橡胶树乳管细胞GPPS基因的克隆与表达分析
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摘要
【目的】克隆橡胶树乳管细胞短链异戊烯基合酶(GPPS)基因,分析其表达特性和编码蛋白的相互作用,为解析该类基因在天然橡胶和萜类合成中的作用提供理论参考。【方法】采用逆转录PCR(RT-PCR)和cDNA末端快速克隆(RACE)从橡胶树乳管细胞胶乳中克隆GPPS基因(HbGPPS1和HbGPPS2),利用生物信息学分析其编码蛋白的特性;采用实时荧光定量PCR(q RT-PCR)检测HbGPPS1和HbGPPS2在树皮和胶乳中的组织表达特异性、在不同橡胶树品系中的表达模式及其对割胶和外源茉莉酸甲酯(MeJA)处理的响应模式,并在原核细胞中进行表达分析;通过酵母双杂交技术检测HbGPPS1和HbGPPS2蛋白的相互作用关系。【结果】从橡胶树乳管细胞乳胶中克隆获得HbGPPS1和HbGPPS2基因,其中HbGPPS1基因编码框长度为1248 bp,编码415个氨基酸,蛋白分子量为45.9 kD,理论等电点为6.77;HbGPPS2基因编码框长度为1239 bp,编码412个氨基酸,蛋白分子量为45.6 kD,理论等电点为6.77。二者的核苷酸序列相似性为89%,且编码蛋白均含有2个典型的异戊烯基转移酶活性结构域DDXXD(D),定位于叶绿体和线粒体。HbGPPS1蛋白自身及其与HbGPPS2蛋白均能形成较弱相互作用的二聚体。qRT-PCR检测结果显示,HbGPPS1和HbGPPS2基因在胶乳中的表达量高于树皮,但均以HbGPPS2基因表达量较高,表明二者表达存在组织特异性。经MeJA处理后橡胶树胶乳中HbGPPS1和HbGPPS2基因表达量较对照高,即外源MeJA可促进内源茉莉酸的合成,激活乳管细胞内的茉莉酸信号从而上调HbGPPS1和HbGPPS2基因表达。在5个橡胶树栽培品系中,HbGPPS2基因的表达量均明显高于HbGPPS1基因,且与PR107、热研7-20-59、RRIM600和热研7-33-97干胶含量趋势一致,但二者仅在热研8-79和热研7-33-97的表达模式存在差异。HbGPPS2基因能在大肠杆菌中成功表达。【结论】HbGPPS2基因可能是乳管细胞中参与天然橡胶和萜类合成的主效基因,而HbGPPS1基因可能是功能冗余基因。HbGPPS2和HbGPPS1基因在不同橡胶树品系中的表达模式与各品系干胶含量呈正相关,可用作为干胶含量评估的筛选标记。
【Objective】The short-chain isopentenyl synthase(GPPS)genes from the laticifer cells of Hevea brasiliensis were cloned and interaction between their expression patterns and encoding proteins were further analyzed in this study for dissecting the function of the GPPS genes in the biosynthesis of natural rubber and terpenoids.【Method】Reverse transcription-PCR(RT-PCR)and rapid amplification of cDNA ends(RACE)methods were performed to clone the GPPS genes from the laticifer cells of H. brasiliensis. The character of GPPS genes encoding proteins were analyzed by bioinformatics. Moreover,real-time fluorescence quantitative PCR(q RT-PCR)was used to assess tissue-specific expression of GPPS genes from H. brasiliensis in the bark and latex and expression pattern of GPPS genes in different clones of H.brasiliensis. q RT-PCR was also used to detect response models of GPPS genes to tapping and exogenous methyl jasmonate(MeJA)treatment. And expression analysis of GPPS-encoding genes was done in prokaryotic cells. The protein interac-tions of HbGPPS1 and HbGPPS2 proteins were detected by using yeast two-hybrid technique.【Result】Genes HbGPPS1 and HbGPPS2 were successfully cloned from the latex of laticifers cells in H. brasiliensis. HbGPPS1 gene harboured a coding frame of 1248 bp in length that encoded 415 amino acids with the protein molecular weight of 45.9 kD and the theoretical isoelectric point(pI)of 6.77. While the gene HbGPPS2 harboured a coding frame of 1239 bp in length that encoded 412 amino acids with the protein molecular weight of 45.6 kD and the pI of 6.77. The similarity of their nucleotide sequence was up to 89%. The encoding proteins harbored two active prenyl transferase domain DDXXD(D)which were located in chloroplast and mitochondria. Protein interaction assay demonstrated that HbGPPS1 protein harbored relatively weak interaction with itself or with HbGPPS2 protein. Detection results of qRT-PCR revealed that both HbGPPS1 and HbGPPS2 had higher expression levels in latex than that in the bark,and the HbGPPS2 gene had higher expression levels than HbGPPS1 gene in both latex and bark,indicating their tissue-specific expression patterns. The expression levels of HbGPPS1 and HbGPPS2 genes after MeJA treatment were higher than control,suggesting that exogenous MeJA treatment could promote the biosynthesis of endogenous jasmonate acid and activate jasmonate acid signaling in laticifer cells,then up-regulate expression levels of HbGPPS1 and HbGPPS2 genes. Among the five H. brasiliensis cultivated clones,the relative expression level of HbGPPS2 gene was largely higher than that of HbGPPS1. And the expression levels of HbGPPS1 and HbGPPS2 genes were consistent with the dry rubber content tendency of PR107,CATAS7-20-59,RRIM600 and CATAS7-33-97,while showed little differences in expression pattern between CATAS8-79 and CATAS7-33-97. And HbGPPS2 gene was successfully expressed in Escherichia coli.【Conclusion】HbGPPS2 gene may be the major gene in the biosynthesis of natural rubber and terpenoids in laticifer cells,while the HbGPPS1 gene may be the functionally redundant gene. The expression patterns of HbGPPS2 and HbGPPS1 genes show a positive correlation with the dry rubber content in different clones of H. brasiliensis,so they could be used as the selective molecular marks for evaluating the dry rubber content of H. brasiliensis.
