菰ZlWRKY72转录因子克隆及其在茎部膨大发育中的表达分析
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摘要
菰黑粉菌(Ustilago esculenta)侵染菰(Zizania latifolia)植株后能够诱导其茎部发生形态转变,但其肉质茎膨大发育的调节机制尚不清楚。本研究克隆获得菰WRKY转录因子72(WRKY transcription factor 72,ZlWRKY72)(Gen Bank登录号:MG365889)的全长序列,其基因序列全长为2 831 bp,含有1个内含子。c DNA全长为1 085 bp,ORF编码框为738 bp,编码245个氨基酸,具有1个典型WRKYGQK结构域和1个Cys2.His2锌指结构,进化树分析结果表明,ZlWRKY72与粳稻(Oryza sativa subsp.japonica)OsWRKY72同源性最高。通过qRT-PCR技术检测ZlWRKY72基因在其茎部膨大发育中的表达,表明ZlWRKY72在菰植株组织生长旺盛的部位表达量较高,分蘖期叶片表达量显著高于茎部(P<0.05),但孕茭起始后茎部表达量显著高于叶片(P<0.05)。茎部ZlWRKY72表达变化与菰黑粉菌菌丝侵染增殖相关,孕茭起始后正常茭白茎部表达量高于灰茭(P<0.05),并在随后的膨大发育过程中表达量逐渐降低,差异显著(P<0.05)。雄茭茎部表达量高于正常茭白,茭白茎部菰黑粉菌菌丝大量侵染增殖后ZlWRKY72表达下调。本研究初步阐明了ZlWRKY72在菰肉质茎膨大发育中的表达响应,探讨了其在菰黑粉菌侵染诱导的茎部膨大发育中的调节作用,为菰孕茭膨大发育机制研究提供了理论依据。
The growth and development of stem could be induced and changed by the infection of Ustilago esculenta in Zizania latifolia,but the regulatory of swollen stem is still unclear.In this research,Z.latifolia WRKY transcription factor 72(ZlWRKY72)(Gen Bank number: MG365889) was cloned with full DNA length of 2 831 bp with one intron.The full length of its c DNA was 1 085 bp with a 738 bp of ORF coding frame encoding 245 amino acids,the typical structure domain of WRKYGQK and a zinc finger structure of Cys2.His2 were found in ZlWRKY72.The phylogenetic analysis showed that the ZlWRKY72 has the highest homology with Oryza sativa subsp.japonica OsWRKY72.Expression analysis showed that the higher expressions of ZlWRKY72 during the development of swollen stem of Z.latifolia,liking the higher expression in leaves than stems at tillering stage seedling(P<0.05),while the higher expression in stems than leaves at stage of initiation swollen using q RT-PCR(P<0.05).The expression of ZlWRKY72 in stem was related to the successfully infection and reproduction of U.esculenta.After the initiation of swollen stem,the higher expression of ZlWRKY72 were detected in the stem of Jiaobai than grey Jiaobai(P<0.05),which was reduced significantly during the development of swollen stem in Jiaobai(P<0.05).The expression of ZlWRKY72 were higher in male Z.latifolia than Jiaobai during the initiation of swollen.It was found that the reduction expression of ZlWRKY72 occurred following with the increasing hypha reproduction of U.esculenta in stem.The expression responses and regulation of ZlWRKY72 are investigated and discussed during the development of swollen stem in Z.latifolia,which might be used in the research of swollen stem.It provides a theoretical basis for studying the mechanism of swelling development of Zizania latifolia.
