梨火疫病菌群体感应相关基因luxS的功能分析
摘要
由梨火疫病菌Erwinia amylovora引起的蔷薇科植物的火疫病是一类毁灭性的细菌病害,造成严重的经济危害。为了有效防治病害的发生及危害,人们对其致病因子的分子机理作了大量的研究。而许多植物病原细菌毒性因子的产生都依赖于群体感应(quorum sensing,QS)系统的调节。
LuxS蛋白参与细菌群体感应信号分子autoinducer-2(AI-2)合成的最后一步。本研究在实验室已有的研究基础上构建了Erwinia amylovora strainNCPPB1665(Era NCPPB1665)的luxS突变体(Ea△luxS)并且对突变体的表型进行了初步的分析。研究发现Ea△luxS影响细菌的游动性、胞外多糖的分泌以及对双氧水的耐受能力。分别将Era NCPPB1665、Ea△luxS、功能互补菌株Ea△luxS(pPRO-luxS)和空载体对照Ea△luxS(pPROEXHta)接种梨树叶柄,发现接种突变体Ea△luxS和空载体对照Ea△luxS(pPROEXHta)的叶片发病情况比野生型和功能互补菌株Ea△luxS(pPRO-luxS)明显降低。这表明luxS基因影响Era NCPPB1665重要毒性基因的表达,因此,干扰luxS基因的表达可能能成为控制梨火疫病菌的一种潜在方式。
为了进一步明确luxS基因是否影响Era NCPPB1665中hrp基因的表达,我们在E.amylovora Ea321的致病岛中挑选了八个hrp相关基因,分别为dspE、hrpW、hrpA、hrpB、hrcJ、hrpL、hrpJ、hrpY。同时,根据Sanger Institute(www.sanger.ac.uk)网站中公布的E.amylovora基因组序列,对八个相关基因分别设计引物。提取野生型Era.NCPPB1665、突变体Ea△luxS、功能互补菌株Ea△luxS(pPRO-luxS)和空载体对照Ea△luxS(pPROEXHta)的总RNA,利用反转录差异显示技术研究luxS基因对八个hrp相关基因表达的影响情况。研究发现,luxS基因并不影响hrp相关基因的表达。
Fire blight,caused by Erwinia amlyvora,is one of the most devastating and difficult-to-control diseases.Till now,a number of researches were focused on the molecular mechanisms of the virulence in E.amlyvora for control of this disease. Interestingly,many virulence factors in different phytopathogenic bacteria are regulated by Quorum sensing(QS) system.Additionally,the autoinducer 2(AI-2) was identified in E.amylovora in our previous study.
The enzyme LuxS catalyses the last step in the production of autoinducer-2 (AI-2).In this study,the luxS mutant of Era NCPPB1665(Ea△luxS) has been constructed and the phenotypes of defined luxS mutant have been characterized based on our laboratory research.Inactivation of luxS in Era NCPPB 1665 impaired motility, extracellular polysaccharide production and tolerance to hydrogen peroxide. Inoculated Era NCPPB1665,Ea△luxS,the complemented strains Ea△luxS (pPRO-luxS) and negative control Ea△luxS(pPROEXHta) respectively to pear petiole,we found lesion length caused by Ea△luxS and Ea△luxS(pPROEXHta) was shorter then Era NCPPB1665 and Ea△luxS(pPRO-luxS).It showed that luxS gene infects expression of some virulence genes in Era NCPPB1665 and interference of luxS might be a potential way for controlling fire blight.
In order to confirm whether luxS gene affect the expression of the hrp gene in Era NCPPB1665,eight hrp related genes were chosen from E.amylovora Ea321 for further study.Meanwhile,total RNA of strain Era.NCPPB1665,Ea△luxS,Ea△luxS (pPRO-luxS) was isolated,respectively,and reverse transcription(RT)-PCR was performed to study the expression of hrp gene.Our results indicated that the mutation on luxS did not affect the expression of hrp related genes.
LuxS蛋白参与细菌群体感应信号分子autoinducer-2(AI-2)合成的最后一步。本研究在实验室已有的研究基础上构建了Erwinia amylovora strainNCPPB1665(Era NCPPB1665)的luxS突变体(Ea△luxS)并且对突变体的表型进行了初步的分析。研究发现Ea△luxS影响细菌的游动性、胞外多糖的分泌以及对双氧水的耐受能力。分别将Era NCPPB1665、Ea△luxS、功能互补菌株Ea△luxS(pPRO-luxS)和空载体对照Ea△luxS(pPROEXHta)接种梨树叶柄,发现接种突变体Ea△luxS和空载体对照Ea△luxS(pPROEXHta)的叶片发病情况比野生型和功能互补菌株Ea△luxS(pPRO-luxS)明显降低。这表明luxS基因影响Era NCPPB1665重要毒性基因的表达,因此,干扰luxS基因的表达可能能成为控制梨火疫病菌的一种潜在方式。
为了进一步明确luxS基因是否影响Era NCPPB1665中hrp基因的表达,我们在E.amylovora Ea321的致病岛中挑选了八个hrp相关基因,分别为dspE、hrpW、hrpA、hrpB、hrcJ、hrpL、hrpJ、hrpY。同时,根据Sanger Institute(www.sanger.ac.uk)网站中公布的E.amylovora基因组序列,对八个相关基因分别设计引物。提取野生型Era.NCPPB1665、突变体Ea△luxS、功能互补菌株Ea△luxS(pPRO-luxS)和空载体对照Ea△luxS(pPROEXHta)的总RNA,利用反转录差异显示技术研究luxS基因对八个hrp相关基因表达的影响情况。研究发现,luxS基因并不影响hrp相关基因的表达。
Fire blight,caused by Erwinia amlyvora,is one of the most devastating and difficult-to-control diseases.Till now,a number of researches were focused on the molecular mechanisms of the virulence in E.amlyvora for control of this disease. Interestingly,many virulence factors in different phytopathogenic bacteria are regulated by Quorum sensing(QS) system.Additionally,the autoinducer 2(AI-2) was identified in E.amylovora in our previous study.
The enzyme LuxS catalyses the last step in the production of autoinducer-2 (AI-2).In this study,the luxS mutant of Era NCPPB1665(Ea△luxS) has been constructed and the phenotypes of defined luxS mutant have been characterized based on our laboratory research.Inactivation of luxS in Era NCPPB 1665 impaired motility, extracellular polysaccharide production and tolerance to hydrogen peroxide. Inoculated Era NCPPB1665,Ea△luxS,the complemented strains Ea△luxS (pPRO-luxS) and negative control Ea△luxS(pPROEXHta) respectively to pear petiole,we found lesion length caused by Ea△luxS and Ea△luxS(pPROEXHta) was shorter then Era NCPPB1665 and Ea△luxS(pPRO-luxS).It showed that luxS gene infects expression of some virulence genes in Era NCPPB1665 and interference of luxS might be a potential way for controlling fire blight.
In order to confirm whether luxS gene affect the expression of the hrp gene in Era NCPPB1665,eight hrp related genes were chosen from E.amylovora Ea321 for further study.Meanwhile,total RNA of strain Era.NCPPB1665,Ea△luxS,Ea△luxS (pPRO-luxS) was isolated,respectively,and reverse transcription(RT)-PCR was performed to study the expression of hrp gene.Our results indicated that the mutation on luxS did not affect the expression of hrp related genes.
引文
Aldridge,P.,Metzger,M.and Geider,K.(1997) Genetics of sorbitol metabolism in Erwinia amylovora and its influence on bacterial virulence.Mol.Gen.Genet.256,611-619.
Alfano,J.R.and Collmer,A.(2004) Type Ⅲ secretion system effector proteins:double agents in bacterial disease and plant defense.Annu.Rev..Phytopathol.42,385-414.
Alfaro,J.F.,Zhang,T.,Wynn,D.P.,et aL(2004).Synthesis of LuxS inhibitors targeting bacterial cell-cell communication.Orglett.6(18):3034
Allison,D.C.,Ruiz,B.,SanJose,C.,et a.(1998) Extracellular products as mediators of the formation and detachment of Pseudomonas fluorescens biofilms.FEMS Microbiol.Lett.167(2),179-184.
Ausubel,F.M.,Brent,R.,Kingston,R.E.,et al.(1995).Short.Protocols in Molecular Biology,3th ed.(Eds.).Wiley,New York
Barny,M.A.(1995) Erwinia amylovora hrpN mutants,blocked in harpin synthesis,express a reduced virulence on host plants and elicit variable hypersensitive reactions on tobacco.Eur.J.Plant Pathol.101,333-340.
Barny,M.A.,Guinebretie're,M.H.,Marcais,B.,et al.(1990) Cloning of a large gene cluster involved in Erwinia amylovora CFBP 1430 virulence.Mol.Microbiol.4,777-786.
Bassler,B.L.,Greenberg,E.P.and Stevens,A.M.(1997).Cross-species induction of luminescence in the quorum sensing bacterium Vibrio harveyi.J.Bacteriol.179(12),4043-4045.
Bassler,B.L.,Wright,M.and Silverman,M.R.(1994).Multiple signaling systems controlling expression of luminescence in Vibrio harveyi:sequence and function of genes encoding a second sensory pathway.Mol.Microbiol.13(2),273-286.
