水稻白叶枯病抗性基因Xa21的分子生物学研究进展
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摘要
由黄单胞杆菌水稻致病变种Xanthomonas oryzae pv.oryzae(Xoo)引起的白叶枯病是水稻重要细菌性病害之一。迄今,已有7个水稻白叶枯病抗性基因被克隆。Xa21是第一个被克隆的白叶枯病抗性基因,因具有广谱抗性而受到广泛的关注。对Xa21的发现、定位及克隆、表达特征、编码产物XA21的生化特性、作用与调控以及XA21介导的免疫反应模式等方面的研究结果进行综述,并对今后的研究方向进行展望。
The bacterial blight of rice caused by Xanthomonas oryzae pv. oryzae(Xoo)is one of the most destructive diseases in the world. Up to now, 7 bacterial blight resistance genes have been cloned. Xa21 is the first bacterial blight disease resistance gene cloned from rice and it has drawn great attention because of its broad spectrum resistance against Xoo. This paper briefly reviewed the discovery, mapping, cloning, expression properties of Xa21, biochemical properties, action and regulation of XA21 and XA21-mediated immunity. Future perspective on Xa21-related research were also discussed.
The bacterial blight of rice caused by Xanthomonas oryzae pv. oryzae(Xoo)is one of the most destructive diseases in the world. Up to now, 7 bacterial blight resistance genes have been cloned. Xa21 is the first bacterial blight disease resistance gene cloned from rice and it has drawn great attention because of its broad spectrum resistance against Xoo. This paper briefly reviewed the discovery, mapping, cloning, expression properties of Xa21, biochemical properties, action and regulation of XA21 and XA21-mediated immunity. Future perspective on Xa21-related research were also discussed.
引文
[1]Huang X,Kurata N,Wei X,et al.A map of rice genome variation reveals the origin of cultivated rice[J].Nature,2012,490(7421):497-501.
[2]Mew TW.Current status and future prospects of research on bacterial blight of rice[J].Annu Rev Phytopathol,1987,25(1):359-382.
[3]章琦.水稻白叶枯病抗性的遗传与改良[M].北京:科学出版社,2007:2.
[4]Ronald PC.The molecular basis of disease resistance in rice[J].Plant Mol Biol,1997,35(1-2):179-186.
[5]Yoshimura S,Yamanouchi U,Katayose Y,et al.Expression of Xa1,a bacterial blight-resistance gene in rice,is induced by bacterial inoculation[J].PNAS,1998,95:1663-1668.
[6]Iyer AS,McCouch SR.The rice bacterial blight resistance gene xa5encodes a novel form of disease resistance[J].Mol Plant Microbe Interact,2004,17:1348-1354.
[7]Chu Z,Ouyang Y,Zhang J,et al.Genome-wide analysis of defenseresponsive genes in bacterial blight resistance of rice mediated by the recessive R gene xa13[J].Mol Genet Genomics,2004,271:111-120.
[8]Song WY,Wang GL,Chen L,et al.A receptor kinase-like protein encoded by the rice disease resistance gene Xa21[J].Science,1995,270:1804-1806.
[9]王春连.水稻抗白叶枯病基因Xa23的图位克隆[D].北京:中国农业科学院研究生院,2006.
[10]Sun X,Cao Y,Yang Z,et al.Xa26,a gene conferring resistance to Xanthomonas oryzae pv.oryzae in rice,encodes an LRR receptor kinase-like protein[J].Plant J,2004,37:517-527.
[11]Gu K,Tian D,Yang F,et al.High-resolution genetic mapping of Xa27(t),a new bacterial blight resistance gene in rice,Oryza sativa L[J].Theor Appl Genet,2004,108(5):800-807.
[12]Khush GS,Bacalangco E,Ogawa T.A new gene for resistance to bacterial blight from O.longistaminate[J].Rice Genet News Lett,1990,7:121-122.
[13]Ronald PC,Albano B,Tabien R,et al.Genetic and physical analysis of the rice bacterial blight disease resistance locus,Xa21[J].Mol Gen Genet,1992,236(1):113-120.
[14]Wang GL,Holsten TE,Song WY,et al.Construction of a rice bacterial artificial chromosome library and identification of clones linked to the Xa27 disease resistance locus[J].Plant J,1995,7:525-533.
[15]Song WY,Pi LY,Wang GL,et al.Evolution of the rice Xa21disease resistance gene family[J].Plant Cell,1997,9:1279-1287.
[16]Goel RK,Gupta AK.Host age in relation to resistance in rice to bacterial blight caused by Xanthomonas campestris pv.oryzae[J].Trop Agric,1990,67:368-370.
