微生物酶法消除黄酒中氨基甲酸乙酯研究进展
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摘要
氨基甲酸乙酯(Ethyl carbamate,EC)具有致癌性,广泛存在于酒精饮料中。我国的黄酒因EC含量高而带来的食品安全问题越来越受到人们的关注。微生物酶法消除黄酒中的EC具有直接、高效的特性而被深入研究。文中从黄酒中EC的形成机制、酸性脲酶研究现状、氨基甲酸乙酯水解酶研究现状等方面概述了微生物酶法消除黄酒中EC的研究进展及存在的问题。并针对这些问题,提出了寻找新型氨基甲酸乙酯水解酶、Fe~(3+)依赖型双功能酸性脲酶食品级表达与定向进化及双酶并用将尿素和EC一起消除的策略。
Ethyl carbamate(EC), a carcinogenic and teratogenic chemical that is widely distributed in various alcoholic beverages, has attracted much attention. Microbial enzymatic degradation of EC in rice wine is always efficient and attractive.In this review, we summarize the research progress and problems of microbial enzymatic elimination of EC in rice wine from three aspects: the mechanisms of EC formation in rice wine, the research progress of acid urease, and the research progress of urethanase. Then, we propose the corresponding strategies to solve the problems: screening new urethanase with satisfied enzyme properties, food-grade expression and directed evolution of the bifunctional Fe~(3+)-dependent acid urease and acid urease used in combination with urethanase to eliminate both urea and EC in rice wine.
Ethyl carbamate(EC), a carcinogenic and teratogenic chemical that is widely distributed in various alcoholic beverages, has attracted much attention. Microbial enzymatic degradation of EC in rice wine is always efficient and attractive.In this review, we summarize the research progress and problems of microbial enzymatic elimination of EC in rice wine from three aspects: the mechanisms of EC formation in rice wine, the research progress of acid urease, and the research progress of urethanase. Then, we propose the corresponding strategies to solve the problems: screening new urethanase with satisfied enzyme properties, food-grade expression and directed evolution of the bifunctional Fe~(3+)-dependent acid urease and acid urease used in combination with urethanase to eliminate both urea and EC in rice wine.
引文
[1]Forkert PG.Mechanisms of lung tumorigenesis by ethyl carbamate and vinyl carbamate.Drug Metab Rev,2010,42(2):355-378.
[2]Weber JV,Sharypov VI.Ethyl carbamate in foods and beverages:a review.Environ Chem Lett,2009,7(3):233-247.
[3]Zhao XR,Du GC,Zou HJ,et al.Progress in preventing the accumulation of ethyl carbamate in alcoholic beverages.Trends Food Sci Technol,201332(2):97-107.
[4]Gowd V,Su HM,Karlovsky P,et al.Ethyl carbamate:an emerging food and environmental toxicant.Food Chem,2018,248:312-321.
[5]Lachenmeier DW.Consequences of IARCre-evaluation of alcoholic beverage consumption and ethyl carbamate on food control.Deut Lebensm Rundsch,2007,103:307-311.
[6]Chen DW,Ren YP,Zhong QD,et al.Ethyl carbamate in alcoholic beverages from China:levels,dietary intake,and risk assessment.Food Control,2017,72:283-288.
[7]Yoshizawa K,Takahashi K,Sato K.Changes of urea content in rice and sake moromi during sake making process.Nippon Jozo Kyokaishi,1988,83:136-141.
[8]Liu SQ,Pilone GJ.A REVIEW:arginine metabolism in wine lactic acid bacteria and its practical significance.J Appl Microbiol,1998,84(3):315-327.
[9]Hasnip S,Caputi A,Crews C,et al.Effects of storage time and temperature on the concentration of ethyl carbamate and its precursors in wine.Food Addit Contam,2004,21(12):1155-1161.
[10]Fang RS.The metabolism mechanism and inhibition method of ethyl caramate formation during traditional Chinese rice wine fermentation[D].HangZhou:Zhejiang University,2017(in Chinese).方若思.传统黄酒发酵中氨基甲酸乙酯产生的代谢规律及抑制方法研究[D].杭州:浙江大学,2017.