Bassler,B.L,Wright,M.,Showalter,R.E.,et al.(1993).Intercellular signaling in Vibrio harveyi-sequence and function of genes regulating expression of luminescence.Mol Microbiol,9,773-786.
Bellemann,P.and Geider,K.(1992).Localization of transposon insertions in pathogenicity mutants of Erwinia amylovora and their biochemical characterization.J.Gen.Microbiol.138,931-940.
Bereswill,S.and Geider,K.(1997).Characterization of the rcsB gene from Erwinia amylovora and its influence on exoploysaccharide synthesis and virulence of the fire blight pathogen.J.Bacteriol.179,1354-1361.
Bereswill,S.,Bugert,P.,Bruchm(u|¨)ller,I.,et al.(1995).Identification of the fire blight pathogen,Erwinia amylovora by PCR assays with chromosomal DNA.Applied and environmental microbiology 61(7),2636-2642.
Bereswill,S.,Pahl,A.,Bellemann,P.,et al.(1992).Sensitive and species-specific detection of Erwinia amylovora by polymerase chain reaction analysis.Appl Environ Microbiol 58,3522-3526.
Bogdanove,A.J.,Bauer,D.W.and Beer,S.V.(1998).Erwinia amylovora secretes DspE,a pathogenicity factor and functional AvrE homolog,through the Hrp(type Ⅲ secretion) pathway.J.Bacteriol.180,2244-2247.
Bogdanove,A.J.,Beer,S.V.,Bonas,U.,et al.(1996).Unified nomenclature for broadly conserved hrp genes of phytopathogenic bacteria.Mol.Microbiol.20,681-683.
Bogdanove,A.J.,Kim,J.F.,Wei,Z.,et al.(1998).Homology and functional similarity of an hrp-linked pathogenicity locus,dspEF,of Erwinia amylovora and the avirulence locus avrE of Pseudomonas syringae pathovar tomato.Proc.Natl.Acad.Sci.USA 95,1325-1330.
Bogdanove,A.J.,Wei,Z.M.,Zhao,L.,et al.(1996) Erwinia amylovora secretes harpin via a type Ⅲ pathway and contains a homolog of yopN of Yersinia.J.Bacteriol.178,1720-1730.
Bonn,W.G.and van der Zwet,T.(2000).Distribution and economic importance of fire blight In:Fire Blight,the Disease and its Causative Agent,Erwinia amylovora(Vanneste,J.L.,Ed.),pp.37-54.CAB International,Wallingford.
Bugert,P.and Geider,K.(1995).Molecular analysis of the ams operon required for exopolysaccharide synthesis of Erwinia amylovora.Mol.Microbiol.15,917-933.
Burgess,N.A.,Kirke,D.F.,Williams,P.,et al.(2002).LuxS-dependent quorum sensing in Porphyromonas gingivalis modulates protease and haemagglutinin activities but is not essential for virulence.Microbiology,148,763-772.
Burrill,T.J.(1883).New species of Micrococcus.Am.Naturalist.17,319.
Burse,A.,Weingart,H.,and Ullrich,M.S.(2004).NorM,an Erwinia amylovora Multidrug Efflux Pump Involved in In Vitro Competition with Other Epiphytic Bacteria.Appl Environ Microbiol.,70(2),693-703.
Buttner,D.,Nennstiel,D.,Klusener,B.et al.(2002).Functional analysis of HrpF,a putative type Ⅲ translocon protein from Xanthomonas campestris pv.vesicatoria.J.Bacteriol.184,2389-2398.
Camara,M.,Daykin,M.,Chhabra,S.R.(1998).Detection,purification and synthesis of N-2-acylhomoserine lactone quorum sensing signal molecules[J].Methods in Microbiol,1998,27:319-330.
Cao,J.G.,Meighen,E.A.(1989).Purification and structural identification of an autoinducer for the luminescence system of Vibrio harveyi.J.Biol.Chem.264(36),21670-21676.
Cha,C.,Gao,P.,Chen,Y.C.,et al.(1998).Production of acyl-homoserine lactone quorum-sensing signals by gram-negative plant-associated bacteria.Mol Plant Microbe Interact 11(11):1119-1129.
Chen,X.,Schauder,S.,Potier,N.,et al.(2002).Structural identification of a bacterial quorum-sensing signal containing boron.Nature 415(6871):545-549.
Clarke,M.B.,Sperandio,V.(2005).Events at the host-microbial interface of the gastrointestinal tract Ⅲ.Cell-to cell signaling among microbial flora,host and pathogens:There is a whole lot of talking going on.Am J Physiol.288(6,Pt.1):G1105-G1109.
Davies,D.C.,Parsek,M.R.,Pearson,J.P.,et al.(1998).The involvement of cell-to-cell signals in the development of a bacterial biofilm.Science 280(5361),295-298.
Day,W.A.,Maurelli,A.T.(2001).Shigella flexneri LuxS quorum-sensing system modulates virB expression but is not essential for virulence.Infect Immun.69(1),15-23.
Winzer,K.,ttardie,K.R.,Burgess,N.,et al.(2002).LuxS:its role in central metabolism and the in vitro synthesis of 4-hydroxy-5-methyl-3(2H)-furanone.Microbiology.Apr;148(Pt 4),909-922.
de Kievit,T.R.,Iglewski,B.H.(2000).Bacterial quorum sensing in pathogenic relationships.Infect Immun,68(9),4839-4849.
DebRoy,S.,Thilmony,R.,Kwack,Y.B.,et al.(2004).A family of conserved bacterial effectors inhibits salicylic acid-mediated basal immunity and promotes disease necrosis in plants.Proc.Natl.Acad.Sci.USA 101,9927-9932.
Dellagi,A.,Brisset,M.N.,Paulin,J.P.,et al.(1998).Dual role of desferrioxamine in Erwinia amylovora pathogenicity.Mol.Plant Microbe Interact.11,734-742.
Doherty,N.,Holden,M.T.G.,Qazi,S.N.,et al.(2006).Functional Analysis of luxS in Staphylococcus aureus Reveals a Role in Metabolism but Not Quorum Sensing.Journal of bacteriology,188(8) 2885-2897.
Dong,H.,Delaney,T.P.,Bauer,D.W.et al.(1999).Harpin induces disease resistance in Arabidopsis through the systemic acquired resistance pathway mediated by salicylic acid and the NIM1 gene.Plant J.20,207-215.
Dong,H.P.,Peng,J.,Bao,Z.,et al.(2004).Downstream divergence of the ethylene signaling pathway for harpin-stimulated Arabidopsis growth and insect defense.Plant Physiol.136,3628-3638.
Dong,Y.H.,Xu,J.L.,Li,X.Z.,et al.(2000).AiiA,an enzyme that inactivates the acylhomoserine lactone quorum sensing signal and attenuates the virulence of Erwinia carotovora.Proc Natl Acad Sci,USA 97(7):3526-3531.
Dong,Y.H.,Wang,L.H.,Xu,J.L.,et al.(2001).Quenching quorum sensing dependent bacterial infection by an N-acyl homoserine lactonase.Nature 411:813-817.
Eastgate,J.A.(2000).Erwinia arnylovora:the molecular basis of fireblight disease.Mol Plant.Pathol.1,325-329.
Eberhard,A.,Burlingame,A.L.,Eberhard,C.,et al.(1981).Structural identification of autoinducer of Photobacterium fischeri luciferase.Biochemistry 20(9),2444-2449.
Eberl,L.,Winson,M.K.,Sternberg,C.,et al.(1996).Involvement of N-acyl-1-homoserine lactone autoinducers in controlling the multicellular behaviour of Serratia liquefaciens.Mol.Microbiol.20(1),127-136.
Eberl,L.,Winson,M.K.,Sternberg,C.,et al.(1996).Involvement of N-acyl-L-hormoserine lactone autoinducers in controlling the multicellular behaviour of Serratia liquefaciens.Mol Microbiol 20(1),127-136.
El-Maarouf,H.,Barny,M.A.,Rona,J.P.,et al..(2001).Harpin,a hypersensitive response elicitor from Erwinia amylovora,regulates ion channel activities in Arabidopsis thaliana suspension cells.FEBS Lett.497,82-84.
Federle,M.J.,Bassler,B.L.(2003) Interspecies communication in bacteria.J Clin Invest 112(9),1291-1299.
Frommberger,M.,Schmitt-Kopplin,P.,Ping,G.,et al.(2004).A simple and robust set-up for on-column sample preconcentration-nano-liquid chromatography-electrospray ionization mass spectrometry for the analysis of N-acylhomoserine lactones.Anal Bioanal Chem,378,1014-1020.
Fuqua,W.C.,Winans,S.C.and Greenberg,E.P.(1994).Quorum sensing in bacteria:the LuxR-LuxI family of cell density-responsive transcriptional regulators.J Bacteriol.176,269-75.
Gao,M.,Teplitski,M.,Robinson,J.B.,et al.(2003).Production of substances by Medicago truncatula that affect bacterial quorum sensing.Mol Plant Microbe Interact 16:827-834.