[17]Kim KD,Hwang BK,Koh YJ.Evaluation of rice cultivars under greenhouse conditions for adult-plant resistance to Pyricularia oryzae[J].J Phytopath,1987,120:310-316.
[18]Ogawa T.Methods and strategy for monitoring race distribution and identification of resistance to bacterial leaf blight(Xanthomonas campestris pv.oryzae)in rice[J].Japan Agric Res Quart,1993,27:71-80.
[19]Yeh WH,Bonman JM,Lee EJ.Effects of temperature,leaf wetness duration,and leaf age on partial resistance to rice blast[J].J Plant Prot Trop,1989,6:223-230.
[20]Century KS,Lagman RA,Adkisson M,et al.Short communication:developmental control of Xa21-mediated disease resistance in rice[J].Plant J,1999,20(2):231-236.
[21]Zhao J,Fu J,Li X,et al.Dissection of the factors affecting development-controlled and race-specific disease resistance conferred by leucine-rich repeat receptor kinase-type R genes in rice[J].TAG Theoretical and Applied Genetics,2009,119:231-239.
[22]Park CJ,Lee SW,Chern M,et al.Ectopic expression of rice Xa21overcomes developmentally controlled resistance to Xanthomonas oryzae pv.Oryzae[J].Plant Sci,2010,179(5):466-471.
[23]万丙良,张献龙.Xa21基因的分子生物学研究进展[J].中国水稻科学,1998,12(2):115-118.
[24]白辉,李莉云,刘国振.水稻抗白叶枯病基因Xa21的研究进展[J].遗传,2006,28(6):745-753.
[25]Liu GZ,Pi LY,Walker JC,et al.Biochemical characterization of the kinase domain of the rice disease resistance receptor-like kinase XA21[J].J Biol Chem,2002,277(23):20264-20269.
[26]Xu WH,Wang YS,Liu GZ,et al.The autophosphorylated Ser686,Thr688,and Ser689 residues in the intracellular juxtamembrane domain of XA21 are implicated in stability control of rice receptorlike kinase[J].Plant J,2006,45(5):740-751.
[27]Flor HH.Inheritance of pathogenicity in Melampsora lini[J].Phytopathology,1942,32:653-669.
[28]Flor HH.Current status of the gene-for-gene concept[J].Annual Review of Phytopathology,1971,9(1):275-296.
[29]王育鹏,刘登义,李征,等.自然植物种群中病原菌与寄主植物相互作用的遗传学[J].生态学杂志,2005,24(2):190-194.
[30]Medzhitov R,Janeway CA Jr.Innate immunity:impact on the adaptive immune response[J].Curr Opin Immunol,1997,9(1):4-9.
[31]Medzhitov R.Toll-like receptors and innate immunity[J].Nat Rev Immunol,2001,1(2):135-145.
[32]Boller T,Felix G.A renaissance of elicitors:perception of microbe-associated molecular patterns and danger signals by pattern-recognition receptors[J].Annu Rev Plant Biol,2009,60:379-406.
[33]Shen Y,Sharma P,da Silva FG,et al.The Xanthomonas oryzae pv.lozengeoryzae raxP and raxQ genes encode an ATP sulphurylase and adenosine-5'-phosphosulphate kinase that are required for AvrXa21 avirulence activity[J].Mol Microbiol,2002,44(1):37-48.
[34]da Silva FG,Shen YW,Dardick C,et al.Bacterial genes involved in type I secretion and sulfation are required to elicit the rice Xa21-mediated innate immune response[J].Mol Plant Microbe Interact,2004,17(6):593-601.
[35]Burdman S,Shen Y,Lee SW,et al.RaxH/RaxR:a two-component regulatory system in Xanthomonas oryzae pv.oryzae required for AvrXa21 activity[J].Mol Plant Microbe Interact,2004,17(6):602-612.
[36]Lee SW,Han SW,Bartley LE,et al.From the Academy:Colloquium review.Unique characteristics of Xanthomonas oryzae pv.oryzae AvrXa21 and implications for plant innate immunity[J].PNAS,2006,103:18395-18400.
[37]Lee SW,Han SW,Sririyanum M,et al.A type I-secreted,sulfated peptide triggers XA21-mediated innate immunity[J].Science,2009,326(5954):850-853.
[38]Dardick C,Ronald P.Plant and animal pathogen recognition receptors signal through non-RD kinases[J].PLoS Patho,2006,2:e2.
[39]Chen X,Shang J,Chen D,et al.A B-lectin receptor kinase gene conferring rice blast resistance[J].Plant J,2006,46:794-804.