【Objective】The short-chain isopentenyl synthase(GPPS)genes from the laticifer cells of Hevea brasiliensis were cloned and interaction between their expression patterns and encoding proteins were further analyzed in this study for dissecting the function of the GPPS genes in the biosynthesis of natural rubber and terpenoids.【Method】Reverse transcription-PCR(RT-PCR)and rapid amplification of cDNA ends(RACE)methods were performed to clone the GPPS genes from the laticifer cells of H. brasiliensis. The character of GPPS genes encoding proteins were analyzed by bioinformatics. Moreover,real-time fluorescence quantitative PCR(q RT-PCR)was used to assess tissue-specific expression of GPPS genes from H. brasiliensis in the bark and latex and expression pattern of GPPS genes in different clones of H.brasiliensis. q RT-PCR was also used to detect response models of GPPS genes to tapping and exogenous methyl jasmonate(MeJA)treatment. And expression analysis of GPPS-encoding genes was done in prokaryotic cells. The protein interac-tions of HbGPPS1 and HbGPPS2 proteins were detected by using yeast two-hybrid technique.【Result】Genes HbGPPS1 and HbGPPS2 were successfully cloned from the latex of laticifers cells in H. brasiliensis. HbGPPS1 gene harboured a coding frame of 1248 bp in length that encoded 415 amino acids with the protein molecular weight of 45.9 kD and the theoretical isoelectric point(pI)of 6.77. While the gene HbGPPS2 harboured a coding frame of 1239 bp in length that encoded 412 amino acids with the protein molecular weight of 45.6 kD and the pI of 6.77. The similarity of their nucleotide sequence was up to 89%. The encoding proteins harbored two active prenyl transferase domain DDXXD(D)which were located in chloroplast and mitochondria. Protein interaction assay demonstrated that HbGPPS1 protein harbored relatively weak interaction with itself or with HbGPPS2 protein. Detection results of qRT-PCR revealed that both HbGPPS1 and HbGPPS2 had higher expression levels in latex than that in the bark,and the HbGPPS2 gene had higher expression levels than HbGPPS1 gene in both latex and bark,indicating their tissue-specific expression patterns. The expression levels of HbGPPS1 and HbGPPS2 genes after MeJA treatment were higher than control,suggesting that exogenous MeJA treatment could promote the biosynthesis of endogenous jasmonate acid and activate jasmonate acid signaling in laticifer cells,then up-regulate expression levels of HbGPPS1 and HbGPPS2 genes. Among the five H. brasiliensis cultivated clones,the relative expression level of HbGPPS2 gene was largely higher than that of HbGPPS1. And the expression levels of HbGPPS1 and HbGPPS2 genes were consistent with the dry rubber content tendency of PR107,CATAS7-20-59,RRIM600 and CATAS7-33-97,while showed little differences in expression pattern between CATAS8-79 and CATAS7-33-97. And HbGPPS2 gene was successfully expressed in Escherichia coli.【Conclusion】HbGPPS2 gene may be the major gene in the biosynthesis of natural rubber and terpenoids in laticifer cells,while the HbGPPS1 gene may be the functionally redundant gene. The expression patterns of HbGPPS2 and HbGPPS1 genes show a positive correlation with the dry rubber content in different clones of H. brasiliensis,so they could be used as the selective molecular marks for evaluating the dry rubber content of H. brasiliensis.
引文
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邓小敏,王靖,陈多坤,吴绍华,田维敏.2018.橡胶树乳管细胞HblJAZ3互作蛋白的分离鉴定[J].热带作物学报,39(2):287-292.[Deng X M,Wang J,Chen D K,Wu S H,Tian W M.2018.Preliminary screening of candidate interactors of HblJAZ3 from Hevea brasiliensis laticifer by using yeast two-hybridization methods[J].Chinese Journal of Tropical Crops,39(2):287-292.]
