不动杆菌属中aidE基因编码高丝氨酸内酯酶
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摘要
群体感应(Quorum sensing,QS)是细菌在进化过程中形成的依赖于群体密度的细菌间交流方式。许多革兰氏阴性细菌以N-酰基高丝氨酸内酯(AHL)为信号分子,感应自身群体密度并调控致病基因表达。因此,淬灭AHLs信号分子可防治此类细菌引起的植物病害。本实验室前期已筛选得到了一株具有AHLs信号降解能力的不动杆菌菌株Acinetobacter sp.77,本研究通过基因组文库筛选,自菌株77中克隆得到具有AHLs降解活性的基因aidE。该基因编码268个氨基酸。序列一致性比较发现aidE的氨基酸序列与吉伦伯不动杆菌Acinetobacter gyllenbergii CIP110306中β-内酰胺酶一致性高达95%,但与已知的AHLs降解酶序列一致性较低,最高为缓黄分支杆菌Mycobacterium lentiflavum中AHL内酯酶Att M/Aii B家族蛋白(CQD23908.1),一致性仅为33%。通过高压液相色谱(HPLC)分析Aid E蛋白处理N-己酰基高丝氨酸内酯(C6-HSL)的反应产物,证明aidE为AHL内酯酶。序列比对研究发现,aidE基因在不动杆菌属中并不保守,其在菌株77基因组中的上下游的基因排列存在菌株水平的特异性,且aidE基因下游存在疑似IS插入序列,上述证据表明aidE基因有可能是通过水平转移进入Acinetobacter sp.77基因组中,或其在基因组中的位置发生过重排。表达aidE的软腐果胶杆菌Z3-3中完全检测不到AHLs信号产生,且致病力明显降低。综上所述,aidE为新发现的AHL内酯酶。在防治依赖QS系统表达致病性的细菌病害中具有应用潜力。
Quorum sensing(QS) is a cell-cell communication mechanism that allows bacterial populations to coordinate gene expression in response to cell density. N-acylhomoserine lactones(AHL) are used as quorum-sensing signal molecules by many Gram negative bacteria. Acinetobacter sp. 77, an AHL-degrading bacterium, was isolated in our previous work. The gene aidE for AHL inactivation was cloned in this study by screening a genomic DNA library. The deduced protein AidE is 268 amino acids in length and shares a high identity(95%) with the beta-lactamase family protein in Acinetobacter gyllenbergii CIP110306, but low identities with known AHL-degrading enzymes. HPLC analysis of the Aid E-degraded C6-HSL products revealed that Aid E functioned as an AHL lactonase. Sequences alignment suggested that the aidE gene is not conserved in Acinetobacter species, flanking sequences of aidE and their arrangement are specific in Acinetobacter sp. 77 genome, and some IS insertion sequences were found downstream of the aidE gene. These evidences indicated that the aidE gene might be foreign DNA taken up via horizontal gene transferring or had changed its relative location due to the genome rear-arrangement. Expression of the aidE gene in Pectobacterium carotovorum subsp. carotovorum Z3-3 significantly reduced its AHL production as well as the pathogenicity on host plants, indicating that Aid E was able to effectively quench quorum sensing-dependent functions in bacteria. In conclusion, aidE is a newfound AHL-lactonase with a potential for suppression of bacterial infections.