Gaudriault,S.,Malandrin,L.,Paulin,J.P.et al.(1997).DspA,an essential pathogenicity factor of Erwinia amylovora showing homology with AvrE of Pseudomonas syringae,is secreted via the Hrp secretion pathway in a DspB-dependent way.Mol.Microbiol.26,1057-1069.
Gaudriault,S.,Paulin,J.P.and Barny,M.A.(2002).The DspB/F protein of Erwinia amylovora is a type Ⅲ secretion chaperone ensuring efficient intrabacterial production of the Hrp-secreted DspA/E pathogenicity factor.Mol.Plant Pathol.3,313-320.
Geier,G.and Geider,K.(1993).Characterization and influence on virulence of the levansucrase gene from the fireblight pathogen Erwinia amylovora.Physiol.Mol.Plant Pathol.42,387-404.
Givskov,M.,de Nys,R.,Manefield,M.,et al.(1996).Eukaryotic interference with homoserine lactone2mediated prokaryotic signalling.J Bacteriol 178,6618-6622.
Goodman,R.N.,Huang,J.S.and Huang,P.Y.(1974).Hostspecific phytotoxic polysaccharide from apple tissue infected by Erwinia amylovora.Science 183,1081-1082.
Greenberg EP,Ulitzur,S and Hastings,J.W.(1979).Induction of luciferase synthesis in Beneckea harveyi by other marine bacteria.Arch Microbiol 120:87-91.
Gross,M.,Geier,G.,Rudolph,K.,et al.(1992).Levan and levansucrase synthesized by the fire blight pathogen Erwinia amylovora.Physiol.Mol.Plant Pathol.40,371-381.
Guiford,P.J.,Taylor,R.K.,Clark,R.G.,et al.(1996).PCR-based techniques for the detection of Erwinia amylovora Acta Horticultural 411:53-56.
Hakansson,S.,Schesser,K.,Persson,C.,et al.(1996).The YopB protein of Yersinia pseudotuberculosis is essential for the translocation of Yop effector proteins across the target cell plasma membrane and displays a contact-dependent membrane disrupting activity.EMBO J.15,5812-5823.
Hale,C.N.and Clark,R.G.(1990).Detection of Erwinia amylovora from apple tissue by DNA hybridization.Acta Horticulturae 273,51-55.
Hastings,J.W.,Nealson,K.H.(1977).Bacterial bioluminescence.Annu RevMicrobiol.31,549-595
Hendricks,C.L.,Ross,J.R.,Piehersky,E.,et al.(2004).An enzyme-coupled colormetric assay for S-adenosylmethionine-dependent rnethyltransferases Anal Biochem,326,100-105.
Morten,H.,Kathrin,R.,Thomas B.R.,et al.(2002).Inhibition of quorum sensing in Fsedomonas aeruginosa biofilm bacteria by a halogenated furanone compound.Microbiology (Reading,United Kingdom),148(1):87-102.
Hentzer,M.,and Givskov,M.(2003).Pharmacological inhibition of quorum sensing for the treatment of chronic bacterial infections.J Clin Invest 112:1300-1307.
Hentzer,M.,Wu,H.,Andersen,J.B.,et al.(2003).Attenuation of Pseudomonas aeruginosa virulence by quorum sensing inhibitors.EMBO J 22:3803-3815.
Hernandez-Guzman,G.and Alvarez-Morales,A.(2001).Isolation and characterization of the gene coding for the amidinotransferase involved in the biosynthesis of phaseolotoxin in Pseudomonas syringae pv.phaseolicola.MoL Plant-Microbe Interact.14,545-554.
Holden,M.T.G.,Ram C.S.,De N.R.,et al.(1999).Quorum-sensing cross talk:isolation and chemical characterization of cyclic dipeptides from Psedomonas aeruginosa and other Gramnegative bacteria.Mol Microbiol.33(6):1254-1266.
Hubner,A.,Revel,A.T.,Nolen,D.M.,et al.(2003).Expression of a luxS gene is not required for Borrelia burgdorferi infection of mice via needle inoculation.Infect Immun 71(5),2892-2896.
Jin,Q.,MeGhee,G.C.,Hart,P.,et al.(2002).Secretion of Hrp and Dsp proteins via the Hrp pilus during type Ⅲ secretion in Erwinia amylovora.Acta Hort.590,441-448.
Johnson,K.B.,and Stockwell,V.O.(1998).Management of fire blight:a case study in microbial ecology.Annu.Rev.Phytopathol.36,227-248.
Joint,I.,Tait,K.,Callow,M.E.,et al.(2002).Cell-to-cell communication across the prokaryote-eukaryote boundary.Science.Nov 8;298(5596):1207.
Kachadourian,R.,Dellagi,A.,Laurent,J.,et al.(1996).Desferrioxamine-dependent iron transport in Erwinia amylovora CFBP 1430:cloning of the gene encoding the ferrioxamine receptor FoxR.Biometals 9,143-150.
Karen,T.E.,and Park,S.F.(2002).Quorum sensing in Campylobacter jejuni:detection of a luxS encoded signaling molecule.Microbiology.148,1475-1481
Kaufmann,G.F.,Sartorio,R.,Lee Ss,H.,et al.(2005).Revisiting quorum sensing:Discovery of additional chemical and diological functions for 3-oxo-N-acyl-homoserine lactones.Proc Natl Acad Sci USA.102(2):309-314.
Kelm,O.,Kiecker,C.,Geider,K.et al.(1997).Interaction of the regulator proteins RcsA and RcsB with the promoter of the operon for amylovoran biosynthesis in Erwinia amylovora.Mol.Gen.Genet.256,72-83.
Kim,J.F.(1997).Molecular characterization of a novel harpin and two hrp secretery operons of Erwinia amylovora,and a hrp operon of E.chrysanthemi.Ph.D.Cornell University,Ithaca,NY.
Kim,J.F.and Beer,S.V.(1998).HrpW of Erwinia amylovora,a new harpin that contains a domain homologous to pectate tyases of a distinct class.J.Bacteriol.180,5203-5210.
Kim,J.F.,Wei,Z.-M.and Beer,S.V.(1997).The hrpA and hrpC operons of Erwinia amylovora encode components of a type Ⅲ pathway that secretes harpin.J.Bacteriol.179,1690-1697.
Kim,W.-S.,Bonasera,J.M.,Meng,X.,et al.(2006).Interaction of DspE/A,a pathogenicity/avirulence protein of Erwinia amylovora,with pre-ferredoxin from apple and its relationship to photosynthetic efficiency.Acta Hort.(in press).
Klars W,Kim R H.(2002).LuxS:its role in central metabolism and the in vitro synthesis of 4-hydroxy-5-methyl-3(2H)-furanone.Microbiology.148:909.
Kleerebezem,M.,Quadri,L.E.,Kuipers,O.P.,et al.(1997).Quorum sensing by peptide pheromones and two-component signal-transduction systems in Gram-positive bacteria.Mol.Microbiol 24(5),895-904.
Lewis,H.A.,Furlong,E.B.,Laubert,B.,et al.(2001).A structural genomics approach to the study of quorum sensing:crystal structures of three LuxS orthologs.Structure.9(6),527-37.
Maes,M.,Garbeva,P.,Crepel,C.(1996).Identification and sensitive endophytic detection of the fire blight pathogen Erwinia amylovora with 23S ribosomal DNA sequences and the polymerase chain reaction.Plant Pathology 45,1139-1149.
Marcus,B.J.,and Blaser,M.J.(2003).Detection of a luxS-Signaling Molecule in Bacillus anthracis.Infection and Immunity.71(7),3914-3919.
Mathesius,U.,Mulders,S.,Gao,M.,et al.(2003).Extensive and specific responses of a eukaryote to bacterial quorum sensing signals.Proc Natl A cad Sci USA 100(3),1444-1449.
Mayville,P.,Ji,G.,Beavis,R.,et al.(1999).Structure-activity analysis of synthetic autoinducing thiolactone peptides from Staphylococcus aureus responsible for virulence.Proc Natl Acad Sci USA.96:1218-1223.
MeClean,K.H.,Winson,M.K.(1997).Quorum sensing and Chromobacterium violaceum:exploitation of violacein production and inhibition for the detection of N-acyl-homoserine lactones.Microbiology 143,3703-3711.
McManus,P.S.and Jones,A.L.(1995).Detection of Erwinia amylovora by nested PCR and PCR-dot-blot and reverse-blot hybridizations.Phytopat.hology 85,618-623.
Meng,X.,Bonasera,J.M.,Kim,J.F.,et al.(2006).Apple proteins that interact with DspE,a pathogenicity effector of Erwinia amylovora,the fire blight pathogen.Mol.Plant-Microbe Interact.(in press).
Merighi,M.,Sandrini,A.(2000).Chemiluminescent and Colorimetric Detection of Erwinia amylovora by Immunoenzymatic Determination of PCR Amplicons from Plasmid pEA29 Plant Disease 84(1),49-54.
Miller,M.B.,Skorupski,K.,et al.(2002).Parallel quorum sensing systems converge to regulate virulence in Vibrio cholerae.Cell.110(3),303-314.
Miller,S.T.,Xavier,K.B.,Campagna,S.R.,et al.(2004) Salmonella typhimurium Recognizes a Chemically Distinct Form of the Bacterial Quorum-Sensing Signal AI-2.Mol.Cell 15,677-687.
