柑橘R2R3-MYB类转录因子CitMYB20的克隆与表达分析
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摘要
为解析柑橘R2R3-MYB转录因子家族成员CitMYB20基因的分子生物学特征及受柑橘黄龙病病菌和溃疡病病菌的诱导表达情况。本研究同源克隆了不同柑橘品种中CitMYB20基因的开放阅读框序列,利用生物信息学方法预测其生物学功能并分析比较该基因在不同品种间的序列差异。在洋葱表皮细胞中瞬时表达观察CitMYB20的亚细胞定位。利用qRT-PCR技术分析黄龙病病菌胁迫和溃疡病病菌胁迫时CitMYB20的表达量变化。研究结果显示柑橘CitMYB20基因在晚锦橙、酸柚、纽荷尔脐橙和四季橘中的相似度达到99.01%,高度保守。晚锦橙中该基因含有3个外显子和2个内含子,其开放阅读框为804 bp,编码267个氨基酸残基。CitMYB20蛋白的N端高度保守,具有两个R结构域,属于典型的R2R3类型的MYB转录因子家族的成员。洋葱表皮细胞亚细胞定位分析结果表明该基因定位于细胞核中。CitMYB20基因受黄龙病病菌胁迫时,在黄龙病敏感品种晚锦橙和耐性品种酸柚中均显著上调表达,其中在耐病品种酸柚中上调表达24倍,在感病品种晚锦橙中上调表达34倍;而在溃疡病病菌胁迫时,CitMYB20基因仅在溃疡病抗性品种四季橘中显著差异表达,感病后上调表达3倍。推测该基因对黄龙病和溃疡病的应答差异可能是因为不同的调控机制造成的,构建CitMYB20基因过表达载体拟对不同病原的应答情况进行深入研究。
In order to analyze the molecular biological characteristics of the CitMYB20 gene, one member of the citrus R2 R3-MYB transcription factor family, and the induced expression in citrus affected with Huanglongbing(HLB) and citrus canker. The open reading frame sequences of the CitMYB20 gene were cloned from different citrus varieties by homology-based cloning. The CitMYB20 sequences from different citrus selections were compared and analyzed to predict the molecular characteristics with bioinformatics methods. The subcellular expression of the CitMYB20 was observed through its transient expression in onion epidermal cells. The differentially expression of CitMYB20 genes was analyzed by qRT-PCR under the biotic stresses of Huanglongbing and citrus canker. The results revealed that the similarity of CitMYB20 gene from Wanjincheng, Sour orange, Newhall Navel Orange and Calamondin was as high as 99.01%, which was highly conservative. CitMYB20 gene in Wanjincheng contained three exons and two introns, and its open reading frame included 804 bp nucleic acids, encoding 267 amino acid residues. The protein CitMYB20 was a typical R2 R3 type of MYB transcription factors with two R domains in the N-terminus. Subcellular localization analysis indicated that the gene was localized in the cell nucleus. When CitMYB20 gene was under the stress of HLB, it was significantly up-regulated not noly in HLB-sensitive citrus but also in HLB-tolerant citrus, which was up-regulated 24 times in the resistant variety Sour orange, and 34 times in the susceptible variety Wanjincheng. However, under the stress of canker, it was significantly up-regulated only in canker-resistant citrus, and the expression was up-regulated 3 times in citrus affected with canker. This indicated that the gene might have different regulatory mechanisms of responsing to Huanglongbing and citrus canker, and an overexpression vector was constructed for advanced research.