Quorum sensing(QS) is a cell-cell communication mechanism that allows bacterial populations to coordinate gene expression in response to cell density. N-acylhomoserine lactones(AHL) are used as quorum-sensing signal molecules by many Gram negative bacteria. Acinetobacter sp. 77, an AHL-degrading bacterium, was isolated in our previous work. The gene aidE for AHL inactivation was cloned in this study by screening a genomic DNA library. The deduced protein AidE is 268 amino acids in length and shares a high identity(95%) with the beta-lactamase family protein in Acinetobacter gyllenbergii CIP110306, but low identities with known AHL-degrading enzymes. HPLC analysis of the Aid E-degraded C6-HSL products revealed that Aid E functioned as an AHL lactonase. Sequences alignment suggested that the aidE gene is not conserved in Acinetobacter species, flanking sequences of aidE and their arrangement are specific in Acinetobacter sp. 77 genome, and some IS insertion sequences were found downstream of the aidE gene. These evidences indicated that the aidE gene might be foreign DNA taken up via horizontal gene transferring or had changed its relative location due to the genome rear-arrangement. Expression of the aidE gene in Pectobacterium carotovorum subsp. carotovorum Z3-3 significantly reduced its AHL production as well as the pathogenicity on host plants, indicating that Aid E was able to effectively quench quorum sensing-dependent functions in bacteria. In conclusion, aidE is a newfound AHL-lactonase with a potential for suppression of bacterial infections.
引文
[1]Fuqua WC,Winans SC.A Lux R-Lux I type regulatory system activates Agrobacterium Ti plasmid conjugal transfer in the presence of a plant tumor metabolite.J Bacteriol,1994,176(10):2796–2806.
[2]Bainton NJ,Stead P,Chhabra SR,et al.N-(3-oxohexanoyl)-L-homoserine lactone regulates carbapenem antibiotic production in Erwinia carotovora.Biochem J,1992,288(3):997–1004.
[3]Blankschien MD,Potrykus K,Grace E,et al.Tra R,a homolog of a RNAP secondary channel interactor,modulates transcription.PLo S Genet,2009,5(1):e1000345.
[4]Ren ZG,Lin M,Jiang WJ,et al.Influence of quorum-sensing system on the virulence of Acidovorax citrulli MH21 on melon seedlings.Acta Phytopathol Sin,2012,42(6):608–619(in Chinese).任争光,林敏,姜文君,等.群体感应系统对甜瓜果斑病菌MH21致病力的影响.植物病理学报,2012,42(6):608–619.
[5]Dumenyo CK,Mukherjee A,Chun W,et al.Genetic and physiological evidence for the production of N-acyl homoserine lactones by Pseudomonas syringae pv.syringae and other fluorescent plant pathogenic Pseudomonas species.Eur J Plant Pathol,1998,104(6):569–582.
[6]Waters CM,Bassler BL.Quorum sensing:cell-to-cell communication in bacteria.Annu Rev Cell Dev Biol,2005,21:319–346.
[7]Dong YH,Zhang LH.Quorum sensing and quorum-quenching enzymes.J Microbiol,2005,43:101–109.
[8]Krysciak D,Schmeisser C,Preu?S,et al.Involvement of multiple loci in quorum quenching of autoinducer I molecules in the nitrogen-fixing symbiont Rhizobium(Sinorhizobium)sp.strain NGR234.Appl Environ Microbiol,2011,77(15):5089–5099.
[9]Dong YH,Xu JL,Li XZ,et al.Aii A,an enzyme that inactivates the acylhomoserine lactone quorum-sensing signal and attenuates the virulence of Erwinia carotovora.Proc Natl Acad Sci USA,2000,97(7):3526–3531.
[10]Mei GY,Yan XX,Turak A,et al.Aid H,an alpha/beta-hydrolase fold family member from an Ochrobactrum sp.strain,is a novel N-acylhomoserine lactonase.Appl Environ Microbiol,2010,76(15):4933–4942.
[11]Uroz S,Oger PM,Chapelle E,et al.A Rhodococcus qsd A-encoded enzyme defines a novel class of large-spectrum quorum-quenching lactonases.Appl Environ Microbiol,2008,74(5):1357–1366.
[12]Riaz K,Elmerich C,Raffoux A,et al.Metagenomics revealed a quorum quenching lactonase Qlc A from yet unculturable soil bacteria.Commun Agric Appl Biol Sci,2008,73(2):3–6.
[13]Schipper C,Hornung C,Bijtenhoorn P,et al.Metagenome-derived clones encoding two novel lactonase family proteins involved in biofilm inhibition in Pseudomonas aeruginosa.Appl Environ Microbiol,2009,75(1):224–233.
