氨氧化古菌及其在氮循环中的重要作用刘晶静

＊刘晶静1　吴伟祥1,2＊＊　丁　颖1,2　石德智1　陈英旭1,2(1,310029;2,310029)　　氨氧化作用作为硝化过程的第一步,是氮素生物地球化学循环的关键步骤.长期以来,变形菌纲β和γ亚群中的氨氧化细菌一直被认为是氨氧化作用的主要承担者.然而,近年来研究发现氨氧化古菌广泛存在于各种生态系统中,并且在数量上占明显优势,在氮素生物地球化学循环中起着非常重要的作用.本文概述了氨氧化古菌的形态、生理生态特性及其系统发育特征,对比分析了氨氧化古菌和氨氧化细菌的氨单加氧酶及其编码基因的异同,综述了氨氧化古菌在水生和陆地等生态系统氮素循环中的作用,同时就氨氧化古菌在应用生态和环境保护领域今后的研究重点进行了展望.　氨氧化古菌　氨单加氧酶　氮循环　1001-9332(2010)08-2154-07　　X172　　AAmmonia-oxidizingarchaeaandtheirimportantrolesinnitrogenbiogeochemicalcycling:Areview.LIUJing-jing1,WUWei-xiang1,2,DINGYing1,2,SHIDe-zhi1,CHENYing-xu1,2(1InstituteofEnvironmentalScienceandTechnology,ZhejiangUniversity,Hangzhou310029,Chi-na;2ZhejiangProvinceKeyLaboratoryforWaterPollutionControlandEnvironmentalSafety,Hang-zhou310029,China).-Chin.J.Appl.Ecol.,2010,21(8):2154-2160.Abstract:Asthefirststepofnitrification,ammoniaoxidationisthekeyprocessinglobalnitrogenbiogeochemicalcycling.Sofar,theautotrophicammonia-oxidizingbacteria(AOB)intheβ-andγ-subgroupsofproteobacteriahavebeenconsideredasthemostimportantcontributorstoammoniaoxidation,buttherecentresearchesindicatedthatammonia-oxidizingarchaea(AOA)arewidelydistributedinvariouskindsofecosystemsandquantitativelypredominant,playingimportantrolesintheglobalnitrogenbiogeochemicalcycling.Thispaperreviewedthemorphological,physiological,andecologicalcharacteristicsandthemolecularphylogeniesofAOA,andcomparedandanalyzedthedifferencesandsimilaritiesoftheammoniamonooxygenase(AMO)anditsencodinggenesbe-tweenAOAandAOB.Inaddition,thepotentialsignificantrolesofAOAinnitrogenbiogeochemicalcyclinginaquaticandterrestrialecosystemsweresummarized,andthefutureresearchdirectionsofAOAinappliedecologyandenvironmentalprotectionwereputforward.Keywords:ammonia-oxidizingarchaea;ammoniamonooxygenase;nitrogencycling.＊“863”(2006AA06Z327)(2006C13066).＊＊.E-mail:[e-mail]2009-12-31,2010-06-01.　　[1-2].,.,(ammonia-oxidizingbacteria,AOB)[3].,[4],———(ammonia-oxidizingarchaea,AOA).、AOB,(Cren-archaeota).AOA[5-7]、[8]、[9-10][11],.AOA、,AOAAOB,、AOA　20108　21　8　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　ChineseJournalofAppliedEcology,Aug.2010,21(8):2154-2160DOI:10.13287/j.1001-9332.2010.0276、、.1　、2004,Venter[12]SargassoSea(AMO),.2005,Treusch[13].,Könneke[14]Nitrosopumilusmaritimus,Francis[15](ammonia-oxidizingarchaea,AOA).,.AOA,,,[7,16].,45℃AOA(mesophilicCrenarchaeota),45℃AOA(thermophilicCren-archaeota)[17-18].[10,18-19].,.AOAAOA[18].,[10,19].,[14,20].,.,1AOAN.maritimus[14].