In order to analyze the molecular biological characteristics of the CitMYB20 gene, one member of the citrus R2 R3-MYB transcription factor family, and the induced expression in citrus affected with Huanglongbing(HLB) and citrus canker. The open reading frame sequences of the CitMYB20 gene were cloned from different citrus varieties by homology-based cloning. The CitMYB20 sequences from different citrus selections were compared and analyzed to predict the molecular characteristics with bioinformatics methods. The subcellular expression of the CitMYB20 was observed through its transient expression in onion epidermal cells. The differentially expression of CitMYB20 genes was analyzed by qRT-PCR under the biotic stresses of Huanglongbing and citrus canker. The results revealed that the similarity of CitMYB20 gene from Wanjincheng, Sour orange, Newhall Navel Orange and Calamondin was as high as 99.01%, which was highly conservative. CitMYB20 gene in Wanjincheng contained three exons and two introns, and its open reading frame included 804 bp nucleic acids, encoding 267 amino acid residues. The protein CitMYB20 was a typical R2 R3 type of MYB transcription factors with two R domains in the N-terminus. Subcellular localization analysis indicated that the gene was localized in the cell nucleus. When CitMYB20 gene was under the stress of HLB, it was significantly up-regulated not noly in HLB-sensitive citrus but also in HLB-tolerant citrus, which was up-regulated 24 times in the resistant variety Sour orange, and 34 times in the susceptible variety Wanjincheng. However, under the stress of canker, it was significantly up-regulated only in canker-resistant citrus, and the expression was up-regulated 3 times in citrus affected with canker. This indicated that the gene might have different regulatory mechanisms of responsing to Huanglongbing and citrus canker, and an overexpression vector was constructed for advanced research.
引文
Bai X.J.,Xu L.Z.,Jia R.R.,Zhou P.F.,Chen M.,He Y.R.,Peng A.H.,Lei T.G.,Li Q.,Yao L.X.,Chen S.C.,and Zou X.P.,2017,Cloning and expression analysis of HLB-associated salicylic acid carboxyl methyltransferase gene CsSAMT-1 in citrus,Yuanyi Xuebao(Acta Horticulturae Sinica),44(12):2265-2274(白晓晶,许兰珍,贾瑞瑞,周鹏飞,陈敏,何永睿,彭爱红,雷天刚,李强,姚利晓,陈善春,邹修平,2017,柑橘黄龙病相关水杨酸羧基甲基转移酶基因CsSAMT-1的克隆与表达分析,园艺学报,44(12):2265-2274)
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Cao Z.H.,Zhang S.Z.,Wang R.K.,Zhang R.F.,and Hao Y.J.,2013,Genome wide analysis of the apple MYB transcription factor family allows the identification of MdoMYB121 gene confering abiotic stress tolerance in plants,PLoS One,8(7):e69955