[11]Wu HM,Chen L,Pan GS,et al.Study on the changing concentration of ethyl carbamate in yellow rice wine during production and storage by gas chromatography/mass spectrometry.Eur Food Res Technol,2012,235(5):779-782.
[12]Park SR,Ha SD,Yoon JH,et al.Exposure to ethyl carbamate in alcohol-drinking and nondrinking adults and its reduction by simple charcoal filtration.Food Control,2009,20(10):946-952.
[13]Liu J,Zhao GA,Xu Y.Directly removal of ethylcarbamate in Chinese rice wine.J Food Sci Biotechnol,2012,31(2):171-176(in Chinese).刘俊,赵光鳌,徐岩.黄酒中氨基甲酸乙酯直接减除技术的研究.食品与生物技术学报,2012,31(2):171-176.
[14]Suizu T,Iimura Y,Gomi K,et al.Construction of urea non-producing yeast Saccharomyces cerevisiae by disruption of the CAR1 gene.Agric Biol Chem,1990,54(2):537-539.
[15]Kitamoto K,Oda K,Gomi K,et al.Genetic engineering of a sake yeast producing no urea by successive disruption of arginase gene.Appl Environ Microbiol,1991,57(1):301-306.
[16]Wu DH,Li XM,Shen C,et al.Decreased ethyl carbamate generation during Chinese rice wine fermentation by disruption of CAR1 in an industrial yeast strain.Int J Food Microbiol,2014,180:19-23.
[17]Wu D,Li X,Lu J,et al.Constitutive expression of the DUR1,2 gene in an industrial yeast strain to minimize ethyl carbamate production during Chinese rice wine fermentation.FEMS Microbiol Lett,2016,363(1):fnv214.
[18]Coulon J,Husnik JI,Inglis DL,et al.Metabolic engineering of Saccharomyces cerevisiae to minimize the production of ethyl carbamate in wine.Am J Enol Viticult,2006,57(2):113-124.
[19]Dahabieh MS,Husnik JI,van Vuuren HJJ.Functional enhancement of Sake yeast strains to minimize the production of ethyl carbamate in Sake wine.J Appl Microbiol,2010,109(3):963-973.
[20]Zhang P,Hu X.Metabolic engineering of arginine permeases to reduce the formation of urea in Saccharomyces cerevisiae.World J Microbiol Biotechnol,2018,34:47.
[21]Mohapatra BR.An Insight into the prevalence and enzymatic abatement of urethane in fermented beverages//Patra JK,Das G,Shin HS,eds.Microbial Biotechnology.Singapore:Springer Singapore,2018:153-170.
[22]Wu DH,Li XM,Sun JY,et al.Effect of citrulline metabolism in Saccharomyces cerevisiae on the formation of ethyl carbamate during Chinese rice wine fermentation.J Inst Brew,2018,124(1):77-84.
[23]An D,Ough CS.Urea excretion and uptake by wine yeasts as affected by various factors.Am J Enol Viticult,1993,44(1):35-40.
[24]Schehl B,Senn T,Lachenmeier DW,et al.Contribution of the fermenting yeast strain to ethyl carbamate generation in stone fruit spirits.Appl Microbiol Biotechnol,2007,74(4):843-850.
[25]Azevedo Z,Couto JA,Hogg T.Citrulline as the main precursor of ethyl carbamate in model fortified wines inoculated with Lactobacillus hilgardii:a marker of the levels in a spoiled fortified wine.Lett Appl Microbiol,2002,34(1):32-36.
[26]Sumner JB.The isolation and crystallization of the enzyme urease.Preliminary paper.J Biol Chem1926,69(2):435-441.
[27]Krajewska B.Ureases I.functional,catalytic and kinetic properties:a review.J Mol Catal B:Enzym,2009,59(1/3):9-21.