[40]Sun X,Cao Y,Yang Z,et al.Xa26,a gene conferring resistance to Xanthomonas oryzae pv.oryzae in rice,encodes an LRR receptor kinase-like protein[J].Plant J,2004,37:517-527.
[41]Chinchilla D,Zipfel C,Robatzek S,et al.A flagellin-induced complex of the receptor FLS2 and BAK1 initiates plant defence[J].Nature,2007,448:497-500.
[42]Wan J,Zhang XC,Neece D,et al.A LysM receptor-like kinase plays a critical role in chitin signaling and fungal resistance in Arabidopsis[J].The Plant Cell,2008,20:471-481.
[43]Wang GL,Ruan DL,Song WY,et al.Xa21D encodes a receptorlike molecule with a leucine-rich repeat domain that determines race-specific recognition and is subject to adaptive evolution[J].The Plant Cell,1998,10:765-779.
[44]Chen X,Chern M,Canlas PE,et al.A conserved threonine residue in the juxtamembrane domain of the XA21 pattern recognition receptor is critical for kinase autophosphorylation and XA21-mediated immunity[J].J Biol Chem,2010,285(14):10454-10463.
[45]Wang Y,Pi L,Chen X,et al.Rice XA21 binding protein 3is a ubiquitin ligase required for full Xa21-mediated disease resistance[J].The Plant Cell,2006,18:3635-3646.
[46]Peng Y,Bartley LE,Chen X,et al.OsWRKY62 is a negative regulator of basal and Xa21-mediated defense against Xanthomonas oryzae pv.oryzae in rice[J].Molecular Plant,2008,1(3):446-458.
[47]Park C,Peng Y,Chen X,et al.Rice XB15,a protein phosphatase2C,negatively regulates cell death and XA21-mediated innate immunity[J].PLoS Biology,2008,6(9):e231.
[48]Chen X,Chern M,Canlas PE,et al.An ATPase promotes autophosphorylation of the pattern recognition receptor XA21 and inhibits XA21-mediated immunity[J].PNAS,2010,107(17):8029-8034.
[49]Park CJ,Bart R,Chern M,et al.Overexpression of the endoplasmic reticulum chaperone BiP3 regulates XA21-mediated innate immunity in rice[J].PLoS One,2010,5(2):e9262.
[50]Peng Y,Bartley LE,Canlas P,et al.OsWRKY IIa transcription factors modulate rice innate immunity[J].Rice,2010,3:36-42.
[51]Park CJ,Han SW,Chen X,et al.Elucidation of XA21-mediated innate immunity[J].Cellular Microbiology,2010,12:1017-1025.
[52]Park CJ,Ronald PC.Cleavage and nuclear localization of the rice XA21 immune receptor[J].Nat Commun,2012,3:920.
[53]曾列先,黄少华,伍尚忠.IRBB21(Xa21)对广东稻白叶枯病菌5个小种的抗性反应[J].植物保护学报,2002,2(29):97-100.
[54]何翎,高利军,邓国富,等.广西水稻白叶枯病有效抗病基因和抗性亲本的筛选[J].安徽农业科学,2011,39(27):16604-16605.
[55]郑康乐,庄杰云,王汉荣.基因聚合提高了水稻对白叶枯病的抗性[J].遗传,1998(4):4-6.
[2]Mew TW.Current status and future prospects of research on bacterial blight of rice[J].Annu Rev Phytopathol,1987,25(1):359-382.
[3]章琦.水稻白叶枯病抗性的遗传与改良[M].北京:科学出版社,2007:2.
[4]Ronald PC.The molecular basis of disease resistance in rice[J].Plant Mol Biol,1997,35(1-2):179-186.
[5]Yoshimura S,Yamanouchi U,Katayose Y,et al.Expression of Xa1,a bacterial blight-resistance gene in rice,is induced by bacterial inoculation[J].PNAS,1998,95:1663-1668.
[6]Iyer AS,McCouch SR.The rice bacterial blight resistance gene xa5encodes a novel form of disease resistance[J].Mol Plant Microbe Interact,2004,17:1348-1354.
[7]Chu Z,Ouyang Y,Zhang J,et al.Genome-wide analysis of defenseresponsive genes in bacterial blight resistance of rice mediated by the recessive R gene xa13[J].Mol Genet Genomics,2004,271:111-120.
[8]Song WY,Wang GL,Chen L,et al.A receptor kinase-like protein encoded by the rice disease resistance gene Xa21[J].Science,1995,270:1804-1806.
[9]王春连.水稻抗白叶枯病基因Xa23的图位克隆[D].北京:中国农业科学院研究生院,2006.
