34(1):1-11.(Wang J Y,Feng J,Hou X D,et al.2017.Effects of bagging treatments with different materials on aroma components and their biosynthetic gene expression in'Shine Muscat'grapeberry[J].Journal of Fruit Science,34(1):1-11.)
武斌,温雪玮,李衫衫,等.2018.稻曲病菌毒素对水稻幼根转录组的影响[J].农业生物技术学报,26(7):1093-1106.(Wu B,Wen X W,Li S S,et al.2018.Influences of mycotoxin of Villosiclava virens on the transcriptome of rice(Oryza sativa)seeding roots[J].Journal of Agricultural Biotechnology,26(7):1093-1106.)
吴琼,孙超,陈士林,等.2010.转录组学在药用植物研究中的应用[J].世界科学技术(中医药现代化),12(3):457-462.(Wu Q,Sun C,Chen S L,et al.2010.Application of transcriptomics in the studies of medicinal plants[J].World Science and Technology(Modernization of Traditional Chinese Medicin),12(3):457-462.)
吴宇锋,吕圭源,张志茹,等.2013.贮存时间对柳叶蜡梅挥发油及桉油精含量的影响[J].中国中药杂志,38(17):2803-2806.(Wu X F,Lv G Y,Zhang J R,et al.2013.The influence of storage time on content of volatile oil and cineole of Chimonanthus salicifolius[J].China Journal of Chinese Materia Medica,38(17):2803-2806.)
徐细建,郭睿,骆群,等.2017.中华蜜蜂幼虫肠道参考转录组的de novo组装及SSR分子标记鉴定[J].中国农业科学,50(6):1157-1166.(Xu X J,Guo R,Luo Q,et al.2017.De novo transcriptome assembly for Apiscerana cerana larval gut and identification of SSR molecular markers[J].Scientia Agricultura Sinica,50(6):1156-1166.)
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