[28]Mobley HL,Hausinger RP.Microbial ureases:significance,regulation,and molecular characterization.Microbiol Rev,1989,53(1):85-108.
[29]Balasubramanian A,Ponnuraj K.Crystal structure of the first plant urease from Jack Bean:83 years of journey from its first crystal to molecular structure.JMol Biol,2010,400(3):274-283.
[30]Carter EL,Flugga N,Boer JL,et al.Interplay of metal ions and urease.Metallomics,2009,1(3):207-221.
[31]Dixon NE,Gazzola C,Blakeley RL,et al.Jack bean urease(EC 3.5.1.5).Metalloenzyme.Simple biological role for nickel.J Am Chem Soc,1975,97(14):4131-4133.
[32]Farrugia MA,Macomber L,Hausinger RP.Biosynthesis of the urease metallocenter.J Biol Chem,2013,288(19):13178-13185.
[33]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.
[34]McDonald JA,Vorhaben JE,Campbell JWInvertebrate urease:purification and properties of the enzyme from a land snail,Otala lactea.Comp Biochem Physiol Part B:Comp Biochem,1980,66(2):223-231.
[35]Suzuki K,Benno Y,Mitsuoka T,et al.Urease-producing species of intestinal anaerobes and their activities.Appl Environ Microbiol,1979,37(3):379-382.
[36]Miyagawa K,Sumida M,Nakao M,et al.Purification,characterization,and application of an acid urease from Arthrobacter mobilis.J Biotechnol,1999,68(2/3):227-236.
[37]Yang LQ,Wang SH,Tian YP.Purification,properties,and application of a novel acid urease from Enterobacter sp.Appl Biochem Biotechnol2010,160(2):303-313.
[38]Mora D,Maguin E,Masiero M,et al.Characterization of urease genes cluster of Streptococcus thermophilus.J Appl Microbiol,2004,96(1):209-219.
[39]Zotta T,Ricciardi A,Rossano R,et al.Urease production by Streptococcus thermophilus.Food Microbiol,2008,25(1):113-119.
[40]Chen YY,Clancy KA,Burne RA.Streptococcus salivarius urease:genetic and biochemical characterization and expression in a dental plaque Streptococcus.Infect Immun,1996,64(2):585-592.
[41]Yamazaki E,Kurasawa T,Kakimoto S,et al Characteristics of acid urease from Streptococcus mitior.Agric Biol Chem,1990,54(9):2433-2435.
[42]Kakimoto S,Sumino Y,Akiyama SI,et al.Purification and characterization of acid urease from Lactobacillus reuteri.Agric Biol Chem,1989,53(4):1119-1125.
[43]Kakimoto S,Sumino Y,Kawahara K,et al.Purification and characterization of acid urease from Lactobacillus fermentum.Appl Microbiol Biotechnol,1990,32(5):538-543.
[44]Yang YQ,Kang Z,Zhou JL,et al.High-level expression and characterization of recombinant acid urease for enzymatic degradation of urea in rice wine.Appl Microbiol Biotechnol,2015,99(1):301-308.
[45]Liu QT,Chen YQ,Yuan ML,et al.A Bacillus paralicheniformis iron-containing urease reduces urea concentrations in rice wine.Appl Environ Microbiol,2017,83(17):e01258-17.
[46]Liu J,Xu Y,Nie Y,et al.Optimization production of acid urease by Enterobacter sp.in an approach to reduce urea in Chinese rice wine.Bioprocess Biosyst Eng,2012,35(4):651-657.
[47]Zhou JL,Kang Z,Liu QT,et al.Degradation of urea and ethyl carbamate in Chinese Rice wine by recombinant acid urease.Chin J Biotech,2016,32(1):74-83(in Chinese).周建立,康振,刘庆涛,等.重组酸性脲酶对黄酒中尿素和氨基甲酸乙酯的降解应用.生物工程学报,2016,32(1):74-83.
[48]Liu XF,Zhang Q,Zhou ND,et al.Expression of an acid urease with urethanase activity in E.coli and analysis of urease gene.Mol Biotechnol,2017,59(2/3):84-97.