[10]Sun X,Cao Y,Yang Z,et al.Xa26,a gene conferring resistance to Xanthomonas oryzae pv.oryzae in rice,encodes an LRR receptor kinase-like protein[J].Plant J,2004,37:517-527.
[11]Gu K,Tian D,Yang F,et al.High-resolution genetic mapping of Xa27(t),a new bacterial blight resistance gene in rice,Oryza sativa L[J].Theor Appl Genet,2004,108(5):800-807.
[12]Khush GS,Bacalangco E,Ogawa T.A new gene for resistance to bacterial blight from O.longistaminate[J].Rice Genet News Lett,1990,7:121-122.
[13]Ronald PC,Albano B,Tabien R,et al.Genetic and physical analysis of the rice bacterial blight disease resistance locus,Xa21[J].Mol Gen Genet,1992,236(1):113-120.
[14]Wang GL,Holsten TE,Song WY,et al.Construction of a rice bacterial artificial chromosome library and identification of clones linked to the Xa27 disease resistance locus[J].Plant J,1995,7:525-533.
[15]Song WY,Pi LY,Wang GL,et al.Evolution of the rice Xa21disease resistance gene family[J].Plant Cell,1997,9:1279-1287.
[16]Goel RK,Gupta AK.Host age in relation to resistance in rice to bacterial blight caused by Xanthomonas campestris pv.oryzae[J].Trop Agric,1990,67:368-370.
[17]Kim KD,Hwang BK,Koh YJ.Evaluation of rice cultivars under greenhouse conditions for adult-plant resistance to Pyricularia oryzae[J].J Phytopath,1987,120:310-316.
[18]Ogawa T.Methods and strategy for monitoring race distribution and identification of resistance to bacterial leaf blight(Xanthomonas campestris pv.oryzae)in rice[J].Japan Agric Res Quart,1993,27:71-80.
[19]Yeh WH,Bonman JM,Lee EJ.Effects of temperature,leaf wetness duration,and leaf age on partial resistance to rice blast[J].J Plant Prot Trop,1989,6:223-230.
[20]Century KS,Lagman RA,Adkisson M,et al.Short communication:developmental control of Xa21-mediated disease resistance in rice[J].Plant J,1999,20(2):231-236.
[21]Zhao J,Fu J,Li X,et al.Dissection of the factors affecting development-controlled and race-specific disease resistance conferred by leucine-rich repeat receptor kinase-type R genes in rice[J].TAG Theoretical and Applied Genetics,2009,119:231-239.
[22]Park CJ,Lee SW,Chern M,et al.Ectopic expression of rice Xa21overcomes developmentally controlled resistance to Xanthomonas oryzae pv.Oryzae[J].Plant Sci,2010,179(5):466-471.
[23]万丙良,张献龙.Xa21基因的分子生物学研究进展[J].中国水稻科学,1998,12(2):115-118.
[24]白辉,李莉云,刘国振.水稻抗白叶枯病基因Xa21的研究进展[J].遗传,2006,28(6):745-753.
[25]Liu GZ,Pi LY,Walker JC,et al.Biochemical characterization of the kinase domain of the rice disease resistance receptor-like kinase XA21[J].J Biol Chem,2002,277(23):20264-20269.
[26]Xu WH,Wang YS,Liu GZ,et al.The autophosphorylated Ser686,Thr688,and Ser689 residues in the intracellular juxtamembrane domain of XA21 are implicated in stability control of rice receptorlike kinase[J].Plant J,2006,45(5):740-751.
[27]Flor HH.Inheritance of pathogenicity in Melampsora lini[J].Phytopathology,1942,32:653-669.
[28]Flor HH.Current status of the gene-for-gene concept[J].Annual Review of Phytopathology,1971,9(1):275-296.
[29]王育鹏,刘登义,李征,等.自然植物种群中病原菌与寄主植物相互作用的遗传学[J].生态学杂志,2005,24(2):190-194.
[30]Medzhitov R,Janeway CA Jr.Innate immunity:impact on the adaptive immune response[J].Curr Opin Immunol,1997,9(1):4-9.
[31]Medzhitov R.Toll-like receptors and innate immunity[J].Nat Rev Immunol,2001,1(2):135-145.
[32]Boller T,Felix G.A renaissance of elicitors:perception of microbe-associated molecular patterns and danger signals by pattern-recognition receptors[J].Annu Rev Plant Biol,2009,60:379-406.
[33]Shen Y,Sharma P,da Silva FG,et al.The Xanthomonas oryzae pv.lozengeoryzae raxP and raxQ genes encode an ATP sulphurylase and adenosine-5'-phosphosulphate kinase that are required for AvrXa21 avirulence activity[J].Mol Microbiol,2002,44(1):37-48.