[14]Dong YH,Wang LH,Xu JL,et al.Quenching quorum-sensing-dependent bacterial infection by an N-acyl homoserine lactonase.Nature,2001,411(6839):813–817.
[15]Ban HF,Chai XL,Lin YJ,et al.Transgenic Amorphophallus konjac expressing synthesized acyl-homoserine lactonase(aii A)gene exhibit enhanced resistance to soft rot disease.Plant Cell Rep,2009,28(12):1847–1855.
[16]Vanjildorj E,Song SY,Yang ZH,et al.Enhancement of tolerance to soft rot disease in the transgenic Chinese cabbage(Brassica rapa L.ssp.pekinensis)inbred line,Kenshin.Plant Cell Rep,2009,28(10):1581–1591.
[17]Chai XL,Zhou Y,Lin YJ,et al.Regeneration of transgenic Amorphophallus konjac expressing synthesized acyl-homoserine lactonase(aii A)gene from Bacillus thuringiensis.Mol Plant Breed,2007,5(5):613–618(in Chinese).柴鑫莉,周盈,林拥军,等.苏云金芽胞杆菌抗软腐病aii A基因转花魔芋研究.分子植物育种,2007,5(5):613–618.
[18]Cha C,Gao P,Chen YC,et al.Production of acyl-homoserine lactone quorum-sensing signals by gram-negative plant-associated bacteria.Mol Plant Microbe Interact,1998,11(11):1119–1129.
[19]Hanahan D.Studies on transformation of Escherichia coli with plasmids.J Mol Biol,1983,166(4):557–580.
[20]Kovach ME,Elzer PH,Hill DS,et al.Four new derivatives of the broad-host-range cloning vector p BBR1MCS,carrying different antibiotic-resistance cassettes.Gene,1995,166(1):175–176.
[21]Kessler B,de Lorenzo V,Timmis KN.A general system to integrate lac Z fusions into the chromosomes of gram-negative eubacteria:regulation of the Pm promoter of the TOL plasmid studied with all controlling elements in monocopy.Mol Gen Genet,1992,233(1/2):293–301.
[22]Keen NT,Tamaki S,Kobayashi D,et al.Improved broad-host-range plasmids for DNA cloning in gram-negative bacteria.Gene,1988,70(1):191–197.
[23]Haudecoeur E,Tannières M,Cirou A,et al.Different regulation and roles of lactonases Aii B and Att M in Agrobacterium tumefaciens C58.Mol Plant Microbe Interact,2009,22(5):529–537.
[24]Khan SR,Farrand SK.The Blc C(Att M)lactonase of Agrobacterium tumefaciens does not quench the quorum-sensing system that regulates Ti plasmid conjugative transfer.J Bacteriol,2009,191(4):1320–1329.
[25]Zhang HB,Wang LH,Zhang LH.Genetic control of quorum-sensing signal turnover in Agrobacterium tumefaciens.Proc Natl Acad Sci USA,2002,99(7):4638–4643.
[26]Carlier A,Uroz S,Smadja B,et al.The Ti plasmid of Agrobacterium tumefaciens harbors an att M-paralogous gene,aii B,also encoding N-acyl homoserine lactonase activity.Appl Environ Microbiol,2003,69(8):4989–4993.
[27]Park SY,Lee SJ,Oh TK,et al.Ahl D,an N-acylhomoserine lactonase in Arthrobacter sp.,and predicted homologues in other bacteria.Microbiology,2003,149(6):1541–1550.
[28]Wang WZ,Morohoshi T,Someya N,et al.Aid C,a novel N-acylhomoserine lactonase from the potato root-associated Cytophaga-Flavobacteria-Bacteroides(CFB)group bacterium Chryseobacterium sp.strain St RB126.Appl Environ Microbiol,2012,78(22):7985–7992.