[49]Wang SM,He J,Cui ZL,et al.Self-formed adaptor PCR:a simple and efficient method for chromosome walking.Appl Environ Microbiol,2007,73(15):5048-5051.
[50]Juminaga D,Baidoo EEK,Redding-Johanson AM,et al.Modular engineering of L-tyrosine production in Escherichia coli.Appl Environ Microbiol,2012,78(1):89-98.
[51]Bauerfeind P,Garner RM,Mobley LT.Allelic exchange mutagenesis of nixA in Helicobacter pylori results in reduced nickel transport and urease activity.Infect Immun,1996,64(7):2877-2880.
[52]Chen YYM,Burne RA.Identification and characterization of the nickel uptake system for urease biogenesis in Streptococcus salivarius 57.I.JBacteriol,2003,185(23):6773-6779.
[53]Kobashi K,Takebe S,Sakai T.Urethane-hydrolyzing enzyme from Citrobacter sp.Chem Pharm Bull1990,38(5):1326-1328.
[54]Akutsu-Shigeno Y,Adachi Y,Yamada C,et al.Isolation of a bacterium that degrades urethane compounds and characterization of its urethane hydrolase.Appl Microbiol Biotechnol,2006,70(4):422-429.
[55]Liu XH,Fang F,Xia XL,et al.Stability enhancement of urethanase from Lysinibacillus fusiformis by site-directed mutagenesis.Chin J Biotech,201632(9):1233-1242(in Chinese).刘晓慧,方芳,夏小乐,等.定点突变改造提高纺锤形赖氨酸芽孢杆菌氨基甲酸乙酯水解酶稳定性.生物工程学报,2016,32(9):1233-1242.
[56]Zhao CJ,Imamura L,Kobashi K.Urethanase of Bacillus licheniformis sp.isolated from mouse gastrointestine.Chem Pharm Bull,1991,39(12):3303-3306.
[57]Zhao C,Kobashi K.Purification and characterization of iron-containing urethanase from Bacillus licheniformis.Biol Pharm Bull,1994,17(6):773-778.
[58]Mohapatra B,Bapuji M.Characterization of urethanase from Micrococcus species associated with the marine sponge(Spirasfrella species).Lett Appl Microbiol,1997,25(6):393-396.
[59]Wu Q,Zhao Y,Wang D,et al.Immobilized Rhodotorula mucilaginosa:a novel urethanase-producing strain for degrading ethyl carbamate.Appl Biochem Biotechnol,2013,171(8):2220-2232.
[60]Zhou ND,Gu X,Tian Y.Isolation and characterization of urethanase from Penicillium variabile and its application to reduce ethyl carbamate contamination in chinese rice wine.Appl Biochem Biotechnol,2013,170(3):718-728.
[61]Bu PP,Chen J,Du GC.Purification and characterization of a halophilic urethanase from Klebsiella pneumoniae.Chin J Biotech,2014,30(12):404-411(in Chinese).卜攀攀,陈坚,堵国成,等.耐盐氨基甲酸乙酯水解酶的分离纯化及酶学性质.生物工程学报,2014,30(12):404-411.
[2]Weber JV,Sharypov VI.Ethyl carbamate in foods and beverages:a review.Environ Chem Lett,2009,7(3):233-247.
[3]Zhao XR,Du GC,Zou HJ,et al.Progress in preventing the accumulation of ethyl carbamate in alcoholic beverages.Trends Food Sci Technol,201332(2):97-107.
[4]Gowd V,Su HM,Karlovsky P,et al.Ethyl carbamate:an emerging food and environmental toxicant.Food Chem,2018,248:312-321.
[5]Lachenmeier DW.Consequences of IARCre-evaluation of alcoholic beverage consumption and ethyl carbamate on food control.Deut Lebensm Rundsch,2007,103:307-311.
[6]Chen DW,Ren YP,Zhong QD,et al.Ethyl carbamate in alcoholic beverages from China:levels,dietary intake,and risk assessment.Food Control,2017,72:283-288.