[34]da Silva FG,Shen YW,Dardick C,et al.Bacterial genes involved in type I secretion and sulfation are required to elicit the rice Xa21-mediated innate immune response[J].Mol Plant Microbe Interact,2004,17(6):593-601.
[35]Burdman S,Shen Y,Lee SW,et al.RaxH/RaxR:a two-component regulatory system in Xanthomonas oryzae pv.oryzae required for AvrXa21 activity[J].Mol Plant Microbe Interact,2004,17(6):602-612.
[36]Lee SW,Han SW,Bartley LE,et al.From the Academy:Colloquium review.Unique characteristics of Xanthomonas oryzae pv.oryzae AvrXa21 and implications for plant innate immunity[J].PNAS,2006,103:18395-18400.
[37]Lee SW,Han SW,Sririyanum M,et al.A type I-secreted,sulfated peptide triggers XA21-mediated innate immunity[J].Science,2009,326(5954):850-853.
[38]Dardick C,Ronald P.Plant and animal pathogen recognition receptors signal through non-RD kinases[J].PLoS Patho,2006,2:e2.
[39]Chen X,Shang J,Chen D,et al.A B-lectin receptor kinase gene conferring rice blast resistance[J].Plant J,2006,46:794-804.
[40]Sun X,Cao Y,Yang Z,et al.Xa26,a gene conferring resistance to Xanthomonas oryzae pv.oryzae in rice,encodes an LRR receptor kinase-like protein[J].Plant J,2004,37:517-527.
[41]Chinchilla D,Zipfel C,Robatzek S,et al.A flagellin-induced complex of the receptor FLS2 and BAK1 initiates plant defence[J].Nature,2007,448:497-500.
[42]Wan J,Zhang XC,Neece D,et al.A LysM receptor-like kinase plays a critical role in chitin signaling and fungal resistance in Arabidopsis[J].The Plant Cell,2008,20:471-481.
[43]Wang GL,Ruan DL,Song WY,et al.Xa21D encodes a receptorlike molecule with a leucine-rich repeat domain that determines race-specific recognition and is subject to adaptive evolution[J].The Plant Cell,1998,10:765-779.
[44]Chen X,Chern M,Canlas PE,et al.A conserved threonine residue in the juxtamembrane domain of the XA21 pattern recognition receptor is critical for kinase autophosphorylation and XA21-mediated immunity[J].J Biol Chem,2010,285(14):10454-10463.
[45]Wang Y,Pi L,Chen X,et al.Rice XA21 binding protein 3is a ubiquitin ligase required for full Xa21-mediated disease resistance[J].The Plant Cell,2006,18:3635-3646.
[46]Peng Y,Bartley LE,Chen X,et al.OsWRKY62 is a negative regulator of basal and Xa21-mediated defense against Xanthomonas oryzae pv.oryzae in rice[J].Molecular Plant,2008,1(3):446-458.
[47]Park C,Peng Y,Chen X,et al.Rice XB15,a protein phosphatase2C,negatively regulates cell death and XA21-mediated innate immunity[J].PLoS Biology,2008,6(9):e231.
[48]Chen X,Chern M,Canlas PE,et al.An ATPase promotes autophosphorylation of the pattern recognition receptor XA21 and inhibits XA21-mediated immunity[J].PNAS,2010,107(17):8029-8034.
[49]Park CJ,Bart R,Chern M,et al.Overexpression of the endoplasmic reticulum chaperone BiP3 regulates XA21-mediated innate immunity in rice[J].PLoS One,2010,5(2):e9262.
[50]Peng Y,Bartley LE,Canlas P,et al.OsWRKY IIa transcription factors modulate rice innate immunity[J].Rice,2010,3:36-42.
[51]Park CJ,Han SW,Chen X,et al.Elucidation of XA21-mediated innate immunity[J].Cellular Microbiology,2010,12:1017-1025.
[52]Park CJ,Ronald PC.Cleavage and nuclear localization of the rice XA21 immune receptor[J].Nat Commun,2012,3:920.
[53]曾列先,黄少华,伍尚忠.IRBB21(Xa21)对广东稻白叶枯病菌5个小种的抗性反应[J].植物保护学报,2002,2(29):97-100.
[54]何翎,高利军,邓国富,等.广西水稻白叶枯病有效抗病基因和抗性亲本的筛选[J].安徽农业科学,2011,39(27):16604-16605.
[55]郑康乐,庄杰云,王汉荣.基因聚合提高了水稻对白叶枯病的抗性[J].遗传,1998(4):4-6.