The growth and development of stem could be induced and changed by the infection of Ustilago esculenta in Zizania latifolia,but the regulatory of swollen stem is still unclear.In this research,Z.latifolia WRKY transcription factor 72(ZlWRKY72)(Gen Bank number: MG365889) was cloned with full DNA length of 2 831 bp with one intron.The full length of its c DNA was 1 085 bp with a 738 bp of ORF coding frame encoding 245 amino acids,the typical structure domain of WRKYGQK and a zinc finger structure of Cys2.His2 were found in ZlWRKY72.The phylogenetic analysis showed that the ZlWRKY72 has the highest homology with Oryza sativa subsp.japonica OsWRKY72.Expression analysis showed that the higher expressions of ZlWRKY72 during the development of swollen stem of Z.latifolia,liking the higher expression in leaves than stems at tillering stage seedling(P<0.05),while the higher expression in stems than leaves at stage of initiation swollen using q RT-PCR(P<0.05).The expression of ZlWRKY72 in stem was related to the successfully infection and reproduction of U.esculenta.After the initiation of swollen stem,the higher expression of ZlWRKY72 were detected in the stem of Jiaobai than grey Jiaobai(P<0.05),which was reduced significantly during the development of swollen stem in Jiaobai(P<0.05).The expression of ZlWRKY72 were higher in male Z.latifolia than Jiaobai during the initiation of swollen.It was found that the reduction expression of ZlWRKY72 occurred following with the increasing hypha reproduction of U.esculenta in stem.The expression responses and regulation of ZlWRKY72 are investigated and discussed during the development of swollen stem in Z.latifolia,which might be used in the research of swollen stem.It provides a theoretical basis for studying the mechanism of swelling development of Zizania latifolia.
引文
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康璐瑶,邢阿宝,崔海峰,等.2017.Zl ADR1基因的克隆及其在茭白孕茭中的表达[J].农业生物技术学报,25(11):1799-1808.(Kang L Y,Xing A B,Cui H F,et al.2017.Cloning and expression of Zl ADR1 gene during the formation of swollen stem of Zizania latifolia[J].Journal of Agricultural Biotechnology,25(11):1799-1808.)
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邢阿宝,崔海峰,俞晓平,等.2017.茭白茎部CTK及ABA含量动态变化分析[J].中国计量大学学报,28(2):261-268.(Xing A B,Cui H F,Yu X P,et al.2017.Dynamic analysis of the contents of CTK and ABA in the stem of Zizania latifolia[J].Journal of China University of Metrology,28(2):261-268.)
闫宁,薛惠民,石林豫,等.2013.茭白“雄茭”和“灰茭”的形成及遗传特性[J].中国蔬菜,(16):35-42.(Yan N,Xue H M,Shi L Y,et al.2013.The growth and heritability of Zizinia latifolia plants infected with sporidial strain of Ustilago esculenta[J].China Vegetables,(16):35-42.)
应荣,崔海峰,倪方群,等.2014.杀菌剂敌磺钠及植物生长调节剂对茭白孕茭的影响[J].植物生理学报,50(7):946-952.(Ying R,Cui H F,Ni F Q,et al.2014.Effects of fungicide fenaminosulf and plant growth regulator on gall formation of Zizania latifolia[J].Plant Physiology Journal,50(7):946-952.)
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尤文雨,邹克琴,俞晓平,等.2010.菰黑粉菌的分离鉴定及其发酵液中植物激素的检测[J].菌物学报,(2):178-184.(You W Y,Zou K Q,Yu X P,et al.2010.Isolation and identification of Ustilago esculenta and detection of plant hormones in the fermentation broth[J].Mycosystema,(2):178-184.)
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Bhattarai K K,Atamian H S,Kaloshian I,et al.2010.WRKY72-type transcription factors contribute to basal immunity in tomato and Arabidopsis as well as gene-forgene resistance mediated by the tomato R gene Mi-1[J].Plant Journal,63(2):229-240.
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Choi C,Wang S H,Fang I R,et al.2015.Molecular characterization of Oryza sativa WRKY6,which binds to Wboxlike element 1 of the Oryza sativa pathogenesis-related(PR)10a promoter and confers reduced susceptibility to pathogens[J].New Phytologist,208(3):846-859.
Chi Y,Yang Y,Zhou Y,et al.2013.Protein-protein interactions in the regulation of WRKY transcription factors[J].Molecular Plant,6(2):287-300.
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Dang F,Wang Y,She J et al.2014.Overexpression of Ca WRKY27,a subgroup IIe WRKY transcription factor of Capsicum annuum,positively regulates tobacco resistance to Ralstonia solanacearum infection[J].Physiologia Plantarum,150(3):397-411.
