细菌脲酶蛋白复合物及其活化机制
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摘要
脲酶能够催化尿素分解生成氨,在农业和医学领域中具有重要的意义。细菌脲酶蛋白包括结构蛋白(UreA、UreB和UreC)和辅助蛋白(UreD/UreH、UreE、UreF和UreG),它们在脲酶活化过程中各自具有独特的作用,结构蛋白形成脲酶活性中心,而辅助蛋白主要负责镍离子的传递。文中综述了细菌脲酶蛋白复合物的结构和功能,以及各蛋白之间如何相互作用完成其活化过程,以期为脲酶活性调控研究及脲酶抑制剂开发等提供理论指导。
Urease decomposes urea to ammonia, and has application potential in agriculture and medical treatment. Urease proteins include structural proteins(UreA, UreB and UreC) and accessory proteins(UreD/UreH, UreE, UreF and UreG), each of them has its own unique role in urease maturation. The structural proteins form the active center of urease, and the accessory proteins are responsible for the delivery of nickel. We review here the structure and function of bacterial urease complexes, and how each protein interacts to complete the activation process. We hope to provide theoretical basis for the regulation of urease activity and the development of urease inhibitors.
Urease decomposes urea to ammonia, and has application potential in agriculture and medical treatment. Urease proteins include structural proteins(UreA, UreB and UreC) and accessory proteins(UreD/UreH, UreE, UreF and UreG), each of them has its own unique role in urease maturation. The structural proteins form the active center of urease, and the accessory proteins are responsible for the delivery of nickel. We review here the structure and function of bacterial urease complexes, and how each protein interacts to complete the activation process. We hope to provide theoretical basis for the regulation of urease activity and the development of urease inhibitors.
引文
[1]Zambelli B,Musiani F,Benini S,et al.Chemistry of Ni2+ in urease:sensing,trafficking,and catalysis.Acc Chem Res,2011,44(7):520-530.
[2]Konieczna I,?arnowiec P,Kwinkowski M,et al.Bacterial urease and its role in long-lasting human diseases.Curr Protein Pept Sci,2012,13(8):789-806.
[3]Zhao SG,Wang JQ,Liu KL,et al.Analysis for the diversity of ureolytic bacterium from dairy rumen based on metagenomics.J China Agric Univ,2010,15(1):55-61(in Chinese).赵圣国,王加启,刘开朗,等.宏基因组学方法分析奶牛瘤胃尿素分解菌的多样性.中国农业大学学报,2010,15(1):55-61.
[4]Jin D,Zhao SG,Zheng N,et al.Differences in ureolytic bacterial composition between the rumen digesta and rumen wall based on ureC Gene Classification.Front Microbiol,2017,8:385.
[5]Jin D,Zhao SG,Wang PP,et al.Insights into abundant rumen ureolytic bacterial community using rumen simulation system.Front Microbiol,2016,7:1006.
[6]Zhao SG,Wang JQ,Bu DP,et al.Biochemistry and molecular biology of bacterial ureases.Microbiology,2008,35(7):1146-1152(in Chinese).赵圣国,王加启,卜登攀,等.细菌尿素酶的生化和分子生物学特点.微生物学通报,2008,35(7):1146-1152.
[7]Toffanin A,Cadahia E,Imperial J,et al.Characterization of the urease gene cluster from Rhizobium leguminosarum cv.viciae.Arch Microbiol,2002,177(4):290-298.
[8]Maeda M,Hidaka M,Nakamura A,et al.Cloning,sequencing,and expression of thermophilic Bacillus sp.strain TB-90 urease gene complex in Escherichia coli.JBacteriol,1994,176(2):432-442.
[9]BosséJT,Gilmour HD,MacInnes JI.Novel genes affecting urease activity in Actinobacillus pleuropneumoniae.J Bacteriol,2001,183(4):1242-1247.
[10]Stoof J,Breijer S,Pot RGJ,et al.Inverse nickel-responsive regulation of two urease enzymes in the gastric pathogen Helicobacter mustelae.Environ Microbiol,2010,10(10):2586-2597.
[11]Kim JK,Mulrooney SB,Hausinger RP.Biosynthesis of active Bacillus subtilis urease in the absence of known urease accessory proteins.J Bacteriol,2005,187(20):7150-7154.
[12]Carter EL,Flugga N,Boer JL,et al.Interplay of metal ions and urease.Metallomics,2009,1(3):207-221.