[29]Wang WZ,Morohoshi T,Ikenoya M,et al.Aii M,a novel class of N-acylhomoserine lactonase from the leaf-associated bacterium Microbacterium testaceum.Appl Environ Microbiol,2010,76(8):2524–2530.
[30]Huang W,Lin YJ,Yi SY,et al.Qsd H,a novel AHL lactonase in the RND-Type inner membrane of marine Pseudoalteromonas byunsanensis strain1A01261.PLo S ONE,2012,7(10):e46587.
[31]Bijtenhoorn P,Schipper C,Hornung C,et al.Bpi B05,a novel metagenome-derived hydrolase acting on N-acylhomoserine lactones.J Biotechnol,2011,155(1):86–94.
[32]Bar-Rogovsky H,Hugenmatter A,Tawfik DS.The evolutionary origins of detoxifying enzymes:the mammalian serum paraoxonases(PONs)relate to bacterial homoserine lactonases.J Biol Chem,2013,288(33):23914–23927.
[33]Bergonzi C,Schwab M,Elias M.The quorum-quenching lactonase from Geobacillus caldoxylosilyticus:purification,characterization,crystallization and crystallographic analysis.Acta Crystallogr F Struct Biol Commun,2016,72(9):681–686.
[34]Garge SS,Nerurkar AS.Attenuation of quorum sensing regulated virulence of Pectobacterium carotovorum subsp.carotovorum through an AHL lactonase produced by Lysinibacillus sp.Gs50.PLo S ONE,2016,11(12):e0167344.
[35]Hiblot J,Gotthard G,Chabriere E,et al.Structural and enzymatic characterization of the lactonase Sis Lac from Sulfolobus islandicus.PLo S ONE,2012,7(10):e47028.
[36]Hiblot J,Gotthard G,Elias M,et al.Differential active site loop conformations mediate promiscuous activities in the lactonase Sso Pox.PLo S ONE,2013,8(9):e75272.
[37]Valera MJ,Mas A,Streit WR,et al.Gqq A,a novel protein in Komagataeibacter europaeus involved in bacterial quorum quenching and cellulose formation.Microb Cell Fact,2016,15(1):88.
[38]Morohoshi T,Sato N,Iizumi T,et al.Identification and characterization of a novel N-acyl-homoserine lactonase gene in Sphingomonas ursincola isolated from industrial cooling water systems.J Biosci Bioeng,2017,123(5):569–575.
[39]Morohoshi T,Tominaga Y,Someya N,et al.Complete genome sequence and characterization of the N-acylhomoserine lactone-degrading gene of the potato leaf-associated Solibacillus silvestris.J Biosci Bioeng,2012,113(1):20–25.
[40]Zhang YH,Liu JW,Tang KH,et al.Genome analysis of Flaviramulus ichthyoenteri Th78T in the family Flavobacteriaceae:insights into its quorum quenching property and potential roles in fish intestine.BMC Genomics,2015,16(1):38.
[41]Yates EA,Philipp B,Buckley C,et al.N-acylhomoserine lactones undergo lactonolysis in a p H-,temperature-,and acyl chain length-dependent manner during growth of Yersinia pseudotuberculosis and Pseudomonas aeruginosa.Infect Immun,2002,70(10):5635–5646.
[42]Abbott DW,Boraston AB.Structural biology of pectin degradation by Enterobacteriaceae.Microbiol Mol Biol Rev,2008,72(2):301–316.
[43]Toth IK,Bell KS,Holeva MC,et al.Soft rot erwiniae:from genes to genomes.Mol Plant Pathol,2003,4(1):17–30.
[44]Jafra S,Jalink H,van der Schoor R,et al.Pectobacterium carotovorum subsp.carotovorum strains show diversity in production of and response to N-acyl homoserine lactones.J Phytopathol,2006,154(11/12):729–739.
[45]Holmquist M.Alpha/Beta-hydrolase fold enzymes:structures,functions and mechanisms.Curr Protein Pept Sci,2000,1(2):209–235.