Chen Q.,Tang H.R.,Dong X.L.,Hou Y.X.,Luo Y.,Jiang Y.,and Huang Q.Y.,2009,Progress in the study of plant Myb transcription factors,Jiyinzuxue Yu Yingyong Shengwuxue(Genomics and Applied Biology),28(2):365-372(陈清,汤浩茹,董晓莉,侯艳霞,罗娅,蒋艳,黄琼瑶,2009,植物Myb转录因子的研究进展,基因组学与应用生物学,28(2):365-372)
Chen Y.H.,Zhang X.B.,Wu W.,Chen Z.L.,Gu H.Y.,and Qu L.J.,2006,Overexpression of the wounding-responsive gene AtMYB15 activates the shikimate pathway in Arabidopsis,J.Integr.Plant Biol.,48(9):1084-1095
Cheng C.Z.,Zeng J.W.,Zhong Y.,Yan H.X.,Jiang B.,and Zhong G.Y.,2013,Research progress on citrus Huanglong bing disease,Yuanyi Xueba(Acta Horticulturae Sinica),40(9):1656-1668(程春振,曾继吾,钟云,闫化学,姜波,钟广炎,2013,柑橘黄龙病研究进展,园艺学报,40(9):1656-1668)
Chezem W.R.,Memon A.,Li F.S.,Weng J.K.,and Clay N.K.,2017,SG2-type R2R3-MYB transcription factor MYB15controls defense-induced lignification and basal immunity in Arabidopsis,Plant Cell,29(8):1907-1926
Dong C.C.,Pang S.P.,Ma Y.Y.,Xie R.J.,Deng L.,and He S.L.,2017,Isolation and expression analysis of Citrus transcription factor CitMYB40,Guoshu Xuebao(Journal of Fruit Science),34(3):257-267(董翠翠,庞少萍,马岩岩,谢让金,邓烈,何绍兰,2017,柑橘CitMYB40转录因子的克隆与表达分析,果树学报,34(3):257-267)
Dubos C.,Stracke R.,Grotewold E.,Weisshaar B.,Martin C.,and Lepiniec L.,2010,MYB transcription factors in Arabidopsis,Trends Plant Sci.,15(10):573-581
He Y.X.,Yang X.D.,Xu C.,Guo D.Q.,Niu L.,Wang Y.,Li J.W.,Yan F.,and Wang Q.Y.,2018,Overexpression of a novel transcriptional repressor GmMYB3a negatively regulates salt-alkali tolerance and stress-related genes in soybean,Biochem.Biophys.Res.Commun.,498:586-591
Jia R.R.,Zhou P.F.,Bai X.J.,Chen S.C.,Xu L.Z.,Peng A.H.,Lei T.G.,Yao L.X.,Chen M.,He Y.R.,and Li Q.,2017,Gene cloning and expression analysis of canker-related transcription factor Cs BZIP40 in citrus,Zhongguo Nongye Kexue(Scientia Agricultura Sinica),50(13):2488-2497(贾瑞瑞,周鹏飞,白晓晶,陈善春,许兰珍,彭爱红,雷天刚,姚利晓,陈敏,何永睿,李强,2017,柑橘响应溃疡病菌转录因子Cs BZIP40的克隆及功能分析,中国农业科学,50(13):2488-2497)
Koops P.,Pelser S.,Ignatz M.,Klose C.,Marrocco-Selden K.,and Kretsch T.,2011,EDL3 is an F-box protein involved in the regulation of abscisic acid signalling in Arabidopsis thaliana,J.Exp.Bot.,62(15):5547-5560
Li H.L.,Guo D.,and Peng S.Q.,2014,Molecular and functional characterization of the JcMYB1,encoding a putative R2R3-MYB transcription factor in Jatropha curcas,Plant Growth Regul.,75(1):45-53
Li Y.J.,Ma Y.Y.,Xie R.J.,He S.L.,Yi S.L.,Lv Q.,Zheng Y.Q.,Wang W.,and Deng L.,2016,Isolation,expression and subcellular localization analysis of citrus transcription factor gene CitMYB22,Yuanyi Xuebao(Acta Horticulturae Sinica),43(6):1057-1068(李永杰,马岩岩,谢让金,何绍兰,易时来,吕强,郑永强,王武,邓烈,2016,柑橘转录因子基因CitMYB22的分离、表达及亚细胞定位分析,园艺学报,43(6):1057-1068)
Liu C.Y.,Long J.M.,Zhu K.J.,Liu L.L.,Yang W.,Zhang H.Y.,Li L.,Xu Q.,and Deng X.X.,2016,Characterization of a citrus R2R3-MYB transcription factor that regulates the flavonol and hydroxycinnamic acid biosynthesis,Scientific Reports,6:25352