[13]Benini S,Rypniewski WR,Wilson KS,et al.A new proposal for urease mechanism based on the crystal structures of the native and inhibited enzyme from Bacillus pasteurii:why urea hydrolysis costs two nickels.Structure,1999,7(2):205-216.
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[16]Park IS,Carr MB,Hausinger RP.In vitro activation of urease apoprotein and role of UreD as a chaperone required for nickel metallocenter assembly.Proc Natl Acad Sci USA,1994,91(8):3233-3237.
[17]Moncrief MB,Hausinger RP.Purification and activation properties of UreD-UreF-urease apoprotein complexes.JBacteriol,1996,178(18):5417-5421.
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[22]Zambelli B,Turano P,Musiani F,et al.Zn2+-linked dimerization of UreG from Helicobacter pylori,a chaperone involved in nickel trafficking and urease activation.Proteins,2010,74(1):222-239.
[23]Leipe DD,Wolf YI,Koonin EV,et al.Classification and evolution of P-loop GTPases and related ATPases.J Mol Biol,2002,317(1):41-72.
[24]Bange G,Sinning I.SIMIBI twins in protein targeting and localization.Nat Struct Mol Biol,2013,20(7):776-780.
[25]Gasper R,Meyer S,Gotthardt K,et al.It takes two to tango:regulation of G proteins by dimerization.Nat Rev Mol Cell Biol,2009,10(6):423-429.
[26]Boer JL,Hausinger RP.Klebsiella aerogenes UreF:identification of the UreG binding site and role in enhancing the fidelity of urease activation.Biochemistry,2012,51(11):2298-2308.
[27]Yang XM,Li HY,Lai TP,et al.UreE-UreG complex facilitates nickel transfer and preactivates GTPase of UreG in Helicobacter pylori.J Biol Chem,2015,290(20):12474-12485.
[28]Lee MH,Pankratz HS,Wang SK,et al.Purification and characterization of Klebsiella aerogenes UreE protein:a nickel-binding protein that functions in urease metallocenter assembly.Protein Sci,2010,2(6):1042-1052.
[29]Brayman TG,Hausinger RP.Purification,characterization,and functional analysis of a truncated Klebsiella aerogenes UreE urease accessory protein lacking the histidine-rich carboxyl terminus.J Bacteriol,1996,178(18):5410-5416.
[30]Banaszak K,Martin-Diaconescu V,Bellucci M,et al.Crystallographic and X-ray absorption spectroscopic characterization of Helicobacter pylori UreE bound to Ni2+and Zn2+reveals a role for the disordered C-terminal arm in metal trafficking.Biochem J,2012,441(3):1017-1035.
[31]Shi R,Munger C,Asinas A,et al.Crystal structures of apo and metal-bound forms of the UreE protein from Helicobacter pylori:role of multiple metal binding sites.Biochemistry,2010,49(33):7080-7088.
[32]Song HK,Mulrooney SB,Huber R,et al.Crystal structure of Klebsiella aerogenes UreE,a nickel-binding metallochaperone for urease activation.J Biol Chem,2001,276(52):49359-49364.
[33]Remaut H,Safarov N,Ciurli S,et al.Structural basis for Ni2+transport and assembly of the urease active site by the metallochaperone UreE from Bacillus pasteurii.JBiol Chem,2001,276(52):49365-49370.
[34]Bellucci M,Zambelli B,Musiani F,et al.Helicobacter pylori UreE,a urease accessory protein:specific Ni2+and Zn2+binding properties and interaction with its cognate UreG.Biochem J,2009,422(1):91-100.
[35]Boer JL,Quiroz-Valenzuela S,Anderson KL,et al.Mutagenesis of Klebsiella aerogenes UreG to probe nickel binding and interactions with other urease-related proteins.Biochemistry,2010,49(28):5859-5869.
[36]Chan KH,Lee KM,Wong KB.Interaction between hydrogenase maturation factors HypA and HypB is required for[NiFe]-hydrogenase maturation.PLoS ONE,2012,7(2):e32592.
[37]Mehta N,Olson JW,Maier RJ.Characterization of Helicobacter pylori nickel metabolism accessory proteins needed for maturation of both urease and hydrogenase.J Bacteriol,2003,185(3):726-734.