[7]Yoshizawa K,Takahashi K,Sato K.Changes of urea content in rice and sake moromi during sake making process.Nippon Jozo Kyokaishi,1988,83:136-141.
[8]Liu SQ,Pilone GJ.A REVIEW:arginine metabolism in wine lactic acid bacteria and its practical significance.J Appl Microbiol,1998,84(3):315-327.
[9]Hasnip S,Caputi A,Crews C,et al.Effects of storage time and temperature on the concentration of ethyl carbamate and its precursors in wine.Food Addit Contam,2004,21(12):1155-1161.
[10]Fang RS.The metabolism mechanism and inhibition method of ethyl caramate formation during traditional Chinese rice wine fermentation[D].HangZhou:Zhejiang University,2017(in Chinese).方若思.传统黄酒发酵中氨基甲酸乙酯产生的代谢规律及抑制方法研究[D].杭州:浙江大学,2017.
[11]Wu HM,Chen L,Pan GS,et al.Study on the changing concentration of ethyl carbamate in yellow rice wine during production and storage by gas chromatography/mass spectrometry.Eur Food Res Technol,2012,235(5):779-782.
[12]Park SR,Ha SD,Yoon JH,et al.Exposure to ethyl carbamate in alcohol-drinking and nondrinking adults and its reduction by simple charcoal filtration.Food Control,2009,20(10):946-952.
[13]Liu J,Zhao GA,Xu Y.Directly removal of ethylcarbamate in Chinese rice wine.J Food Sci Biotechnol,2012,31(2):171-176(in Chinese).刘俊,赵光鳌,徐岩.黄酒中氨基甲酸乙酯直接减除技术的研究.食品与生物技术学报,2012,31(2):171-176.
[14]Suizu T,Iimura Y,Gomi K,et al.Construction of urea non-producing yeast Saccharomyces cerevisiae by disruption of the CAR1 gene.Agric Biol Chem,1990,54(2):537-539.
[15]Kitamoto K,Oda K,Gomi K,et al.Genetic engineering of a sake yeast producing no urea by successive disruption of arginase gene.Appl Environ Microbiol,1991,57(1):301-306.
[16]Wu DH,Li XM,Shen C,et al.Decreased ethyl carbamate generation during Chinese rice wine fermentation by disruption of CAR1 in an industrial yeast strain.Int J Food Microbiol,2014,180:19-23.
[17]Wu D,Li X,Lu J,et al.Constitutive expression of the DUR1,2 gene in an industrial yeast strain to minimize ethyl carbamate production during Chinese rice wine fermentation.FEMS Microbiol Lett,2016,363(1):fnv214.
[18]Coulon J,Husnik JI,Inglis DL,et al.Metabolic engineering of Saccharomyces cerevisiae to minimize the production of ethyl carbamate in wine.Am J Enol Viticult,2006,57(2):113-124.
[19]Dahabieh MS,Husnik JI,van Vuuren HJJ.Functional enhancement of Sake yeast strains to minimize the production of ethyl carbamate in Sake wine.J Appl Microbiol,2010,109(3):963-973.
[20]Zhang P,Hu X.Metabolic engineering of arginine permeases to reduce the formation of urea in Saccharomyces cerevisiae.World J Microbiol Biotechnol,2018,34:47.
[21]Mohapatra BR.An Insight into the prevalence and enzymatic abatement of urethane in fermented beverages//Patra JK,Das G,Shin HS,eds.Microbial Biotechnology.Singapore:Springer Singapore,2018:153-170.
[22]Wu DH,Li XM,Sun JY,et al.Effect of citrulline metabolism in Saccharomyces cerevisiae on the formation of ethyl carbamate during Chinese rice wine fermentation.J Inst Brew,2018,124(1):77-84.
[23]An D,Ough CS.Urea excretion and uptake by wine yeasts as affected by various factors.Am J Enol Viticult,1993,44(1):35-40.