Miller,M.B.,Bassler,B.L.(2001) Quorum sensing in bacteria.Annu Rev Microbiol.55,165-199.
Mohammadi,M.,Geider,K.(2007).Autoinducer-2 of the fire blight pathogen Erwinia amylovora and other plant associated bacteria,FEMS Microbiol Lett 266:34-41.
Mok,K.C.,Wingreen,N.S.,Bassler,B.L.(2003).Vibrio harveyi quorum sensing:a coincidence detector for two autoinducers controls gene expression The EMBO Journal 2(4),870-881.
Molina,L.,Constantineseu,F.,Michel,L.,et al.(2003).Degradation of pathogen quorum-sensing molecules by soil bacteria:a preventive and curative biological control mechanism.FEMS Microbiol Ecol 45:71-81.
Molina,L.,Rezzonico,F.,Defago,G.et al.(2005).Autoinduction in Erwinia amylovora:Evidence of an Acyl-Homoserine Lactone Signal in the Fire Blight Pathogen.Journal of Bacteriology.187,3206-3213.
Oh,C.-S.(2005).Characterization of HrpN-interacting proteins from plants,the Hrp pathogenicity island of Erwinia amylovora,and its proteins that affect the hypersensitive response.Ph.D.Cornell University,Ithaca,NY.
Oh,C.-S.,Kim,J.F.and Beer,S.V.(2005) The Hrp pathogenicity island of Erwinia amylovora and the identification of three novel genes required for systemic infection.Mol.Plant Pathol.6,125-138.
Oh,C.-S.and Beer,S.V.(2005).Molecular genetics of Erwinia amylovora involved in the development of fire blight FEMS Microbiology Letters 253,185-192.
Peng,J.-L.,Dong,H.-S.,Dong,H.-P.,et al.(2003).Harpin-elicited hypersensitive cell death and pathogen resistance requires the NDR1 and EDS1 genes.Physiol.Mol.Plant Pathol.62,317-326.
Perino,C.,Gaudriault,S.,Vian,B.,et al.(1999).Visualization of harpin secretion in planta during infection of apple seedlings by Erwinia amylovora.Cell Microbiol.1,131-141.
Petnieki-Oewieja,T.,van Dijk,K.and Alfano,J.R.(2005).The hrpK operon of Pseudomonas syringae pv.tomato DC3000 encodes two proteins secreted by the type Ⅲ(Hrp) protein secretion system:HopB1 and HrpK,a putative type Ⅲ translocator.J.Bacteriol.187,649-663.
Ramos,A.R.(2004).Hrp proteins and harpins:defining their roles in the type Ⅲ protein secretion system in Pseudomonas syringae.Ph.D.Cornell University,Ithaca,NY.
Raseh,M.,Rasmussen,T.B.(2007).Andersen J B.Well-known quorum sensing inhibitors do not affect bacterial quorum sensing-regulated bean spourt spoilage.J Appl microbial.102(3):826-837.
Ratledge,C.and Dover,L.G.(2000).Iron metabolism in pathogenic bacteria.Annu.Rev.Microbiol.54,881-941.
Ren,D.,Sins,J.J.(2001).Wood T K.Inhibition of biofilm formation and swarming of Escherichia coli by(5Z)-4-bromo-5-(bromomethylene)-3-buty-2(5H)-furanone.Environ Microbiol.3:731
Ren,D.,Sins J.J.(2001).Wood T K.Inhibition of biofilm formation and swarming of Bacillus subtilis by(5Z)-4-bromo-5-(bromomethylene)-3-buty-2(5H)-furanone.Lett Appl Microbiol,34:293.
Ren,D.,Sims J.J.,and Wood,T.K.(2002).Inhibition of biofilm formation and swarming of Bacillus subtilis by(5Z)-4-bromo-5-(bromomethylene)-3- butyl-2(5H)-furanone.Lett.Appl.Microbiol.34,293-299.
Ren,D.,Sims,J.J.and Wood,T.K.(2001).Inhibition of biofilm formation and swarming of Escherichia coli by(5Z)-4-bromo-5-(bromomethylene)-3-butyl-2(5H)-furanone.Environ.Microbiol.3,731-736.
Rezzonico F.& Duffy B.(2007).The role of luxS in the fire blight pathogen Erwinia amyIovora is limited to metabolism and does not involve quorum sensing.Molecular Plant-Microbe Interactions 20:1284-1297.
Samebrook,J.,Russell,D.W.(2001).Molecular cloning:a laboratory mannual,3~(rd) ed.Cold Spring Harbor Laboratory.Press,NY.
Sehauder,S.,Bassler,B.L.(2001).The languages of bacteria.Genes Dev 15(12),1468-1480.
Schmidt,H.and Hensel,M.(2004).Pathogenicity islands in bacterial pathogenesis.Clin.Microbiol.Rev.17,14-56.
Sehnabel,E.L.,and Jones,A.L.(1998).Instability of a pEA29 marker in Erwinia amylovora previously used for strain classification.Plant Dis.83,1334-1336.
Seemuller,E.A.and Beer,S.V.(1976).Absence of cell wall polysaccharide degradation of Erwinia amylovora.Phytopathology 66,433-436.
Shaw,P.D.,Ping,G.,Daly,S.L.,et al.(1997) Detecting and characterizing N—acyl-homoserine lactone signal molecules by thin-layer chromatography.Proc.Natl Acad Sci USA 94(12),6036-6041.
Shiner E K,Rumbaugh K P,S C.(2005).Interkingdom signaling:deciphering the language of acyl homoserine lactones.FEMS Microbiol Rev.29(5):935-947
Sjulin,T.M.and Beer,S.V.(1978).Mechanism of wilt induction by amylovoran in Cotoneaster shoots and its relation to wilting of shoots infected by Erwinia amylovora.Phytopathology 68,89-94.
Smith K M,Bu Y,Suga,H.(2003).Induction and inhibition of Pseudomonas aeruginosa quorum sensing by synthetic autoinducer analogs.Chem Biol.10(1):81-89
Sperandio,V.,Torres,A.G.,Jarvis,B.,et al.(2003).Bacteria-host communication:The language of hormones PNAS 100(15),8951-8956
Staskawicz,B.J.,Mudgett,M.B.,Dangl,J.L.et al.(2001).Common and contrasting themes of plant and animal diseases.Science 292,2285-2289.
Steinberger,E.M.and Beer,S.V.(1988).Creation and complementation of pathogenicity mutants of Erwinia amylovora.Mol.Plant-Microbe Interact.1,135-144.
Steinbrenner,B.,Belleman,P.,Zeller,W.,et al.(1990) DNA-hybridization,a specific method for the diagnosis of fireblight.Acta Horticulturae 273,91-93.
Sun,J.,Daniel,R.,Wagner-D(o|¨)bler,I.,et al.(2004).Is autoinducer-2 a universal signal for interspecies communication:a comparative genomic and phylogenetic analysis of the synthesis and signal transduction pathways.BMC Evolutionary Biology,4:Art.No.36.
Surette,M.G.& Bassler,B.L.(1998).Quorum-sensing in Escherichia coli and Salmonella typhimurium.Proc Natl Acad Sci USA 95:7046-7050.
Surette,M.G.& Bassler,B.L.(1999).Regulation of autoinducer production in Salmonella typhimurium.Mol Microbiol 31:585-595.
Swift,S.,Winson,M.K.,Chan,P.F.,et al.(1993).A novel strategy for the isolation of luxl homologues:evidence for the widespread distribution of a LuxR:LuxI superfamily in enteric bacteria.Mol Microbiol.10(3),511-20.
Swift,S.,Downie,J.A.,Whitehead,N.A.,et al.(2001).Quorum sensing as a population-density-dependent determinant of bacterial physiology.Adv Microb Phylosiol.45,199-270
Taga,M.E.and Bassler,B.L.(2003) Chemical communication among bacteria.Proc Natl Acad Sci USA.100(Suppl 2),14549-14554.
Van der Zwet,T.and Beer,S.V.(1991).Fire Blight:Its nature,prevention,and control-A practical guide to integrated disease management.US Dep.Agric.,Agric.Inform.Bull.No.631,34PP.
Van der Zwet,T.,Keil,H.L.(1979).Fire Blight:a bacterial disease of rosaceous plants.Agriculture handbook no.510.U.S.Government Printing Office,Washington,D.C..
Vanneste,J.L.(2000).Fire blight:the disease and its causative agent,Erwinia amylovora.CABI Publishing,Wallingford,Oxon,United Kingdom.
Vanneste,J.L.,Eden-Green,S.(2000).Migration of Erwinia amylovora in host plant tissue,p.9-36.In J.L.Vanneste(ed.),Fire Blight:the disease and its causative agent,Erwinia amylovora.CABI Publishing,Oxon,United Kingdom.
Venisse,J.S.,Barny,M.A.,Paulin,J.P.et al.(2003).Involvement of three pathogenicity factors of Erwinia amylovora in the oxidative stress associated with compatible interaction in pear.FEBS Lett.537,198-202.