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江解增,邱届娟,韩秀芹,等.2004.茭白生育过程中地上各部位内源激素的含量变化[J].植物科学学报,22(30):245-250.(Jiang J Z,Qiu J J,Han X Q,et al.2004.Changes of endogenous hormone contents of different parts during development of water bamboo(Zizania latifolia)[J].Journal of Wuhan Botanical Research,22(30):245-250.)
江解增,曹碚生,黄凯丰,等.2005.茭白肉质茎膨大过程中的糖代谢与激素含量变化[J].园艺学报,32(1):134-137.(Jiang J Z,Cao P S,Huang K F,et al.2005.Changes of NSC,enzymes and endogenous hormones during Zizania Gall's Swelling[J].Acta Horticulturae Sinica,32(1):134-137.)
金晔.2015.茭白茎部膨大前后应激响应的蛋白质组分析[D].硕士学位论文,中国计量大学,导师:叶子弘,pp.49-50.(Jin Y.2015.Proteomic analysis of the stress response in pre-and post-swollen stem in Zizania latifolia[D].Thesis for M.S.,China Jiliang University,Suppervisor:Ye Z H,pp.49-50.)
康璐瑶,邢阿宝,崔海峰,等.2017.Zl ADR1基因的克隆及其在茭白孕茭中的表达[J].农业生物技术学报,25(11):1799-1808.(Kang L Y,Xing A B,Cui H F,et al.2017.Cloning and expression of Zl ADR1 gene during the formation of swollen stem of Zizania latifolia[J].Journal of Agricultural Biotechnology,25(11):1799-1808.)
柯卫东,孔庆东,余家林.1997.双季茭白品种资源材料的系统聚类分析[J].华中农业大学学报,(46):598-602.(Ke W D,Kong Q D,Yu J L.1997.Hierachical clustering analysis of double season water bamboo germplasm resources[J].Journey of Huazhong Agricultural University,(46):598-602.)
李蕾,谢丙炎,戴小枫,等.2005.WRKY转录因子及其在植物防御反应中的作用[J].分子植物育种,3(3):401-408.(Li L,Xie B Y,Dai X F,et al.2005.WRKY transcription factors and their roles in plant defense responses[J].Molecular Plant Breeding,3(3):401-408.)
李帅.2016.茭白Zl-RPM1和Zl-ADR1抗病基因的克隆及表达分析[D].硕士学位论文,中国计量大学,导师:叶子弘,pp.1-2.(Li S.2016.Cloning and expression analysis of Zl-RPM1 and Zl-ADR1 resistance genes in Zizania latifolia[D].Thesis for M.S.,China Jiliang University,Suppervisor:Ye Z H,pp.1-2.)
宋钰,刘冬梅,余迪求.2008.高表达水稻WRKY72基因影响拟南芥生长素信号转导[J].云南植物研究,30(6):699-705.(Song Y,Liu D M,Yu D Q.2008.Overexpression of Os WRKY72 gene causes auxin signaling passway in Arabidopsis[J].Acta Botanica Yunnanica,30(6):699-705.)
宋钰,荆邵娟,余迪求.2009.水稻WRKY转录调控因子基因功能研究进展[J].中国水稻科学,23(5):447-455.(Song Y,Jing S J,Yu D Q.2009.Research progress on function analysis of rice WRKY genes[J].Chinese Journal of Rice Science,23(5):447-455.)
伍林涛,杜才富,张敏琴,等.2013.WRKY转录因子的结构及其在植物抗逆境胁迫中的功能[J].分子植物育种,11(4):634-638.(Wu L T,Du C F,Zhang M Q,et al.2013.The structure and function of WRKY transcription factors in abiotic and biotic stress[J].Molecular Plant Breeding,11(4):634-638.)
邢阿宝,崔海峰,俞晓平,等.2017.茭白茎部CTK及ABA含量动态变化分析[J].中国计量大学学报,28(2):261-268.(Xing A B,Cui H F,Yu X P,et al.2017.Dynamic analysis of the contents of CTK and ABA in the stem of Zizania latifolia[J].Journal of China University of Metrology,28(2):261-268.)