Liu C.Y.,Wang X.,Xu Y.T.,Deng X.X.,and Xu Q.,2013,Whole genome level analysis of R2R3-MYB transcription factor family in sweet orange,Yuanyi Xuebao(Acta Horticulturae Sinica),40(S):2623(刘朝阳,王霞,徐远涛,邓秀新,徐强,2013,甜橙中R2R3-MYB转录因子家族的全基因组水平分析,园艺学报,40(S):2623)
Peng Z.C.,Gong G.Z.,Chen S.C.,Zhang G.B.,and Hong Q.B.,2010,Screening of molecular marker putatively related with citrus canker resistance,Yanyi Xuebao(Acta Horticulturae Sinica),37(3):383-389(彭祝春,龚桂芝,陈善春,张戈壁,洪棋斌,2010,柑橘溃疡病抗性相关的SSR标记筛选,园艺学报,37(3):383-389)
Ramadugu C.,Keremane M.L.,Halbert S.E.,Duan Y.P.,Roose M.L.,Stover E.,and Lee R.F.,2016,Long-term field evaluation reveals Huanglongbing resistance in citrus relatives,Plant Disease,100(9):1858-1869
Romero-Romero J.L.,Inostroza-Blancheteau C.,Orellana D.,Aquea F.,Reyes-Díaz M.,Gil P.M.,Matte J.P.,and ArceJohnson P.,2018,Stomata regulation by tissue-specific expression of the Citrus sinensis MYB61 transcription factor improves water-use efficiency in Arabidopsis,Plant Physiol.Biochem.,130:54-60
Shan T.L.,Hong Y.T.,Du L.P.,Xu H.J.,Wei X.N.,and Zhang Z.Y.,2016,Development and characterization of TaMYB86-overexpressing transgenic wheat lines with resistance to common root rot,Zuowu Xuebao(Acta Agronomica Sinica),42(10):1429-1436(单天雷,洪彦涛,杜丽璞,徐惠君,魏学宁,张增艳,2016,抗根腐病的TaMYB86过表达转基因小麦的创制与分子功能鉴定,作物学报,42(10):1429-1436)
Shi H.J.,2010,Function analysis of transcription factor AtMYB44 and At MYB15 during defense response in Arabidopsis,Dissertation for Ph.D.,Nanjing Agricultural University,Supervisor:Dong H.S.,and Hu S.B.,pp.79(时浩杰,2010,转录因子At MYB44和At MYB15在拟南芥防卫反应中的功能研究,博士学位论文,南京农业大学,导师:董汉松,胡白石,pp.79)
Stracke R.,Werber M.,and Weisshaar B.,2001,The R2R3-MYBgene family in Arabidopsis thaliana,Curr.Opin.Plant Biol.,4(5):447-456
Sun L.F.,Xing S.C.,Zhang J.,Yang J.F.,Wang X.Z.,and Dong Y.S.,2009,Function of the transcription factors in plant domestication and stress resistance,Jiyinzuxue Yu Yingyong Shengwuxue(Genomics and Applied Biology),28(3):569-577(孙丽芳,邢少辰,张君,杨建福,王兴智,董英山,2009,转录因子在植物进化和抗逆中的作用,基因组学与应用生物学,28(3):569-577)
Wang L.J.,Ran L.Y.,Hou Y.S.,Tian Q.Y.,Li C.F.,Liu R.,Fan D.,and Luo K.M.,2017,The transcription factor MYB115contributes to the regulation of proanthocyanidin biosynthesis and enhances fungal resistance in poplar,New Phytologist,215:351-367
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Xu J.,2009,Transcription factor At MYB13 inhibits Arabidopsis resistance to a bacterial pathogen,Dissertation for Ph.D.,Nanjing Agricultural University,Supervisor:Dong H.S.,pp.20-21(徐珏,2009,转录因子At MYB13抑制拟南芥防卫反应的研究,博士学位论文,南京农业大学,导师:董汉松,pp.20-21)