Venisse,J.S.,Gullner,G.and Brisset,M.N.(2001).Evidence for the involvement of an oxidative stress in the initiation of infection of pear by Erwinia amylovora.Plant Physiol.125,2164-2172.
Venturi,V.,Venuti,C.,Deveseovi,G.,et al.(2004).The plant pathogen Erwinia amylovora produces acyl-homoserine lactone signal molecules in vitro and in planta.FENS Microbiology letters.241,179-183
Wang,L.(2004).Virulence genes of Erwinia amylovora:explorations,Ph.D.Cornell University,Ithaca,NY.
Wei,Z.,Kim,J.F.and Beer,S.V.(2000).Regulation of hrp genes and type Ⅲ protein secretion in Erwinia amylovora by HrpX/HrpY,a novel two-component system,and HrpS.Mol.Plant-Microbe interact.13,1251-1262.
Wei,Z.-M.and Beer,S.V.(1995).hrpL activates Erwinia amylovora hrp gene transcription and is a member of the ECF subfamily of s factors.J.Bacteriol.177,6201-6210.
Wei,Z.M.,Laby,R.J.,Zumoff,C.It.,et al.(1992).Harpin,elicitor of the hypersensitive response produced by the plant pathogen Erwinia amylovora.Science 257,85-88.
Wei,Z.,Kim,J.F.,Beer,S.V.(2000).Regulation of hrp genes and type Ⅲ protein secretion in Erwinia amylovora by HrpX\HrpY,a novel two-component system and HrpS.Mol.Plant Microbe Interact.13(11),1251-1262.
Winson,M.K.,Swift,S.,Fish,L.,et al.(1998).Construction and analysis of luxCDABE-based plasmid sensors for investigating N-acyl homoserine lactone-mediated quorum sensing.FEMS Microbiol lett.163(2),185-92.
Winzer,K.,Williams,P.(2001).Quorum sensing and the regulation of virulence gene expression in pathogenic bacteria.Int J Med Microbiol.291(2),131-43.
Xavier,K.B.,Bassler,B.L.(2003).LuxS quorum sensing:more than just a numbers game.CurrOp inMicrobiol 6,191-197.
Xie,Z.and Chen,Z.(2000).Harpin-induced hypersensitive cell death is associated with altered mitochondrial functions in tobacco cells.Mol.Plant-Microbe Interact.13,183-190.
Zaccardelfi,M.,and Bazzi,C.(1997).Reliability of PCR with primers from plasmid pEA29 for the detection of Erwinia amylovora.Phytopathol.Mediterr.36,116-120.
Zhang,Y.and Geider,K.(1999).Molecular analysis of the rlsA gene regulating levan production by the fireblight pathogen Erwinia amylovora.Physiol.Mol.Plant Pathol.54,187-201.
Zhang,Y.,Bak,D.D.,Heid,H.,et al.(1999).Molecular characterization of a protease secreted by Erwinia amylovora.J.Mol.Biol.289,1239-1251.
Zhang,L.H.,Murphy,P.J.,Kerr,A.,et al.(1993).Agrobacterium conjugation and gene regulation by N-acyl-1-homoserine lactones.Nature,362(6419),446-447.
李华林,闻玉梅(2004) AI-2,一种新的细菌自体诱导分子,生命科学,16(3):138-143
钱国良,胡白石,卢玲,刘凤权,许志刚(2006)梨火疫病菌的实时荧光PCR检测,植物病理学报,36(2):123-128
陶金莉,迟莉丽,沈亚领,魏东芝(2004)细菌的群体行为调控机制-Quorum sensing,微生物学通报 31(4),106-110
吴红,宋志军,Niels Hoiby(2003)细菌与细菌之间的信息交流——革兰氏阴性细菌的Quorum-sensing系统,自然科学进展,13(7),679-687
高岩,梨火疫病菌群体感应系统的初步研究,硕士学位论文,2006
Alfano,J.R.and Collmer,A.(2004) Type Ⅲ secretion system effector proteins:double agents in bacterial disease and plant defense.Annu.Rev..Phytopathol.42,385-414.
Alfaro,J.F.,Zhang,T.,Wynn,D.P.,et aL(2004).Synthesis of LuxS inhibitors targeting bacterial cell-cell communication.Orglett.6(18):3034
Allison,D.C.,Ruiz,B.,SanJose,C.,et a.(1998) Extracellular products as mediators of the formation and detachment of Pseudomonas fluorescens biofilms.FEMS Microbiol.Lett.167(2),179-184.
Ausubel,F.M.,Brent,R.,Kingston,R.E.,et al.(1995).Short.Protocols in Molecular Biology,3th ed.(Eds.).Wiley,New York
Barny,M.A.(1995) Erwinia amylovora hrpN mutants,blocked in harpin synthesis,express a reduced virulence on host plants and elicit variable hypersensitive reactions on tobacco.Eur.J.Plant Pathol.101,333-340.
Barny,M.A.,Guinebretie're,M.H.,Marcais,B.,et al.(1990) Cloning of a large gene cluster involved in Erwinia amylovora CFBP 1430 virulence.Mol.Microbiol.4,777-786.
Bassler,B.L.,Greenberg,E.P.and Stevens,A.M.(1997).Cross-species induction of luminescence in the quorum sensing bacterium Vibrio harveyi.J.Bacteriol.179(12),4043-4045.
Bassler,B.L.,Wright,M.and Silverman,M.R.(1994).Multiple signaling systems controlling expression of luminescence in Vibrio harveyi:sequence and function of genes encoding a second sensory pathway.Mol.Microbiol.13(2),273-286.
Bassler,B.L,Wright,M.,Showalter,R.E.,et al.(1993).Intercellular signaling in Vibrio harveyi-sequence and function of genes regulating expression of luminescence.Mol Microbiol,9,773-786.
Bellemann,P.and Geider,K.(1992).Localization of transposon insertions in pathogenicity mutants of Erwinia amylovora and their biochemical characterization.J.Gen.Microbiol.138,931-940.
Bereswill,S.and Geider,K.(1997).Characterization of the rcsB gene from Erwinia amylovora and its influence on exoploysaccharide synthesis and virulence of the fire blight pathogen.J.Bacteriol.179,1354-1361.
Bereswill,S.,Bugert,P.,Bruchm(u|¨)ller,I.,et al.(1995).Identification of the fire blight pathogen,Erwinia amylovora by PCR assays with chromosomal DNA.Applied and environmental microbiology 61(7),2636-2642.
Bereswill,S.,Pahl,A.,Bellemann,P.,et al.(1992).Sensitive and species-specific detection of Erwinia amylovora by polymerase chain reaction analysis.Appl Environ Microbiol 58,3522-3526.
Bogdanove,A.J.,Bauer,D.W.and Beer,S.V.(1998).Erwinia amylovora secretes DspE,a pathogenicity factor and functional AvrE homolog,through the Hrp(type Ⅲ secretion) pathway.J.Bacteriol.180,2244-2247.
Bogdanove,A.J.,Beer,S.V.,Bonas,U.,et al.(1996).Unified nomenclature for broadly conserved hrp genes of phytopathogenic bacteria.Mol.Microbiol.20,681-683.
Bogdanove,A.J.,Kim,J.F.,Wei,Z.,et al.(1998).Homology and functional similarity of an hrp-linked pathogenicity locus,dspEF,of Erwinia amylovora and the avirulence locus avrE of Pseudomonas syringae pathovar tomato.Proc.Natl.Acad.Sci.USA 95,1325-1330.
Bogdanove,A.J.,Wei,Z.M.,Zhao,L.,et al.(1996) Erwinia amylovora secretes harpin via a type Ⅲ pathway and contains a homolog of yopN of Yersinia.J.Bacteriol.178,1720-1730.
Bonn,W.G.and van der Zwet,T.(2000).Distribution and economic importance of fire blight In:Fire Blight,the Disease and its Causative Agent,Erwinia amylovora(Vanneste,J.L.,Ed.),pp.37-54.CAB International,Wallingford.
Bugert,P.and Geider,K.(1995).Molecular analysis of the ams operon required for exopolysaccharide synthesis of Erwinia amylovora.Mol.Microbiol.15,917-933.
Burgess,N.A.,Kirke,D.F.,Williams,P.,et al.(2002).LuxS-dependent quorum sensing in Porphyromonas gingivalis modulates protease and haemagglutinin activities but is not essential for virulence.Microbiology,148,763-772.
Burrill,T.J.(1883).New species of Micrococcus.Am.Naturalist.17,319.
Burse,A.,Weingart,H.,and Ullrich,M.S.(2004).NorM,an Erwinia amylovora Multidrug Efflux Pump Involved in In Vitro Competition with Other Epiphytic Bacteria.Appl Environ Microbiol.,70(2),693-703.
Buttner,D.,Nennstiel,D.,Klusener,B.et al.(2002).Functional analysis of HrpF,a putative type Ⅲ translocon protein from Xanthomonas campestris pv.vesicatoria.J.Bacteriol.184,2389-2398.
Camara,M.,Daykin,M.,Chhabra,S.R.(1998).Detection,purification and synthesis of N-2-acylhomoserine lactone quorum sensing signal molecules[J].Methods in Microbiol,1998,27:319-330.