[38]Yang XM,Li HY,Cheng TF,et al.Nickel translocation between metallochaperones HypA and UreE in Helicobacter pylori.Metallomics,2014,6(9):1731-1736.
[39]Sydor AM,Lebrette H,Ariyakumaran R,et al.Relationship between Ni(II)and Zn(II)coordination and nucleotide binding by the Helicobacter pylori[NiFe]-hydrogenase and urease maturation factor HypB.J Biol Chem,2014,289(7):3828-3841.
[40]Benoit SL,Zbell AL,Maier RJ.Nickel enzyme maturation in Helicobacter hepaticus:roles of accessory proteins in hydrogenase and urease activities.Microbiology,2007,153(11):3748-3756.
[41]Strugatsky D,McNulty R,Munson K,et al.Structure of the proton-gated urea channel from the gastric pathogen Helicobacter pylori.Nature,2013,493(7431):255-258.
[42]Cáceres-Delpiano J,Teneb J,Mansilla R,et al.Variations in periplasmic loop interactions determine the pH-dependent activity of the hexameric urea transporter UreI from Helicobacter pylori:a molecular dynamics study.BMC Struct Biol,2015,15(1):1-12.
[43]Sebbane F,Mandrand-Berthelot MA,Simonet M.Genes encoding specific nickel transport systems flank the chromosomal urease locus of Pathogenic yersiniae.JBacteriol,2002,184(20):5706-5713.
[44]Chen YYM,Burne RA.Identification and characterization of the nickel uptake system for urease biogenesis in Streptococcus salivarius 57.I.Virology,2003,183(23):6773-6779.
[45]Sigel A,Sigel H,Sigel RKO.Nickel and Its Surprising Impact in Nature,Volume 2.Hoboken,NJ:John Wiley&Sons,Ltd,2007:519-544.
[46]Musiani F,Gioia D,Masetti M,et al.Protein tunnels:the case of urease accessory proteins.J Chem Theory Comput,2017,13(5):2322-2331.
[47]Zambelli B,Berardi A,Martin-Diaconescu V,et al.Nickel binding properties of Helicobacter pylori UreF,an accessory protein in the nickel-based activation of urease.J Biol Inorg Chem,2014,19(3):319-334.
[48]Chang ZZ,Kuchar J,Hausinger RP.Chemical cross-linking and mass spectrometric identification of sites of interaction for UreD,UreF,and urease.J Biol Chem,2004,279(15):15305-15313.
[49]Quiroz-Valenzuela S,Sukuru SCK,Hausinger RP,et al.The structure of urease activation complexes examined by flexibility analysis,mutagenesis,and small-angle X-ray scattering.Arch Biochem Biophys,2008,480(1):51-57.
[50]Park IS,Hausinger RP.Requirement of carbon dioxide for in vitro assembly of the urease nickel metallocenter.Science,1995,267(5201):1156-1158.
[51]Carter EL,Boer JL,Farrugia MA,et al.Function of UreB in Klebsiella aerogenes urease.Biochemistry,2011,50(43):9296-9308.
[52]Soriano A,Colpas GJ,Hausinger RP.UreE stimulation of GTP-dependent urease activation in the UreD-UreF-UreG-urease apoprotein complex.Biochemistry,2000,39(40):12435-12440.
[53]Farrugia MA,Macomber L,Hausinger RP.Biosynthesis of the urease metallocenter.J Biol Chem,2013,288(19):13178-13185.
[54]Soriano A,Hausinger RP.GTP-dependent activation of urease apoprotein in complex with the UreD,UreF,and UreG accessory proteins.Proc Natl Acad Sci USA,1999,96(20):11140-11144.
[55]Fong YH,Wong HC,Yuen MH,et al.Structure of UreG/UreF/UreH complex reveals how urease accessory proteins facilitate maturation of Helicobacter pylori urease.PLoS Biol,2013,11(10):e1001678.
[56]Stingl K,Schauer K,Ecobichon C,et al.In vivo interactome of Helicobacter pylori urease revealed by tandem affinity purification.Mol Cell Proteomics,2008,7(12):2429-2441
[57]Farrugia MA,Han LJ,Zhong YY,et al.Analysis of a soluble(UreD:UreF:UreG)2 accessory protein complex and its interactions with Klebsiella aerogenes urease by mass spectrometry.J Am Soc Mass Spectrom,2013,24(9):1328-1337.