[24]Schehl B,Senn T,Lachenmeier DW,et al.Contribution of the fermenting yeast strain to ethyl carbamate generation in stone fruit spirits.Appl Microbiol Biotechnol,2007,74(4):843-850.
[25]Azevedo Z,Couto JA,Hogg T.Citrulline as the main precursor of ethyl carbamate in model fortified wines inoculated with Lactobacillus hilgardii:a marker of the levels in a spoiled fortified wine.Lett Appl Microbiol,2002,34(1):32-36.
[26]Sumner JB.The isolation and crystallization of the enzyme urease.Preliminary paper.J Biol Chem1926,69(2):435-441.
[27]Krajewska B.Ureases I.functional,catalytic and kinetic properties:a review.J Mol Catal B:Enzym,2009,59(1/3):9-21.
[28]Mobley HL,Hausinger RP.Microbial ureases:significance,regulation,and molecular characterization.Microbiol Rev,1989,53(1):85-108.
[29]Balasubramanian A,Ponnuraj K.Crystal structure of the first plant urease from Jack Bean:83 years of journey from its first crystal to molecular structure.JMol Biol,2010,400(3):274-283.
[30]Carter EL,Flugga N,Boer JL,et al.Interplay of metal ions and urease.Metallomics,2009,1(3):207-221.
[31]Dixon NE,Gazzola C,Blakeley RL,et al.Jack bean urease(EC 3.5.1.5).Metalloenzyme.Simple biological role for nickel.J Am Chem Soc,1975,97(14):4131-4133.
[32]Farrugia MA,Macomber L,Hausinger RP.Biosynthesis of the urease metallocenter.J Biol Chem,2013,288(19):13178-13185.
[33]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.
[34]McDonald JA,Vorhaben JE,Campbell JWInvertebrate urease:purification and properties of the enzyme from a land snail,Otala lactea.Comp Biochem Physiol Part B:Comp Biochem,1980,66(2):223-231.
[35]Suzuki K,Benno Y,Mitsuoka T,et al.Urease-producing species of intestinal anaerobes and their activities.Appl Environ Microbiol,1979,37(3):379-382.
[36]Miyagawa K,Sumida M,Nakao M,et al.Purification,characterization,and application of an acid urease from Arthrobacter mobilis.J Biotechnol,1999,68(2/3):227-236.
[37]Yang LQ,Wang SH,Tian YP.Purification,properties,and application of a novel acid urease from Enterobacter sp.Appl Biochem Biotechnol2010,160(2):303-313.
[38]Mora D,Maguin E,Masiero M,et al.Characterization of urease genes cluster of Streptococcus thermophilus.J Appl Microbiol,2004,96(1):209-219.
[39]Zotta T,Ricciardi A,Rossano R,et al.Urease production by Streptococcus thermophilus.Food Microbiol,2008,25(1):113-119.
[40]Chen YY,Clancy KA,Burne RA.Streptococcus salivarius urease:genetic and biochemical characterization and expression in a dental plaque Streptococcus.Infect Immun,1996,64(2):585-592.
[41]Yamazaki E,Kurasawa T,Kakimoto S,et al Characteristics of acid urease from Streptococcus mitior.Agric Biol Chem,1990,54(9):2433-2435.
[42]Kakimoto S,Sumino Y,Akiyama SI,et al.Purification and characterization of acid urease from Lactobacillus reuteri.Agric Biol Chem,1989,53(4):1119-1125.
[43]Kakimoto S,Sumino Y,Kawahara K,et al.Purification and characterization of acid urease from Lactobacillus fermentum.Appl Microbiol Biotechnol,1990,32(5):538-543.
[44]Yang YQ,Kang Z,Zhou JL,et al.High-level expression and characterization of recombinant acid urease for enzymatic degradation of urea in rice wine.Appl Microbiol Biotechnol,2015,99(1):301-308.
[45]Liu QT,Chen YQ,Yuan ML,et al.A Bacillus paralicheniformis iron-containing urease reduces urea concentrations in rice wine.Appl Environ Microbiol,2017,83(17):e01258-17.