Cao,J.G.,Meighen,E.A.(1989).Purification and structural identification of an autoinducer for the luminescence system of Vibrio harveyi.J.Biol.Chem.264(36),21670-21676.
Cha,C.,Gao,P.,Chen,Y.C.,et al.(1998).Production of acyl-homoserine lactone quorum-sensing signals by gram-negative plant-associated bacteria.Mol Plant Microbe Interact 11(11):1119-1129.
Chen,X.,Schauder,S.,Potier,N.,et al.(2002).Structural identification of a bacterial quorum-sensing signal containing boron.Nature 415(6871):545-549.
Clarke,M.B.,Sperandio,V.(2005).Events at the host-microbial interface of the gastrointestinal tract Ⅲ.Cell-to cell signaling among microbial flora,host and pathogens:There is a whole lot of talking going on.Am J Physiol.288(6,Pt.1):G1105-G1109.
Davies,D.C.,Parsek,M.R.,Pearson,J.P.,et al.(1998).The involvement of cell-to-cell signals in the development of a bacterial biofilm.Science 280(5361),295-298.
Day,W.A.,Maurelli,A.T.(2001).Shigella flexneri LuxS quorum-sensing system modulates virB expression but is not essential for virulence.Infect Immun.69(1),15-23.
Winzer,K.,ttardie,K.R.,Burgess,N.,et al.(2002).LuxS:its role in central metabolism and the in vitro synthesis of 4-hydroxy-5-methyl-3(2H)-furanone.Microbiology.Apr;148(Pt 4),909-922.
de Kievit,T.R.,Iglewski,B.H.(2000).Bacterial quorum sensing in pathogenic relationships.Infect Immun,68(9),4839-4849.
DebRoy,S.,Thilmony,R.,Kwack,Y.B.,et al.(2004).A family of conserved bacterial effectors inhibits salicylic acid-mediated basal immunity and promotes disease necrosis in plants.Proc.Natl.Acad.Sci.USA 101,9927-9932.
Dellagi,A.,Brisset,M.N.,Paulin,J.P.,et al.(1998).Dual role of desferrioxamine in Erwinia amylovora pathogenicity.Mol.Plant Microbe Interact.11,734-742.
Doherty,N.,Holden,M.T.G.,Qazi,S.N.,et al.(2006).Functional Analysis of luxS in Staphylococcus aureus Reveals a Role in Metabolism but Not Quorum Sensing.Journal of bacteriology,188(8) 2885-2897.
Dong,H.,Delaney,T.P.,Bauer,D.W.et al.(1999).Harpin induces disease resistance in Arabidopsis through the systemic acquired resistance pathway mediated by salicylic acid and the NIM1 gene.Plant J.20,207-215.
Dong,H.P.,Peng,J.,Bao,Z.,et al.(2004).Downstream divergence of the ethylene signaling pathway for harpin-stimulated Arabidopsis growth and insect defense.Plant Physiol.136,3628-3638.
Dong,Y.H.,Xu,J.L.,Li,X.Z.,et al.(2000).AiiA,an enzyme that inactivates the acylhomoserine lactone quorum sensing signal and attenuates the virulence of Erwinia carotovora.Proc Natl Acad Sci,USA 97(7):3526-3531.
Dong,Y.H.,Wang,L.H.,Xu,J.L.,et al.(2001).Quenching quorum sensing dependent bacterial infection by an N-acyl homoserine lactonase.Nature 411:813-817.
Eastgate,J.A.(2000).Erwinia arnylovora:the molecular basis of fireblight disease.Mol Plant.Pathol.1,325-329.
Eberhard,A.,Burlingame,A.L.,Eberhard,C.,et al.(1981).Structural identification of autoinducer of Photobacterium fischeri luciferase.Biochemistry 20(9),2444-2449.
Eberl,L.,Winson,M.K.,Sternberg,C.,et al.(1996).Involvement of N-acyl-1-homoserine lactone autoinducers in controlling the multicellular behaviour of Serratia liquefaciens.Mol.Microbiol.20(1),127-136.
Eberl,L.,Winson,M.K.,Sternberg,C.,et al.(1996).Involvement of N-acyl-L-hormoserine lactone autoinducers in controlling the multicellular behaviour of Serratia liquefaciens.Mol Microbiol 20(1),127-136.
El-Maarouf,H.,Barny,M.A.,Rona,J.P.,et al..(2001).Harpin,a hypersensitive response elicitor from Erwinia amylovora,regulates ion channel activities in Arabidopsis thaliana suspension cells.FEBS Lett.497,82-84.
Federle,M.J.,Bassler,B.L.(2003) Interspecies communication in bacteria.J Clin Invest 112(9),1291-1299.
Frommberger,M.,Schmitt-Kopplin,P.,Ping,G.,et al.(2004).A simple and robust set-up for on-column sample preconcentration-nano-liquid chromatography-electrospray ionization mass spectrometry for the analysis of N-acylhomoserine lactones.Anal Bioanal Chem,378,1014-1020.
Fuqua,W.C.,Winans,S.C.and Greenberg,E.P.(1994).Quorum sensing in bacteria:the LuxR-LuxI family of cell density-responsive transcriptional regulators.J Bacteriol.176,269-75.
Gao,M.,Teplitski,M.,Robinson,J.B.,et al.(2003).Production of substances by Medicago truncatula that affect bacterial quorum sensing.Mol Plant Microbe Interact 16:827-834.
Gaudriault,S.,Malandrin,L.,Paulin,J.P.et al.(1997).DspA,an essential pathogenicity factor of Erwinia amylovora showing homology with AvrE of Pseudomonas syringae,is secreted via the Hrp secretion pathway in a DspB-dependent way.Mol.Microbiol.26,1057-1069.
Gaudriault,S.,Paulin,J.P.and Barny,M.A.(2002).The DspB/F protein of Erwinia amylovora is a type Ⅲ secretion chaperone ensuring efficient intrabacterial production of the Hrp-secreted DspA/E pathogenicity factor.Mol.Plant Pathol.3,313-320.
Geier,G.and Geider,K.(1993).Characterization and influence on virulence of the levansucrase gene from the fireblight pathogen Erwinia amylovora.Physiol.Mol.Plant Pathol.42,387-404.
Givskov,M.,de Nys,R.,Manefield,M.,et al.(1996).Eukaryotic interference with homoserine lactone2mediated prokaryotic signalling.J Bacteriol 178,6618-6622.
Goodman,R.N.,Huang,J.S.and Huang,P.Y.(1974).Hostspecific phytotoxic polysaccharide from apple tissue infected by Erwinia amylovora.Science 183,1081-1082.
Greenberg EP,Ulitzur,S and Hastings,J.W.(1979).Induction of luciferase synthesis in Beneckea harveyi by other marine bacteria.Arch Microbiol 120:87-91.
Gross,M.,Geier,G.,Rudolph,K.,et al.(1992).Levan and levansucrase synthesized by the fire blight pathogen Erwinia amylovora.Physiol.Mol.Plant Pathol.40,371-381.
Guiford,P.J.,Taylor,R.K.,Clark,R.G.,et al.(1996).PCR-based techniques for the detection of Erwinia amylovora Acta Horticultural 411:53-56.
Hakansson,S.,Schesser,K.,Persson,C.,et al.(1996).The YopB protein of Yersinia pseudotuberculosis is essential for the translocation of Yop effector proteins across the target cell plasma membrane and displays a contact-dependent membrane disrupting activity.EMBO J.15,5812-5823.
Hale,C.N.and Clark,R.G.(1990).Detection of Erwinia amylovora from apple tissue by DNA hybridization.Acta Horticulturae 273,51-55.
Hastings,J.W.,Nealson,K.H.(1977).Bacterial bioluminescence.Annu RevMicrobiol.31,549-595
Hendricks,C.L.,Ross,J.R.,Piehersky,E.,et al.(2004).An enzyme-coupled colormetric assay for S-adenosylmethionine-dependent rnethyltransferases Anal Biochem,326,100-105.
Morten,H.,Kathrin,R.,Thomas B.R.,et al.(2002).Inhibition of quorum sensing in Fsedomonas aeruginosa biofilm bacteria by a halogenated furanone compound.Microbiology (Reading,United Kingdom),148(1):87-102.
Hentzer,M.,and Givskov,M.(2003).Pharmacological inhibition of quorum sensing for the treatment of chronic bacterial infections.J Clin Invest 112:1300-1307.
Hentzer,M.,Wu,H.,Andersen,J.B.,et al.(2003).Attenuation of Pseudomonas aeruginosa virulence by quorum sensing inhibitors.EMBO J 22:3803-3815.
Hernandez-Guzman,G.and Alvarez-Morales,A.(2001).Isolation and characterization of the gene coding for the amidinotransferase involved in the biosynthesis of phaseolotoxin in Pseudomonas syringae pv.phaseolicola.MoL Plant-Microbe Interact.14,545-554.
Holden,M.T.G.,Ram C.S.,De N.R.,et al.(1999).Quorum-sensing cross talk:isolation and chemical characterization of cyclic dipeptides from Psedomonas aeruginosa and other Gramnegative bacteria.Mol Microbiol.33(6):1254-1266.