[58]Merloni A,Dobrovolska O,Zambelli B,et al.Molecular landscape of the interaction between the urease accessory proteins UreE and UreG.Biochim Biophys Acta,2014,1844(9):1662-1674.
[59]Yuen MH,Fong YH,Nim YS,et al.Structural insights into how GTP-dependent conformational changes in a metallochaperone UreG facilitate urease maturation.Proc Natl Acad Sci USA,2017,114(51):E10890-E10898.
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[2]Konieczna I,?arnowiec P,Kwinkowski M,et al.Bacterial urease and its role in long-lasting human diseases.Curr Protein Pept Sci,2012,13(8):789-806.
[3]Zhao SG,Wang JQ,Liu KL,et al.Analysis for the diversity of ureolytic bacterium from dairy rumen based on metagenomics.J China Agric Univ,2010,15(1):55-61(in Chinese).赵圣国,王加启,刘开朗,等.宏基因组学方法分析奶牛瘤胃尿素分解菌的多样性.中国农业大学学报,2010,15(1):55-61.
[4]Jin D,Zhao SG,Zheng N,et al.Differences in ureolytic bacterial composition between the rumen digesta and rumen wall based on ureC Gene Classification.Front Microbiol,2017,8:385.
[5]Jin D,Zhao SG,Wang PP,et al.Insights into abundant rumen ureolytic bacterial community using rumen simulation system.Front Microbiol,2016,7:1006.
[6]Zhao SG,Wang JQ,Bu DP,et al.Biochemistry and molecular biology of bacterial ureases.Microbiology,2008,35(7):1146-1152(in Chinese).赵圣国,王加启,卜登攀,等.细菌尿素酶的生化和分子生物学特点.微生物学通报,2008,35(7):1146-1152.
[7]Toffanin A,Cadahia E,Imperial J,et al.Characterization of the urease gene cluster from Rhizobium leguminosarum cv.viciae.Arch Microbiol,2002,177(4):290-298.
[8]Maeda M,Hidaka M,Nakamura A,et al.Cloning,sequencing,and expression of thermophilic Bacillus sp.strain TB-90 urease gene complex in Escherichia coli.JBacteriol,1994,176(2):432-442.
[9]BosséJT,Gilmour HD,MacInnes JI.Novel genes affecting urease activity in Actinobacillus pleuropneumoniae.J Bacteriol,2001,183(4):1242-1247.
[10]Stoof J,Breijer S,Pot RGJ,et al.Inverse nickel-responsive regulation of two urease enzymes in the gastric pathogen Helicobacter mustelae.Environ Microbiol,2010,10(10):2586-2597.
[11]Kim JK,Mulrooney SB,Hausinger RP.Biosynthesis of active Bacillus subtilis urease in the absence of known urease accessory proteins.J Bacteriol,2005,187(20):7150-7154.
[12]Carter EL,Flugga N,Boer JL,et al.Interplay of metal ions and urease.Metallomics,2009,1(3):207-221.
[13]Benini S,Rypniewski WR,Wilson KS,et al.A new proposal for urease mechanism based on the crystal structures of the native and inhibited enzyme from Bacillus pasteurii:why urea hydrolysis costs two nickels.Structure,1999,7(2):205-216.
[14]Carter EL,Hausinger RP.Characterization of the Klebsiella aerogenes urease accessory protein UreD in fusion with the maltose binding protein.J Bacteriol,2010,192(9):2294-2304.
[15]Farrugia MA,Wang BB,Feig M,et al.Mutational and computational evidence that a nickel-transfer tunnel in UreD is used for activation of Klebsiella aerogenes urease.Biochemistry,2015,54(41):6392-6401.
[16]Park IS,Carr MB,Hausinger RP.In vitro activation of urease apoprotein and role of UreD as a chaperone required for nickel metallocenter assembly.Proc Natl Acad Sci USA,1994,91(8):3233-3237.
[17]Moncrief MB,Hausinger RP.Purification and activation properties of UreD-UreF-urease apoprotein complexes.JBacteriol,1996,178(18):5417-5421.
[18]Fong YH,Wong HC,Chuck CP,et al.Assembly of preactivation complex for urease maturation in Helicobacter pylori:crystal structure of UreF-UreHprotein complex.J Biol Chem,2011,286(50):43241-43249.