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Yao T.,Hu J.,Tang K.,Ran C.,Li Z.A.,and Zhou C.Y.,2010,Primary analysis on genomic diversities of Xanthomonas axnopodis pv.citri in nine provinces of China,Guoshu Xuebao(Journal of Fruit Science),27(5):819-822(姚廷山,胡军华,唐科志,冉春,李中安,周常勇,2010,利用rep-PCR技术研究我国9省柑橘溃疡病菌遗传多样性,果树学报,27(5):819-822)
Zhu F.,Luo T.,Liu C.Y.,Wang Y.,Yang H.B.,Yang W.,Zheng L.,Xiao X.,Zhang M.F.,Xu R.W.,Xu J.G.,Zeng Y.L.,Xu J.,Xu Q.,Guo W.W.,Larkin R.M.,Deng X.X.,and Cheng Y.J.,2017,An R2R3-MYB transcription factor represses the transformation ofα-andβ-branch carotenoids by negatively regulating expression of CrBCH2 and CrNCED5 in flavedo of Citrus reticulate,New Phytologist,216(1):178-192
Butelli E.,Garcia-Lor A.,Licciardello C.,Casas G.L.,Hill L.,Recupero G.R.,Keremane M.L.,Ramadugu C.,Krueger R.,Xu Q.,Deng X.X.,Fanciullino A.L.,Froelicher Y.,Navarro L.,and Martin C.,2017,Changes in anthocyanin production during domestication of citrus,Plant Physiol.,173(4):2225-2242
Cao Z.H.,Zhang S.Z.,Wang R.K.,Zhang R.F.,and Hao Y.J.,2013,Genome wide analysis of the apple MYB transcription factor family allows the identification of MdoMYB121 gene confering abiotic stress tolerance in plants,PLoS One,8(7):e69955
Chen Q.,Tang H.R.,Dong X.L.,Hou Y.X.,Luo Y.,Jiang Y.,and Huang Q.Y.,2009,Progress in the study of plant Myb transcription factors,Jiyinzuxue Yu Yingyong Shengwuxue(Genomics and Applied Biology),28(2):365-372(陈清,汤浩茹,董晓莉,侯艳霞,罗娅,蒋艳,黄琼瑶,2009,植物Myb转录因子的研究进展,基因组学与应用生物学,28(2):365-372)
Chen Y.H.,Zhang X.B.,Wu W.,Chen Z.L.,Gu H.Y.,and Qu L.J.,2006,Overexpression of the wounding-responsive gene AtMYB15 activates the shikimate pathway in Arabidopsis,J.Integr.Plant Biol.,48(9):1084-1095
Cheng C.Z.,Zeng J.W.,Zhong Y.,Yan H.X.,Jiang B.,and Zhong G.Y.,2013,Research progress on citrus Huanglong bing disease,Yuanyi Xueba(Acta Horticulturae Sinica),40(9):1656-1668(程春振,曾继吾,钟云,闫化学,姜波,钟广炎,2013,柑橘黄龙病研究进展,园艺学报,40(9):1656-1668)
Chezem W.R.,Memon A.,Li F.S.,Weng J.K.,and Clay N.K.,2017,SG2-type R2R3-MYB transcription factor MYB15controls defense-induced lignification and basal immunity in Arabidopsis,Plant Cell,29(8):1907-1926
Dong C.C.,Pang S.P.,Ma Y.Y.,Xie R.J.,Deng L.,and He S.L.,2017,Isolation and expression analysis of Citrus transcription factor CitMYB40,Guoshu Xuebao(Journal of Fruit Science),34(3):257-267(董翠翠,庞少萍,马岩岩,谢让金,邓烈,何绍兰,2017,柑橘CitMYB40转录因子的克隆与表达分析,果树学报,34(3):257-267)
Dubos C.,Stracke R.,Grotewold E.,Weisshaar B.,Martin C.,and Lepiniec L.,2010,MYB transcription factors in Arabidopsis,Trends Plant Sci.,15(10):573-581
He Y.X.,Yang X.D.,Xu C.,Guo D.Q.,Niu L.,Wang Y.,Li J.W.,Yan F.,and Wang Q.Y.,2018,Overexpression of a novel transcriptional repressor GmMYB3a negatively regulates salt-alkali tolerance and stress-related genes in soybean,Biochem.Biophys.Res.Commun.,498:586-591