黄德宝,秦云霞,唐朝荣.2010.橡胶树三个品系(热研8-79、热研7-33-97和PR107)胶乳生理参数的比较研究[J].热带亚热带植物学报,18(2):170-175.[Huang D B,Qin YX,Tang C R.2010.Physiological characters of latex from three Hevea clones(Reyan 8-79,Reyan 7-33-97 and PR107)[J].Journal of Tropical and Subtropical Botany,18(2):170-175.]
鞠岩峰,张剑,吴润,师君,王砚峰.2014.天然橡胶种植现状及市场需求预测分析[J].林业资源管理,(1):152-157.[Ju Y F,Zhang J,Wu R,Shi J,Wang Y F.2014.Situation of natural rubber planting and analysis of market demand forecast[J].Forest Resources Management,(1):152-157.]
梅志刚,刘实忠,校现周,罗世巧,杨文凤,牛静明,黄荣辉.2011.巴西橡胶树胶乳化学成分的GC-MS分析[J].热带作物学报,32(2):315-319.[Mei Z G,Liu S Z,Xiao XZ,Luo S Q,Yang W F,Niu J M,Huang R H.2011.Analysis of chemical components of the latex from Hevea brasiliensis by GC-MS[J].Chinese Journal of Tropical Crops,32(2):315-319.]
屠李婵,张逸风,苏平,胡添源,童宇茹,关红雨,赵瑜君,张夏楠,袁媛,高伟,黄璐琦.2017.雷公藤牻牛儿基焦磷酸合酶基因TwGPPS克隆与表达分析[J].中国中药杂志,42(2):220-225.[Tu L C,Zhang Y F,Su P,Hu T Y,Tong Y R,Guan H Y,Zhao Y J,Zhang X N,Yuan Y,Gao W,Huang L Q.2017.Cloning and protein expression analysis of geranyl diphosphate synthase genes in Tripterygium wilfordii[J].China Journal of Chinese Materia Medica,42(2):220-225.]
王彩云,李富生,李涛,李彩霞,张晓东,王元忠.2014.滇龙胆GrGPPS基因的克隆及其序列分析与原核表达[J].中草药,45(14):2060-2068.[Wang C Y,Li F S,Li T,Li CX,Zhang X D,Wang Y Z.2014.Cloning,sequence analysis,and prokaryotic expression of GrGPPS gene in Gentiana rigescens[J].Chinese Traditional and Herbal Drugs,45(14):2060-2068.]
王靖,邓小敏,田维敏.2016.巴西橡胶树乳管细胞HblMYC3互作蛋白筛选与功能分析[J].热带作物学报,37(7):1-6.[Wang J,Deng X M,Tian W M.2016.Screening and in vitro functional analysis of the HblMYC3 interacted proteins in laticifer cells of Hevea brasiliensis[J].Chinese Journal of Tropical Crops,37(7):1-6.]
魏芳,校现周.2008.巴西橡胶树热研7-33-97?PR107?RRIM600生理特性比较[J].安徽农业科学,36(18):7561-7563.[Wei F,Xiao X Z.2008.Comparision on the physiological characters of three clones Reyan7-33-97,PR107,RRIM600 of Brazil Hevea brasiliensis[J].Journal of Anhui Agricultural Sciences,36(18):7561-7563.]
吴明,刘实忠,杨文凤,校现周,罗世巧.2013.橡胶树热研7-20-59、PR107、RRIM600品种生理特性比较[J].安徽农业科学,41(8):3465-3467.[Wu M,Liu S Z,Yang W F,Xiao X Z,Luo S Q.2013.Comparison on the physiological characters of three clones Reyan7-20-59,PR107,RRIM600 on Hevea brasiliensis[J].Journal of Anhui A-gricultural Sciences,41(8):3465-3467.]
于盱,王海棠,冯美英,梁呈元,李维林.2013.薄荷GPPS基因的克隆与表达分析[J].江西农业学报,25(7):25-29.[Yu X,Wang H T,Feng M Y,Liang C Y,Li W L.2013.Cloning and expression analysis of gene GPPS in mint[J].Acta Agriculturae Jiangxi,25(7):25-29.]
Bouvier F,Suire C,d'Harlingue A,Backhaus R A,Camara B.2000.Molecular cloning of geranyl diphosphate synthase and compartmentation of monoterpene synthesis in plant cells[J].The Plant Journal,24(2):241-252.
Burke C C,Wildung M R,Croteau R.1999.Geranyl diphosphate synthase:Cloning,expression,and characterization of this prenyltransferase as a heterodimer[J].Proceedings of the National Academy of Sciences of the United States of America,96(23):13062-13067.
Buke C,Croteau R.2002.Geranyl diphosphate synthase from Abies grandis:cDNA isolation,functional expression,and characterization[J].Archives of Biochemistry and Biophysics,405(1):130-136.
Chow K S,Mat-Isa M N,Bahari A,Ghazali A K,Alias H,Mohd-Zainuddin Z,Hoh C C,Wan K L.2012.Metabolic routes affecting rubber biosynthesis in Hevea brasiliensis latex[J].Journal of Experimental Botany,63(5):1863-1871.
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