[46]Liu J,Xu Y,Nie Y,et al.Optimization production of acid urease by Enterobacter sp.in an approach to reduce urea in Chinese rice wine.Bioprocess Biosyst Eng,2012,35(4):651-657.
[47]Zhou JL,Kang Z,Liu QT,et al.Degradation of urea and ethyl carbamate in Chinese Rice wine by recombinant acid urease.Chin J Biotech,2016,32(1):74-83(in Chinese).周建立,康振,刘庆涛,等.重组酸性脲酶对黄酒中尿素和氨基甲酸乙酯的降解应用.生物工程学报,2016,32(1):74-83.
[48]Liu XF,Zhang Q,Zhou ND,et al.Expression of an acid urease with urethanase activity in E.coli and analysis of urease gene.Mol Biotechnol,2017,59(2/3):84-97.
[49]Wang SM,He J,Cui ZL,et al.Self-formed adaptor PCR:a simple and efficient method for chromosome walking.Appl Environ Microbiol,2007,73(15):5048-5051.
[50]Juminaga D,Baidoo EEK,Redding-Johanson AM,et al.Modular engineering of L-tyrosine production in Escherichia coli.Appl Environ Microbiol,2012,78(1):89-98.
[51]Bauerfeind P,Garner RM,Mobley LT.Allelic exchange mutagenesis of nixA in Helicobacter pylori results in reduced nickel transport and urease activity.Infect Immun,1996,64(7):2877-2880.
[52]Chen YYM,Burne RA.Identification and characterization of the nickel uptake system for urease biogenesis in Streptococcus salivarius 57.I.JBacteriol,2003,185(23):6773-6779.
[53]Kobashi K,Takebe S,Sakai T.Urethane-hydrolyzing enzyme from Citrobacter sp.Chem Pharm Bull1990,38(5):1326-1328.
[54]Akutsu-Shigeno Y,Adachi Y,Yamada C,et al.Isolation of a bacterium that degrades urethane compounds and characterization of its urethane hydrolase.Appl Microbiol Biotechnol,2006,70(4):422-429.
[55]Liu XH,Fang F,Xia XL,et al.Stability enhancement of urethanase from Lysinibacillus fusiformis by site-directed mutagenesis.Chin J Biotech,201632(9):1233-1242(in Chinese).刘晓慧,方芳,夏小乐,等.定点突变改造提高纺锤形赖氨酸芽孢杆菌氨基甲酸乙酯水解酶稳定性.生物工程学报,2016,32(9):1233-1242.
[56]Zhao CJ,Imamura L,Kobashi K.Urethanase of Bacillus licheniformis sp.isolated from mouse gastrointestine.Chem Pharm Bull,1991,39(12):3303-3306.
[57]Zhao C,Kobashi K.Purification and characterization of iron-containing urethanase from Bacillus licheniformis.Biol Pharm Bull,1994,17(6):773-778.
[58]Mohapatra B,Bapuji M.Characterization of urethanase from Micrococcus species associated with the marine sponge(Spirasfrella species).Lett Appl Microbiol,1997,25(6):393-396.
[59]Wu Q,Zhao Y,Wang D,et al.Immobilized Rhodotorula mucilaginosa:a novel urethanase-producing strain for degrading ethyl carbamate.Appl Biochem Biotechnol,2013,171(8):2220-2232.
[60]Zhou ND,Gu X,Tian Y.Isolation and characterization of urethanase from Penicillium variabile and its application to reduce ethyl carbamate contamination in chinese rice wine.Appl Biochem Biotechnol,2013,170(3):718-728.
[61]Bu PP,Chen J,Du GC.Purification and characterization of a halophilic urethanase from Klebsiella pneumoniae.Chin J Biotech,2014,30(12):404-411(in Chinese).卜攀攀,陈坚,堵国成,等.耐盐氨基甲酸乙酯水解酶的分离纯化及酶学性质.生物工程学报,2014,30(12):404-411.