[46]Wang LH,Weng LX,Dong YH,et al.Specificity and enzyme kinetics of the quorum-quenching N-acyl homoserine lactone lactonase(AHL-lactonase).J Biol Chem,2004,279(14):13645–13651.
[47]Ochiai S,Yasumoto S,Morohoshi T,et al.Ami E,a novel N-acylhomoserine Lactone acylase belonging to the amidase family,from the activated-sludge isolate Acinetobacter sp.strain Ooi24.Appl Environ Microbiol,2014,80(22):6919–6925.
[2]Bainton NJ,Stead P,Chhabra SR,et al.N-(3-oxohexanoyl)-L-homoserine lactone regulates carbapenem antibiotic production in Erwinia carotovora.Biochem J,1992,288(3):997–1004.
[3]Blankschien MD,Potrykus K,Grace E,et al.Tra R,a homolog of a RNAP secondary channel interactor,modulates transcription.PLo S Genet,2009,5(1):e1000345.
[4]Ren ZG,Lin M,Jiang WJ,et al.Influence of quorum-sensing system on the virulence of Acidovorax citrulli MH21 on melon seedlings.Acta Phytopathol Sin,2012,42(6):608–619(in Chinese).任争光,林敏,姜文君,等.群体感应系统对甜瓜果斑病菌MH21致病力的影响.植物病理学报,2012,42(6):608–619.
[5]Dumenyo CK,Mukherjee A,Chun W,et al.Genetic and physiological evidence for the production of N-acyl homoserine lactones by Pseudomonas syringae pv.syringae and other fluorescent plant pathogenic Pseudomonas species.Eur J Plant Pathol,1998,104(6):569–582.
[6]Waters CM,Bassler BL.Quorum sensing:cell-to-cell communication in bacteria.Annu Rev Cell Dev Biol,2005,21:319–346.
[7]Dong YH,Zhang LH.Quorum sensing and quorum-quenching enzymes.J Microbiol,2005,43:101–109.
[8]Krysciak D,Schmeisser C,Preu?S,et al.Involvement of multiple loci in quorum quenching of autoinducer I molecules in the nitrogen-fixing symbiont Rhizobium(Sinorhizobium)sp.strain NGR234.Appl Environ Microbiol,2011,77(15):5089–5099.
[9]Dong YH,Xu JL,Li XZ,et al.Aii A,an enzyme that inactivates the acylhomoserine lactone quorum-sensing signal and attenuates the virulence of Erwinia carotovora.Proc Natl Acad Sci USA,2000,97(7):3526–3531.
[10]Mei GY,Yan XX,Turak A,et al.Aid H,an alpha/beta-hydrolase fold family member from an Ochrobactrum sp.strain,is a novel N-acylhomoserine lactonase.Appl Environ Microbiol,2010,76(15):4933–4942.
[11]Uroz S,Oger PM,Chapelle E,et al.A Rhodococcus qsd A-encoded enzyme defines a novel class of large-spectrum quorum-quenching lactonases.Appl Environ Microbiol,2008,74(5):1357–1366.
[12]Riaz K,Elmerich C,Raffoux A,et al.Metagenomics revealed a quorum quenching lactonase Qlc A from yet unculturable soil bacteria.Commun Agric Appl Biol Sci,2008,73(2):3–6.
[13]Schipper C,Hornung C,Bijtenhoorn P,et al.Metagenome-derived clones encoding two novel lactonase family proteins involved in biofilm inhibition in Pseudomonas aeruginosa.Appl Environ Microbiol,2009,75(1):224–233.
[14]Dong YH,Wang LH,Xu JL,et al.Quenching quorum-sensing-dependent bacterial infection by an N-acyl homoserine lactonase.Nature,2001,411(6839):813–817.
[15]Ban HF,Chai XL,Lin YJ,et al.Transgenic Amorphophallus konjac expressing synthesized acyl-homoserine lactonase(aii A)gene exhibit enhanced resistance to soft rot disease.Plant Cell Rep,2009,28(12):1847–1855.