Hubner,A.,Revel,A.T.,Nolen,D.M.,et al.(2003).Expression of a luxS gene is not required for Borrelia burgdorferi infection of mice via needle inoculation.Infect Immun 71(5),2892-2896.
Jin,Q.,MeGhee,G.C.,Hart,P.,et al.(2002).Secretion of Hrp and Dsp proteins via the Hrp pilus during type Ⅲ secretion in Erwinia amylovora.Acta Hort.590,441-448.
Johnson,K.B.,and Stockwell,V.O.(1998).Management of fire blight:a case study in microbial ecology.Annu.Rev.Phytopathol.36,227-248.
Joint,I.,Tait,K.,Callow,M.E.,et al.(2002).Cell-to-cell communication across the prokaryote-eukaryote boundary.Science.Nov 8;298(5596):1207.
Kachadourian,R.,Dellagi,A.,Laurent,J.,et al.(1996).Desferrioxamine-dependent iron transport in Erwinia amylovora CFBP 1430:cloning of the gene encoding the ferrioxamine receptor FoxR.Biometals 9,143-150.
Karen,T.E.,and Park,S.F.(2002).Quorum sensing in Campylobacter jejuni:detection of a luxS encoded signaling molecule.Microbiology.148,1475-1481
Kaufmann,G.F.,Sartorio,R.,Lee Ss,H.,et al.(2005).Revisiting quorum sensing:Discovery of additional chemical and diological functions for 3-oxo-N-acyl-homoserine lactones.Proc Natl Acad Sci USA.102(2):309-314.
Kelm,O.,Kiecker,C.,Geider,K.et al.(1997).Interaction of the regulator proteins RcsA and RcsB with the promoter of the operon for amylovoran biosynthesis in Erwinia amylovora.Mol.Gen.Genet.256,72-83.
Kim,J.F.(1997).Molecular characterization of a novel harpin and two hrp secretery operons of Erwinia amylovora,and a hrp operon of E.chrysanthemi.Ph.D.Cornell University,Ithaca,NY.
Kim,J.F.and Beer,S.V.(1998).HrpW of Erwinia amylovora,a new harpin that contains a domain homologous to pectate tyases of a distinct class.J.Bacteriol.180,5203-5210.
Kim,J.F.,Wei,Z.-M.and Beer,S.V.(1997).The hrpA and hrpC operons of Erwinia amylovora encode components of a type Ⅲ pathway that secretes harpin.J.Bacteriol.179,1690-1697.
Kim,W.-S.,Bonasera,J.M.,Meng,X.,et al.(2006).Interaction of DspE/A,a pathogenicity/avirulence protein of Erwinia amylovora,with pre-ferredoxin from apple and its relationship to photosynthetic efficiency.Acta Hort.(in press).
Klars W,Kim R H.(2002).LuxS:its role in central metabolism and the in vitro synthesis of 4-hydroxy-5-methyl-3(2H)-furanone.Microbiology.148:909.
Kleerebezem,M.,Quadri,L.E.,Kuipers,O.P.,et al.(1997).Quorum sensing by peptide pheromones and two-component signal-transduction systems in Gram-positive bacteria.Mol.Microbiol 24(5),895-904.
Lewis,H.A.,Furlong,E.B.,Laubert,B.,et al.(2001).A structural genomics approach to the study of quorum sensing:crystal structures of three LuxS orthologs.Structure.9(6),527-37.
Maes,M.,Garbeva,P.,Crepel,C.(1996).Identification and sensitive endophytic detection of the fire blight pathogen Erwinia amylovora with 23S ribosomal DNA sequences and the polymerase chain reaction.Plant Pathology 45,1139-1149.
Marcus,B.J.,and Blaser,M.J.(2003).Detection of a luxS-Signaling Molecule in Bacillus anthracis.Infection and Immunity.71(7),3914-3919.
Mathesius,U.,Mulders,S.,Gao,M.,et al.(2003).Extensive and specific responses of a eukaryote to bacterial quorum sensing signals.Proc Natl A cad Sci USA 100(3),1444-1449.
Mayville,P.,Ji,G.,Beavis,R.,et al.(1999).Structure-activity analysis of synthetic autoinducing thiolactone peptides from Staphylococcus aureus responsible for virulence.Proc Natl Acad Sci USA.96:1218-1223.
MeClean,K.H.,Winson,M.K.(1997).Quorum sensing and Chromobacterium violaceum:exploitation of violacein production and inhibition for the detection of N-acyl-homoserine lactones.Microbiology 143,3703-3711.
McManus,P.S.and Jones,A.L.(1995).Detection of Erwinia amylovora by nested PCR and PCR-dot-blot and reverse-blot hybridizations.Phytopat.hology 85,618-623.
Meng,X.,Bonasera,J.M.,Kim,J.F.,et al.(2006).Apple proteins that interact with DspE,a pathogenicity effector of Erwinia amylovora,the fire blight pathogen.Mol.Plant-Microbe Interact.(in press).
Merighi,M.,Sandrini,A.(2000).Chemiluminescent and Colorimetric Detection of Erwinia amylovora by Immunoenzymatic Determination of PCR Amplicons from Plasmid pEA29 Plant Disease 84(1),49-54.
Miller,M.B.,Skorupski,K.,et al.(2002).Parallel quorum sensing systems converge to regulate virulence in Vibrio cholerae.Cell.110(3),303-314.
Miller,S.T.,Xavier,K.B.,Campagna,S.R.,et al.(2004) Salmonella typhimurium Recognizes a Chemically Distinct Form of the Bacterial Quorum-Sensing Signal AI-2.Mol.Cell 15,677-687.
Miller,M.B.,Bassler,B.L.(2001) Quorum sensing in bacteria.Annu Rev Microbiol.55,165-199.
Mohammadi,M.,Geider,K.(2007).Autoinducer-2 of the fire blight pathogen Erwinia amylovora and other plant associated bacteria,FEMS Microbiol Lett 266:34-41.
Mok,K.C.,Wingreen,N.S.,Bassler,B.L.(2003).Vibrio harveyi quorum sensing:a coincidence detector for two autoinducers controls gene expression The EMBO Journal 2(4),870-881.
Molina,L.,Constantineseu,F.,Michel,L.,et al.(2003).Degradation of pathogen quorum-sensing molecules by soil bacteria:a preventive and curative biological control mechanism.FEMS Microbiol Ecol 45:71-81.
Molina,L.,Rezzonico,F.,Defago,G.et al.(2005).Autoinduction in Erwinia amylovora:Evidence of an Acyl-Homoserine Lactone Signal in the Fire Blight Pathogen.Journal of Bacteriology.187,3206-3213.
Oh,C.-S.(2005).Characterization of HrpN-interacting proteins from plants,the Hrp pathogenicity island of Erwinia amylovora,and its proteins that affect the hypersensitive response.Ph.D.Cornell University,Ithaca,NY.
Oh,C.-S.,Kim,J.F.and Beer,S.V.(2005) The Hrp pathogenicity island of Erwinia amylovora and the identification of three novel genes required for systemic infection.Mol.Plant Pathol.6,125-138.
Oh,C.-S.and Beer,S.V.(2005).Molecular genetics of Erwinia amylovora involved in the development of fire blight FEMS Microbiology Letters 253,185-192.
Peng,J.-L.,Dong,H.-S.,Dong,H.-P.,et al.(2003).Harpin-elicited hypersensitive cell death and pathogen resistance requires the NDR1 and EDS1 genes.Physiol.Mol.Plant Pathol.62,317-326.
Perino,C.,Gaudriault,S.,Vian,B.,et al.(1999).Visualization of harpin secretion in planta during infection of apple seedlings by Erwinia amylovora.Cell Microbiol.1,131-141.
Petnieki-Oewieja,T.,van Dijk,K.and Alfano,J.R.(2005).The hrpK operon of Pseudomonas syringae pv.tomato DC3000 encodes two proteins secreted by the type Ⅲ(Hrp) protein secretion system:HopB1 and HrpK,a putative type Ⅲ translocator.J.Bacteriol.187,649-663.
Ramos,A.R.(2004).Hrp proteins and harpins:defining their roles in the type Ⅲ protein secretion system in Pseudomonas syringae.Ph.D.Cornell University,Ithaca,NY.
Raseh,M.,Rasmussen,T.B.(2007).Andersen J B.Well-known quorum sensing inhibitors do not affect bacterial quorum sensing-regulated bean spourt spoilage.J Appl microbial.102(3):826-837.
Ratledge,C.and Dover,L.G.(2000).Iron metabolism in pathogenic bacteria.Annu.Rev.Microbiol.54,881-941.
Ren,D.,Sins,J.J.(2001).Wood T K.Inhibition of biofilm formation and swarming of Escherichia coli by(5Z)-4-bromo-5-(bromomethylene)-3-buty-2(5H)-furanone.Environ Microbiol.3:731
Ren,D.,Sins J.J.(2001).Wood T K.Inhibition of biofilm formation and swarming of Bacillus subtilis by(5Z)-4-bromo-5-(bromomethylene)-3-buty-2(5H)-furanone.Lett Appl Microbiol,34:293.