、;,0.5～0.9μm,0.17～0.22μm;.AOANitrososphaeragargensis[18]Nitrosocaldusyel-lowstonii[19]AOACenarchaeumsymbio-sum[20].AOA,20h,AOB[21].N.gargensis,3.08mmol·L-1,AOB200mmol·L-1[22-23].C.symbiosum3-,[24].C.symbiosumamoA、amoBamoC,、、、,AOBc554cm552.　　,AOAAOB.16SrRNAAOA(1).、AOAGroup1.1a(A);AOAGroup1.1b(B)[25].AOAamoA,Weidler[10],“thermalspringcluster”.,1.,AOA1　[25]Fig.1　Phylogenetictreeofmajorcrenarchaeotaandbranchstructure.A:Marinewaterandsediments(Group1.1a);B:Ubiquitousinmostsoilandotherenviron-ments(Group1.1b);C:Grasslandandco-niferousforestsoils;D:Sediments;E:、、Floodedsoils,sediments,freshwaterandotherenvironments;F:BMarinebenthicgroupB;G:CMarinebenthicgroupC;H:Subsurfaceplantroots;I:Terrestrialhotspringsandhydrothermalvents.21558　　　　　　　　　　　　　:　　　　　　,AOA.2　(ammoniamonooxygenase,AMO)AmoA、AmoBAmoC3[26],.3amoA、amoBamoCAOAAOB,AOAAOBAMO,25%,40%[25],[24].AOAAOBAmoA,,10[13].,AOAAOBAMO,.AmoAAMO[27-28],amoA、,、.AOAAOBamoA60%.,Arch-amoAFArch-amoARAOA[15],amoA-1FamoA-2RAOB[29].AOAAOBamoA.β-AOB2～3amoA,γ-AOB1;AOA2～3,AOA1[5,30].amoAAOAAOB.amo,AOAAOBamo,[25],amoA、amoBamoCAOAAOB(2).AOBamo,amoAamoB,amoBamoA,amoCamoA[30].AOAamo[25],.N.maritimusAOAamo,amoBamoC,amoAamoC,amoCamoA.AOAamoC,amoBamoAAOA[13].AOAamoamoC,amoAamoB[18-19].N.yellowstoniiAmoBAOAAmoB50%,30,　2　AOAAOBamo[19]Fig.2　Schematicrepresentationofamogenesindifferentspe-ciesofAOAandAOBNitrosomonaseuropaea:;SargassoSeascaffoldCH004643:AMO;SoilCrenarchaeotafos-mid54d9:AMO;Cenarchaeumsymbiosum,Ni-trosopumilusmaritimus,Candidatus“Nitrosocaldusyellowstonii”:AOATheAOAsobtainedbyenrichmentculture;D:Con-servedhypothetical;E:Connection.AOA[19].3　,AOA,.AOA,、、16SrRNAamoAAOA,.3.1　AOA,[5-6,31][7,32-33]、[32,34]、[35]、[36-37]、[38]、[37],amoA[39].,AOBamoA[5-6].AOBamoAAOA,[37]、[38][40],AOBamoAAOA.,2156　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　21,AOAamoA10～105copy·ml-1,107～108copy·g-1,AOBamoA1～3.AOAAOBamoA11071[5-6,34,39].,AOA,amoA,AOAAOB.,[41]、[42]、[31][43],amoA.9,amoA,AOBamoA[37].Wuchter[5]12,amoA.,Lam[7]AOA.,,AOA[44].AOA.AOAKmKm[16].10～20nmol·L-1,AOA,AOB(1μmol·L-1)100[45].N.maritimus,68700g-1·h-1,AOB200,30.,,AOAAOB,AOB、,AOA.3.2　AOA.,[46]、[8]、[47]、[48-49]、[50]、[51-52]、Littorellauniflora[11][53]AOA.Zhang[53](5700masl),AOBamoAAOA1～2.,AOAamoA104～108copy·g-1,AOBamoA2～3.、pHAOA.AOAamoA[13];AOA[52].AOA.Nicol[54]pH4.9～7.5,AOAamoApH;pH,AOAAOBamoA;,pH(8.3～8.7)AOA[46].(10℃～30℃)AOA[55].AOA.Tourna[55],AOAamoAAOB,AOB.Offre[56]AOAAOB,AOA,AOA.Chen[52],AOA,amoAAOB.,,AOB.,AOAamoA,AOBamoA[48].,AOA,AOB,[49].AOA;AOB,AOA,.3.3　AOA、,.[57]、[58][59-60]amoA.21578　　　　　　　　　　　　　:　　　　　　AOA,AOA.AOA.AOApH,38℃～97℃pH2.5～9.0、[9-10,61-62].,AOBAOA.4　AOA,、、、.AOAAOB,AOB,,AOAAOB.AOA.,AOA.,AOAAOAAOB,,,:1)、AOA