[16]Vanjildorj E,Song SY,Yang ZH,et al.Enhancement of tolerance to soft rot disease in the transgenic Chinese cabbage(Brassica rapa L.ssp.pekinensis)inbred line,Kenshin.Plant Cell Rep,2009,28(10):1581–1591.
[17]Chai XL,Zhou Y,Lin YJ,et al.Regeneration of transgenic Amorphophallus konjac expressing synthesized acyl-homoserine lactonase(aii A)gene from Bacillus thuringiensis.Mol Plant Breed,2007,5(5):613–618(in Chinese).柴鑫莉,周盈,林拥军,等.苏云金芽胞杆菌抗软腐病aii A基因转花魔芋研究.分子植物育种,2007,5(5):613–618.
[18]Cha C,Gao P,Chen YC,et al.Production of acyl-homoserine lactone quorum-sensing signals by gram-negative plant-associated bacteria.Mol Plant Microbe Interact,1998,11(11):1119–1129.
[19]Hanahan D.Studies on transformation of Escherichia coli with plasmids.J Mol Biol,1983,166(4):557–580.
[20]Kovach ME,Elzer PH,Hill DS,et al.Four new derivatives of the broad-host-range cloning vector p BBR1MCS,carrying different antibiotic-resistance cassettes.Gene,1995,166(1):175–176.
[21]Kessler B,de Lorenzo V,Timmis KN.A general system to integrate lac Z fusions into the chromosomes of gram-negative eubacteria:regulation of the Pm promoter of the TOL plasmid studied with all controlling elements in monocopy.Mol Gen Genet,1992,233(1/2):293–301.
[22]Keen NT,Tamaki S,Kobayashi D,et al.Improved broad-host-range plasmids for DNA cloning in gram-negative bacteria.Gene,1988,70(1):191–197.
[23]Haudecoeur E,Tannières M,Cirou A,et al.Different regulation and roles of lactonases Aii B and Att M in Agrobacterium tumefaciens C58.Mol Plant Microbe Interact,2009,22(5):529–537.
[24]Khan SR,Farrand SK.The Blc C(Att M)lactonase of Agrobacterium tumefaciens does not quench the quorum-sensing system that regulates Ti plasmid conjugative transfer.J Bacteriol,2009,191(4):1320–1329.
[25]Zhang HB,Wang LH,Zhang LH.Genetic control of quorum-sensing signal turnover in Agrobacterium tumefaciens.Proc Natl Acad Sci USA,2002,99(7):4638–4643.
[26]Carlier A,Uroz S,Smadja B,et al.The Ti plasmid of Agrobacterium tumefaciens harbors an att M-paralogous gene,aii B,also encoding N-acyl homoserine lactonase activity.Appl Environ Microbiol,2003,69(8):4989–4993.
[27]Park SY,Lee SJ,Oh TK,et al.Ahl D,an N-acylhomoserine lactonase in Arthrobacter sp.,and predicted homologues in other bacteria.Microbiology,2003,149(6):1541–1550.
[28]Wang WZ,Morohoshi T,Someya N,et al.Aid C,a novel N-acylhomoserine lactonase from the potato root-associated Cytophaga-Flavobacteria-Bacteroides(CFB)group bacterium Chryseobacterium sp.strain St RB126.Appl Environ Microbiol,2012,78(22):7985–7992.
[29]Wang WZ,Morohoshi T,Ikenoya M,et al.Aii M,a novel class of N-acylhomoserine lactonase from the leaf-associated bacterium Microbacterium testaceum.Appl Environ Microbiol,2010,76(8):2524–2530.
[30]Huang W,Lin YJ,Yi SY,et al.Qsd H,a novel AHL lactonase in the RND-Type inner membrane of marine Pseudoalteromonas byunsanensis strain1A01261.PLo S ONE,2012,7(10):e46587.