闫宁,薛惠民,石林豫,等.2013.茭白“雄茭”和“灰茭”的形成及遗传特性[J].中国蔬菜,(16):35-42.(Yan N,Xue H M,Shi L Y,et al.2013.The growth and heritability of Zizinia latifolia plants infected with sporidial strain of Ustilago esculenta[J].China Vegetables,(16):35-42.)
应荣,崔海峰,倪方群,等.2014.杀菌剂敌磺钠及植物生长调节剂对茭白孕茭的影响[J].植物生理学报,50(7):946-952.(Ying R,Cui H F,Ni F Q,et al.2014.Effects of fungicide fenaminosulf and plant growth regulator on gall formation of Zizania latifolia[J].Plant Physiology Journal,50(7):946-952.)
尤文雨,叶子弘,刘倩,等.2008.我国茭白的生物学研究[J].长江蔬菜,(11):35-38.(You W Y,Ye Z H,Liu Q,et al.2008.Progress of biological study on Zizania latifolia in China[J].Journal of Changjiang Vegetables,(11):35-38.)
尤文雨,邹克琴,俞晓平,等.2010.菰黑粉菌的分离鉴定及其发酵液中植物激素的检测[J].菌物学报,(2):178-184.(You W Y,Zou K Q,Yu X P,et al.2010.Isolation and identification of Ustilago esculenta and detection of plant hormones in the fermentation broth[J].Mycosystema,(2):178-184.)
周鹏飞,贾瑞瑞,陈善春,等.2017.柑橘4个WRKY转录因子基因的克隆及其响应柑橘溃疡病菌侵染的表达分析[J].园艺学报,44(3):452-462.(Zhou P F,Jia R R,Chen S C,et al.2017.Cloning and expression analysis of four citrus WRKY genes responding to Xanthomon asaxonopodis pv.citri[J].Acta Horticulturae Sinica,44(3):452-462.)
Bakshi M,Oelmüller R.2014.WRKY transcription factors:Jack of many trades in plants[J].Plant Signaling&Behavior,9(2):e27700.
Bhattarai K K,Atamian H S,Kaloshian I,et al.2010.WRKY72-type transcription factors contribute to basal immunity in tomato and Arabidopsis as well as gene-forgene resistance mediated by the tomato R gene Mi-1[J].Plant Journal,63(2):229-240.
Chan Y S,Thrower L B.1980.The host-parasite relationship between Zizania caduciflora Turcz.and Ustilago esculenta P.Henn.IV.growth substances in the host-parasite combination[J].New Phytologist,85(2):225-233.
Cheng Y,Zhou Y,Yang Y,et al.2012.Structural and functional analysis of VQ motif-containing proteins in Arabidopsis as interacting proteins of WRKY transcription factors[J].Plant Physiology,159(2):810-825.
Choi C,Wang S H,Fang I R,et al.2015.Molecular characterization of Oryza sativa WRKY6,which binds to Wboxlike element 1 of the Oryza sativa pathogenesis-related(PR)10a promoter and confers reduced susceptibility to pathogens[J].New Phytologist,208(3):846-859.
Chi Y,Yang Y,Zhou Y,et al.2013.Protein-protein interactions in the regulation of WRKY transcription factors[J].Molecular Plant,6(2):287-300.
Chung K R,Tzeng D D.2004.Nutritional requirements of the edible gall-producing fungus Ustilago esculenta[J].Journal of Biological Sciences,4(2):246-252.
Dang F,Wang Y,She J et al.2014.Overexpression of Ca WRKY27,a subgroup IIe WRKY transcription factor of Capsicum annuum,positively regulates tobacco resistance to Ralstonia solanacearum infection[J].Physiologia Plantarum,150(3):397-411.
Dong J X,Chen C H,Chen Z X.2003.Expression profiles of the Arabidopsis WRKY gene superfamily during plant defense response[J].Plant Molecular Biology,51(1):21-37.
Gl?ckner G,Eichinger L,Szafranski K,et al.2002.Sequence and analysis of chromosome 2 of Dictyostelium discoideum[J].Nature,418(6893):79-85.
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