Ren,D.,Sims J.J.,and Wood,T.K.(2002).Inhibition of biofilm formation and swarming of Bacillus subtilis by(5Z)-4-bromo-5-(bromomethylene)-3- butyl-2(5H)-furanone.Lett.Appl.Microbiol.34,293-299.
Ren,D.,Sims,J.J.and Wood,T.K.(2001).Inhibition of biofilm formation and swarming of Escherichia coli by(5Z)-4-bromo-5-(bromomethylene)-3-butyl-2(5H)-furanone.Environ.Microbiol.3,731-736.
Rezzonico F.& Duffy B.(2007).The role of luxS in the fire blight pathogen Erwinia amyIovora is limited to metabolism and does not involve quorum sensing.Molecular Plant-Microbe Interactions 20:1284-1297.
Samebrook,J.,Russell,D.W.(2001).Molecular cloning:a laboratory mannual,3~(rd) ed.Cold Spring Harbor Laboratory.Press,NY.
Sehauder,S.,Bassler,B.L.(2001).The languages of bacteria.Genes Dev 15(12),1468-1480.
Schmidt,H.and Hensel,M.(2004).Pathogenicity islands in bacterial pathogenesis.Clin.Microbiol.Rev.17,14-56.
Sehnabel,E.L.,and Jones,A.L.(1998).Instability of a pEA29 marker in Erwinia amylovora previously used for strain classification.Plant Dis.83,1334-1336.
Seemuller,E.A.and Beer,S.V.(1976).Absence of cell wall polysaccharide degradation of Erwinia amylovora.Phytopathology 66,433-436.
Shaw,P.D.,Ping,G.,Daly,S.L.,et al.(1997) Detecting and characterizing N—acyl-homoserine lactone signal molecules by thin-layer chromatography.Proc.Natl Acad Sci USA 94(12),6036-6041.
Shiner E K,Rumbaugh K P,S C.(2005).Interkingdom signaling:deciphering the language of acyl homoserine lactones.FEMS Microbiol Rev.29(5):935-947
Sjulin,T.M.and Beer,S.V.(1978).Mechanism of wilt induction by amylovoran in Cotoneaster shoots and its relation to wilting of shoots infected by Erwinia amylovora.Phytopathology 68,89-94.
Smith K M,Bu Y,Suga,H.(2003).Induction and inhibition of Pseudomonas aeruginosa quorum sensing by synthetic autoinducer analogs.Chem Biol.10(1):81-89
Sperandio,V.,Torres,A.G.,Jarvis,B.,et al.(2003).Bacteria-host communication:The language of hormones PNAS 100(15),8951-8956
Staskawicz,B.J.,Mudgett,M.B.,Dangl,J.L.et al.(2001).Common and contrasting themes of plant and animal diseases.Science 292,2285-2289.
Steinberger,E.M.and Beer,S.V.(1988).Creation and complementation of pathogenicity mutants of Erwinia amylovora.Mol.Plant-Microbe Interact.1,135-144.
Steinbrenner,B.,Belleman,P.,Zeller,W.,et al.(1990) DNA-hybridization,a specific method for the diagnosis of fireblight.Acta Horticulturae 273,91-93.
Sun,J.,Daniel,R.,Wagner-D(o|¨)bler,I.,et al.(2004).Is autoinducer-2 a universal signal for interspecies communication:a comparative genomic and phylogenetic analysis of the synthesis and signal transduction pathways.BMC Evolutionary Biology,4:Art.No.36.
Surette,M.G.& Bassler,B.L.(1998).Quorum-sensing in Escherichia coli and Salmonella typhimurium.Proc Natl Acad Sci USA 95:7046-7050.
Surette,M.G.& Bassler,B.L.(1999).Regulation of autoinducer production in Salmonella typhimurium.Mol Microbiol 31:585-595.
Swift,S.,Winson,M.K.,Chan,P.F.,et al.(1993).A novel strategy for the isolation of luxl homologues:evidence for the widespread distribution of a LuxR:LuxI superfamily in enteric bacteria.Mol Microbiol.10(3),511-20.
Swift,S.,Downie,J.A.,Whitehead,N.A.,et al.(2001).Quorum sensing as a population-density-dependent determinant of bacterial physiology.Adv Microb Phylosiol.45,199-270
Taga,M.E.and Bassler,B.L.(2003) Chemical communication among bacteria.Proc Natl Acad Sci USA.100(Suppl 2),14549-14554.
Van der Zwet,T.and Beer,S.V.(1991).Fire Blight:Its nature,prevention,and control-A practical guide to integrated disease management.US Dep.Agric.,Agric.Inform.Bull.No.631,34PP.
Van der Zwet,T.,Keil,H.L.(1979).Fire Blight:a bacterial disease of rosaceous plants.Agriculture handbook no.510.U.S.Government Printing Office,Washington,D.C..
Vanneste,J.L.(2000).Fire blight:the disease and its causative agent,Erwinia amylovora.CABI Publishing,Wallingford,Oxon,United Kingdom.
Vanneste,J.L.,Eden-Green,S.(2000).Migration of Erwinia amylovora in host plant tissue,p.9-36.In J.L.Vanneste(ed.),Fire Blight:the disease and its causative agent,Erwinia amylovora.CABI Publishing,Oxon,United Kingdom.
Venisse,J.S.,Barny,M.A.,Paulin,J.P.et al.(2003).Involvement of three pathogenicity factors of Erwinia amylovora in the oxidative stress associated with compatible interaction in pear.FEBS Lett.537,198-202.
Venisse,J.S.,Gullner,G.and Brisset,M.N.(2001).Evidence for the involvement of an oxidative stress in the initiation of infection of pear by Erwinia amylovora.Plant Physiol.125,2164-2172.
Venturi,V.,Venuti,C.,Deveseovi,G.,et al.(2004).The plant pathogen Erwinia amylovora produces acyl-homoserine lactone signal molecules in vitro and in planta.FENS Microbiology letters.241,179-183
Wang,L.(2004).Virulence genes of Erwinia amylovora:explorations,Ph.D.Cornell University,Ithaca,NY.
Wei,Z.,Kim,J.F.and Beer,S.V.(2000).Regulation of hrp genes and type Ⅲ protein secretion in Erwinia amylovora by HrpX/HrpY,a novel two-component system,and HrpS.Mol.Plant-Microbe interact.13,1251-1262.
Wei,Z.-M.and Beer,S.V.(1995).hrpL activates Erwinia amylovora hrp gene transcription and is a member of the ECF subfamily of s factors.J.Bacteriol.177,6201-6210.
Wei,Z.M.,Laby,R.J.,Zumoff,C.It.,et al.(1992).Harpin,elicitor of the hypersensitive response produced by the plant pathogen Erwinia amylovora.Science 257,85-88.
Wei,Z.,Kim,J.F.,Beer,S.V.(2000).Regulation of hrp genes and type Ⅲ protein secretion in Erwinia amylovora by HrpX\HrpY,a novel two-component system and HrpS.Mol.Plant Microbe Interact.13(11),1251-1262.
Winson,M.K.,Swift,S.,Fish,L.,et al.(1998).Construction and analysis of luxCDABE-based plasmid sensors for investigating N-acyl homoserine lactone-mediated quorum sensing.FEMS Microbiol lett.163(2),185-92.
Winzer,K.,Williams,P.(2001).Quorum sensing and the regulation of virulence gene expression in pathogenic bacteria.Int J Med Microbiol.291(2),131-43.
Xavier,K.B.,Bassler,B.L.(2003).LuxS quorum sensing:more than just a numbers game.CurrOp inMicrobiol 6,191-197.
Xie,Z.and Chen,Z.(2000).Harpin-induced hypersensitive cell death is associated with altered mitochondrial functions in tobacco cells.Mol.Plant-Microbe Interact.13,183-190.
Zaccardelfi,M.,and Bazzi,C.(1997).Reliability of PCR with primers from plasmid pEA29 for the detection of Erwinia amylovora.Phytopathol.Mediterr.36,116-120.
Zhang,Y.and Geider,K.(1999).Molecular analysis of the rlsA gene regulating levan production by the fireblight pathogen Erwinia amylovora.Physiol.Mol.Plant Pathol.54,187-201.
Zhang,Y.,Bak,D.D.,Heid,H.,et al.(1999).Molecular characterization of a protease secreted by Erwinia amylovora.J.Mol.Biol.289,1239-1251.
Zhang,L.H.,Murphy,P.J.,Kerr,A.,et al.(1993).Agrobacterium conjugation and gene regulation by N-acyl-1-homoserine lactones.Nature,362(6419),446-447.
李华林,闻玉梅(2004) AI-2,一种新的细菌自体诱导分子,生命科学,16(3):138-143
钱国良,胡白石,卢玲,刘凤权,许志刚(2006)梨火疫病菌的实时荧光PCR检测,植物病理学报,36(2):123-128
陶金莉,迟莉丽,沈亚领,魏东芝(2004)细菌的群体行为调控机制-Quorum sensing,微生物学通报 31(4),106-110
吴红,宋志军,Niels Hoiby(2003)细菌与细菌之间的信息交流——革兰氏阴性细菌的Quorum-sensing系统,自然科学进展,13(7),679-687
高岩,梨火疫病菌群体感应系统的初步研究,硕士学位论文,2006