[31]Bijtenhoorn P,Schipper C,Hornung C,et al.Bpi B05,a novel metagenome-derived hydrolase acting on N-acylhomoserine lactones.J Biotechnol,2011,155(1):86–94.
[32]Bar-Rogovsky H,Hugenmatter A,Tawfik DS.The evolutionary origins of detoxifying enzymes:the mammalian serum paraoxonases(PONs)relate to bacterial homoserine lactonases.J Biol Chem,2013,288(33):23914–23927.
[33]Bergonzi C,Schwab M,Elias M.The quorum-quenching lactonase from Geobacillus caldoxylosilyticus:purification,characterization,crystallization and crystallographic analysis.Acta Crystallogr F Struct Biol Commun,2016,72(9):681–686.
[34]Garge SS,Nerurkar AS.Attenuation of quorum sensing regulated virulence of Pectobacterium carotovorum subsp.carotovorum through an AHL lactonase produced by Lysinibacillus sp.Gs50.PLo S ONE,2016,11(12):e0167344.
[35]Hiblot J,Gotthard G,Chabriere E,et al.Structural and enzymatic characterization of the lactonase Sis Lac from Sulfolobus islandicus.PLo S ONE,2012,7(10):e47028.
[36]Hiblot J,Gotthard G,Elias M,et al.Differential active site loop conformations mediate promiscuous activities in the lactonase Sso Pox.PLo S ONE,2013,8(9):e75272.
[37]Valera MJ,Mas A,Streit WR,et al.Gqq A,a novel protein in Komagataeibacter europaeus involved in bacterial quorum quenching and cellulose formation.Microb Cell Fact,2016,15(1):88.
[38]Morohoshi T,Sato N,Iizumi T,et al.Identification and characterization of a novel N-acyl-homoserine lactonase gene in Sphingomonas ursincola isolated from industrial cooling water systems.J Biosci Bioeng,2017,123(5):569–575.
[39]Morohoshi T,Tominaga Y,Someya N,et al.Complete genome sequence and characterization of the N-acylhomoserine lactone-degrading gene of the potato leaf-associated Solibacillus silvestris.J Biosci Bioeng,2012,113(1):20–25.
[40]Zhang YH,Liu JW,Tang KH,et al.Genome analysis of Flaviramulus ichthyoenteri Th78T in the family Flavobacteriaceae:insights into its quorum quenching property and potential roles in fish intestine.BMC Genomics,2015,16(1):38.
[41]Yates EA,Philipp B,Buckley C,et al.N-acylhomoserine lactones undergo lactonolysis in a p H-,temperature-,and acyl chain length-dependent manner during growth of Yersinia pseudotuberculosis and Pseudomonas aeruginosa.Infect Immun,2002,70(10):5635–5646.
[42]Abbott DW,Boraston AB.Structural biology of pectin degradation by Enterobacteriaceae.Microbiol Mol Biol Rev,2008,72(2):301–316.
[43]Toth IK,Bell KS,Holeva MC,et al.Soft rot erwiniae:from genes to genomes.Mol Plant Pathol,2003,4(1):17–30.
[44]Jafra S,Jalink H,van der Schoor R,et al.Pectobacterium carotovorum subsp.carotovorum strains show diversity in production of and response to N-acyl homoserine lactones.J Phytopathol,2006,154(11/12):729–739.
[45]Holmquist M.Alpha/Beta-hydrolase fold enzymes:structures,functions and mechanisms.Curr Protein Pept Sci,2000,1(2):209–235.
[46]Wang LH,Weng LX,Dong YH,et al.Specificity and enzyme kinetics of the quorum-quenching N-acyl homoserine lactone lactonase(AHL-lactonase).J Biol Chem,2004,279(14):13645–13651.
[47]Ochiai S,Yasumoto S,Morohoshi T,et al.Ami E,a novel N-acylhomoserine Lactone acylase belonging to the amidase family,from the activated-sludge isolate Acinetobacter sp.strain Ooi24.Appl Environ Microbiol,2014,80(22):6919–6925.