[19]Moncrief MB,Hausinger RP.Characterization of UreG,identification of a UreD-UreF-UreG complex,and evidence suggesting that a nucleotide-binding site in UreG is required for in vivo metallocenter assembly of Klebsiella aerogenes urease.J Bacteriol,1997,179(13):4081-4086.
[20]Zambelli B,Stola M,Musiani F,et al.UreG,a chaperone in the urease assembly process,is an intrinsically unstructured GTPase that specifically binds Zn2+.J Biol Chem,2005,280(6):4684-4695.
[21]Zambelli B,Musiani F,Savini M,et al.Biochemical studies on Mycobacterium tuberculosis UreG and comparative modeling reveal structural and functional conservation among the bacterial UreG family.Biochemistry,2007,46(11):3171-3182.
[22]Zambelli B,Turano P,Musiani F,et al.Zn2+-linked dimerization of UreG from Helicobacter pylori,a chaperone involved in nickel trafficking and urease activation.Proteins,2010,74(1):222-239.
[23]Leipe DD,Wolf YI,Koonin EV,et al.Classification and evolution of P-loop GTPases and related ATPases.J Mol Biol,2002,317(1):41-72.
[24]Bange G,Sinning I.SIMIBI twins in protein targeting and localization.Nat Struct Mol Biol,2013,20(7):776-780.
[25]Gasper R,Meyer S,Gotthardt K,et al.It takes two to tango:regulation of G proteins by dimerization.Nat Rev Mol Cell Biol,2009,10(6):423-429.
[26]Boer JL,Hausinger RP.Klebsiella aerogenes UreF:identification of the UreG binding site and role in enhancing the fidelity of urease activation.Biochemistry,2012,51(11):2298-2308.
[27]Yang XM,Li HY,Lai TP,et al.UreE-UreG complex facilitates nickel transfer and preactivates GTPase of UreG in Helicobacter pylori.J Biol Chem,2015,290(20):12474-12485.
[28]Lee MH,Pankratz HS,Wang SK,et al.Purification and characterization of Klebsiella aerogenes UreE protein:a nickel-binding protein that functions in urease metallocenter assembly.Protein Sci,2010,2(6):1042-1052.
[29]Brayman TG,Hausinger RP.Purification,characterization,and functional analysis of a truncated Klebsiella aerogenes UreE urease accessory protein lacking the histidine-rich carboxyl terminus.J Bacteriol,1996,178(18):5410-5416.
[30]Banaszak K,Martin-Diaconescu V,Bellucci M,et al.Crystallographic and X-ray absorption spectroscopic characterization of Helicobacter pylori UreE bound to Ni2+and Zn2+reveals a role for the disordered C-terminal arm in metal trafficking.Biochem J,2012,441(3):1017-1035.
[31]Shi R,Munger C,Asinas A,et al.Crystal structures of apo and metal-bound forms of the UreE protein from Helicobacter pylori:role of multiple metal binding sites.Biochemistry,2010,49(33):7080-7088.
[32]Song HK,Mulrooney SB,Huber R,et al.Crystal structure of Klebsiella aerogenes UreE,a nickel-binding metallochaperone for urease activation.J Biol Chem,2001,276(52):49359-49364.
[33]Remaut H,Safarov N,Ciurli S,et al.Structural basis for Ni2+transport and assembly of the urease active site by the metallochaperone UreE from Bacillus pasteurii.JBiol Chem,2001,276(52):49365-49370.
[34]Bellucci M,Zambelli B,Musiani F,et al.Helicobacter pylori UreE,a urease accessory protein:specific Ni2+and Zn2+binding properties and interaction with its cognate UreG.Biochem J,2009,422(1):91-100.
[35]Boer JL,Quiroz-Valenzuela S,Anderson KL,et al.Mutagenesis of Klebsiella aerogenes UreG to probe nickel binding and interactions with other urease-related proteins.Biochemistry,2010,49(28):5859-5869.
[36]Chan KH,Lee KM,Wong KB.Interaction between hydrogenase maturation factors HypA and HypB is required for[NiFe]-hydrogenase maturation.PLoS ONE,2012,7(2):e32592.
[37]Mehta N,Olson JW,Maier RJ.Characterization of Helicobacter pylori nickel metabolism accessory proteins needed for maturation of both urease and hydrogenase.J Bacteriol,2003,185(3):726-734.