Jia R.R.,Zhou P.F.,Bai X.J.,Chen S.C.,Xu L.Z.,Peng A.H.,Lei T.G.,Yao L.X.,Chen M.,He Y.R.,and Li Q.,2017,Gene cloning and expression analysis of canker-related transcription factor Cs BZIP40 in citrus,Zhongguo Nongye Kexue(Scientia Agricultura Sinica),50(13):2488-2497(贾瑞瑞,周鹏飞,白晓晶,陈善春,许兰珍,彭爱红,雷天刚,姚利晓,陈敏,何永睿,李强,2017,柑橘响应溃疡病菌转录因子Cs BZIP40的克隆及功能分析,中国农业科学,50(13):2488-2497)
Koops P.,Pelser S.,Ignatz M.,Klose C.,Marrocco-Selden K.,and Kretsch T.,2011,EDL3 is an F-box protein involved in the regulation of abscisic acid signalling in Arabidopsis thaliana,J.Exp.Bot.,62(15):5547-5560
Li H.L.,Guo D.,and Peng S.Q.,2014,Molecular and functional characterization of the JcMYB1,encoding a putative R2R3-MYB transcription factor in Jatropha curcas,Plant Growth Regul.,75(1):45-53
Li Y.J.,Ma Y.Y.,Xie R.J.,He S.L.,Yi S.L.,Lv Q.,Zheng Y.Q.,Wang W.,and Deng L.,2016,Isolation,expression and subcellular localization analysis of citrus transcription factor gene CitMYB22,Yuanyi Xuebao(Acta Horticulturae Sinica),43(6):1057-1068(李永杰,马岩岩,谢让金,何绍兰,易时来,吕强,郑永强,王武,邓烈,2016,柑橘转录因子基因CitMYB22的分离、表达及亚细胞定位分析,园艺学报,43(6):1057-1068)
Liu C.Y.,Long J.M.,Zhu K.J.,Liu L.L.,Yang W.,Zhang H.Y.,Li L.,Xu Q.,and Deng X.X.,2016,Characterization of a citrus R2R3-MYB transcription factor that regulates the flavonol and hydroxycinnamic acid biosynthesis,Scientific Reports,6:25352
Liu C.Y.,Wang X.,Xu Y.T.,Deng X.X.,and Xu Q.,2013,Whole genome level analysis of R2R3-MYB transcription factor family in sweet orange,Yuanyi Xuebao(Acta Horticulturae Sinica),40(S):2623(刘朝阳,王霞,徐远涛,邓秀新,徐强,2013,甜橙中R2R3-MYB转录因子家族的全基因组水平分析,园艺学报,40(S):2623)
Peng Z.C.,Gong G.Z.,Chen S.C.,Zhang G.B.,and Hong Q.B.,2010,Screening of molecular marker putatively related with citrus canker resistance,Yanyi Xuebao(Acta Horticulturae Sinica),37(3):383-389(彭祝春,龚桂芝,陈善春,张戈壁,洪棋斌,2010,柑橘溃疡病抗性相关的SSR标记筛选,园艺学报,37(3):383-389)
Ramadugu C.,Keremane M.L.,Halbert S.E.,Duan Y.P.,Roose M.L.,Stover E.,and Lee R.F.,2016,Long-term field evaluation reveals Huanglongbing resistance in citrus relatives,Plant Disease,100(9):1858-1869
Romero-Romero J.L.,Inostroza-Blancheteau C.,Orellana D.,Aquea F.,Reyes-Díaz M.,Gil P.M.,Matte J.P.,and ArceJohnson P.,2018,Stomata regulation by tissue-specific expression of the Citrus sinensis MYB61 transcription factor improves water-use efficiency in Arabidopsis,Plant Physiol.Biochem.,130:54-60
Shan T.L.,Hong Y.T.,Du L.P.,Xu H.J.,Wei X.N.,and Zhang Z.Y.,2016,Development and characterization of TaMYB86-overexpressing transgenic wheat lines with resistance to common root rot,Zuowu Xuebao(Acta Agronomica Sinica),42(10):1429-1436(单天雷,洪彦涛,杜丽璞,徐惠君,魏学宁,张增艳,2016,抗根腐病的TaMYB86过表达转基因小麦的创制与分子功能鉴定,作物学报,42(10):1429-1436)
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