[38]Yang XM,Li HY,Cheng TF,et al.Nickel translocation between metallochaperones HypA and UreE in Helicobacter pylori.Metallomics,2014,6(9):1731-1736.
[39]Sydor AM,Lebrette H,Ariyakumaran R,et al.Relationship between Ni(II)and Zn(II)coordination and nucleotide binding by the Helicobacter pylori[NiFe]-hydrogenase and urease maturation factor HypB.J Biol Chem,2014,289(7):3828-3841.
[40]Benoit SL,Zbell AL,Maier RJ.Nickel enzyme maturation in Helicobacter hepaticus:roles of accessory proteins in hydrogenase and urease activities.Microbiology,2007,153(11):3748-3756.
[41]Strugatsky D,McNulty R,Munson K,et al.Structure of the proton-gated urea channel from the gastric pathogen Helicobacter pylori.Nature,2013,493(7431):255-258.
[42]Cáceres-Delpiano J,Teneb J,Mansilla R,et al.Variations in periplasmic loop interactions determine the pH-dependent activity of the hexameric urea transporter UreI from Helicobacter pylori:a molecular dynamics study.BMC Struct Biol,2015,15(1):1-12.
[43]Sebbane F,Mandrand-Berthelot MA,Simonet M.Genes encoding specific nickel transport systems flank the chromosomal urease locus of Pathogenic yersiniae.JBacteriol,2002,184(20):5706-5713.
[44]Chen YYM,Burne RA.Identification and characterization of the nickel uptake system for urease biogenesis in Streptococcus salivarius 57.I.Virology,2003,183(23):6773-6779.
[45]Sigel A,Sigel H,Sigel RKO.Nickel and Its Surprising Impact in Nature,Volume 2.Hoboken,NJ:John Wiley&Sons,Ltd,2007:519-544.
[46]Musiani F,Gioia D,Masetti M,et al.Protein tunnels:the case of urease accessory proteins.J Chem Theory Comput,2017,13(5):2322-2331.
[47]Zambelli B,Berardi A,Martin-Diaconescu V,et al.Nickel binding properties of Helicobacter pylori UreF,an accessory protein in the nickel-based activation of urease.J Biol Inorg Chem,2014,19(3):319-334.
[48]Chang ZZ,Kuchar J,Hausinger RP.Chemical cross-linking and mass spectrometric identification of sites of interaction for UreD,UreF,and urease.J Biol Chem,2004,279(15):15305-15313.
[49]Quiroz-Valenzuela S,Sukuru SCK,Hausinger RP,et al.The structure of urease activation complexes examined by flexibility analysis,mutagenesis,and small-angle X-ray scattering.Arch Biochem Biophys,2008,480(1):51-57.
[50]Park IS,Hausinger RP.Requirement of carbon dioxide for in vitro assembly of the urease nickel metallocenter.Science,1995,267(5201):1156-1158.
[51]Carter EL,Boer JL,Farrugia MA,et al.Function of UreB in Klebsiella aerogenes urease.Biochemistry,2011,50(43):9296-9308.
[52]Soriano A,Colpas GJ,Hausinger RP.UreE stimulation of GTP-dependent urease activation in the UreD-UreF-UreG-urease apoprotein complex.Biochemistry,2000,39(40):12435-12440.
[53]Farrugia MA,Macomber L,Hausinger RP.Biosynthesis of the urease metallocenter.J Biol Chem,2013,288(19):13178-13185.
[54]Soriano A,Hausinger RP.GTP-dependent activation of urease apoprotein in complex with the UreD,UreF,and UreG accessory proteins.Proc Natl Acad Sci USA,1999,96(20):11140-11144.
[55]Fong YH,Wong HC,Yuen MH,et al.Structure of UreG/UreF/UreH complex reveals how urease accessory proteins facilitate maturation of Helicobacter pylori urease.PLoS Biol,2013,11(10):e1001678.
[56]Stingl K,Schauer K,Ecobichon C,et al.In vivo interactome of Helicobacter pylori urease revealed by tandem affinity purification.Mol Cell Proteomics,2008,7(12):2429-2441
[57]Farrugia MA,Han LJ,Zhong YY,et al.Analysis of a soluble(UreD:UreF:UreG)2 accessory protein complex and its interactions with Klebsiella aerogenes urease by mass spectrometry.J Am Soc Mass Spectrom,2013,24(9):1328-1337.
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