海洋蓝细菌生物固氮的研究进展

20320053ADVANCESINEARTHSCIENCEVol.20No.3Mar.,2005:100128166(2005)032031220831,2,1(1.,,361005;2.,445000):,CO2,,,,,:;;;;:Q178153:A,,,,[1](biogeochemicalcycle),[2],2:,;,,[1,2],[24](fixednitrogen),,[2]11000,20,,25100kg/hm2[3,5],,[3]1961(Trichodesmium)[2]Zehr,cyanobacteria(,),proteobacteria(),[2,3,5]3:2004203209;:2004207221.3:(:G1999043706);(:40076031);.:(19672),,,,,.E2mail:[e-mail]:(19642),,,,,.E2mail:[e-mail]©1995-2005TsinghuaTongfangOpticalDiscCo.,Ltd.Allrightsreserved.[6]15N(Trichodesmiumerythraeum)Trichodesmiumthiebautii,T.tenueT.contortum(T.hildebrandtii),Katagnymene,[5,6],Synechococcus[6],2(34m):Cyanothecespp.[7]Erythrosphaeramarina[8]3,3,[9],[10]Ze2hr[11]2000725m15N,320m,1050/mL,1000/mL[12],Trichodesmium(Trichodesmium:137mol/(m2d)[13]Zehr[11],,TrichodesmiumRhizosoleniaRichelia,nifH,[14]2(15N),[5,6],,,,[14,15],nifH(Fe),PCRDNA,[16],22(nifH),ALOHA,[15,17](BATS),(ALOHA),[17],[18][17,19],,[20],,,RT2PCR(PCR),[15],,,,,,,[15]3,10%,36%Trichodesmium87.4TgN[21],ZehrCO23133:©1995-2005TsinghuaTongfangOpticalDiscCo.,Ltd.Allrightsreserved.,CO2,CO2CO2,[5][22]CO2[15]N2NH3,[23]1961Trichodesmium,Trichodesmium[24],,,CO2[10,25],,Trichodesmium[10,25],Dugdale[2]:()(),,,(nif,N2NH3)[15],nif,,nif[14],CO2,,CO24NP,PFeO2,CO2[14,15,26],[27]4.1,PFe,PFe,(200250mol/L)[14],Fe[28],[26,29,30](DIP)(150m),DIPDOPP(P)PPNO-3N,P,()()2,Fe(DOFe)Fe,Fe[27,28]PNALOHA,N:P(161),,,NP(161),NPNP,,N,[26,27,29,30]4.2()Fu[31]2003,()Tri241320©1995-2005TsinghuaTongfangOpticalDiscCo.,Ltd.Allrightsreserved.chodesmiumsp.,200m[32],1525m510m[33],,4.2.1Ohki[34]Fu[31],GBRTRLI10168h,0,,,,[31],[24,31,34,35]Ohki[34],,,0Ohki[34],,Ber2man2Frank[36],,,TrichodesmiumO2:,,,O2,;,,,ATP,,,[34,37]4.2.2Fu[31],Trichodesmium;(10mol/(m2s)),8h,,,ATPNADPH[38]Bergman[36],ATP,FeFe,[39]4.2.3(),:[31]T,,,Kumar[40](Anabaenasp.)Fay[41],I,90%a(Chla)I,a,,,a,Trichodesmi2um,Trichodesmium,/,[21]4.3[42],,[43],[44],O2,[21,4146],,[42]5133:©1995-2005TsinghuaTongfangOpticalDiscCo.,Ltd.Allrightsreserved.NH3NH3[26,41,45,46],,,[26,41],,CO2,CO2N2CO2NOx,(35)[45,46],,[26,41],FeMoV(ROS),Fe4S4Fe2S,,[23,26,36,38],FeFe2+Fe3+,Mo,90%[23,26,47](FeVMo),Mo2[47]55.1,[43],,,:;(PPC),(DAP)(GAP)1,62(FBP),(PEP),[44,48,49],,NADPH,:NADP(FNR)(FdxH),2(Fe2FeMo2)NAD(P)H(RET),ATP,,I(PSI2)ATPPSINAD(P)H,[44,48]5.2,(nif)DNAnifHDK,nifEnifNnifBnifQFeMoco,nifSnifUnifMnifFnifJnifLA,nif,NH+4O2[44,50,51]nif,2nifLA,(ntr)nif2nifLnifA,,nifA,(NifA)57000,nif,NifA,35NifAnifNtrAnifL50000,NifLNifA,,(GS)ntr3ntrAntrBntrCnifA,84000GSntrBntrC,ntrBntrC3600056000ntrntrAntrC,nifLA,ntrBntrC[5153],,NtrA[53]61320©1995-2005TsinghuaTongfangOpticalDiscCo.,Ltd.Allrightsreserved.6,CO2,[2,3,14,15],,,[15]:(1)??(2)???(3)?(4)?(5)??(6)??,,,,,(References):[1]AllenCB,KandaJEA.NewproductionandphotosyntheticrateswithinandoutsideacyclonicmesoscaleeddyintheNorthPacificsubtropicalgyre[J].DeepSeaResearch,1996,43:9172936.[2]DugdaleRC,GoeringJJ.Uptakeofnewandregeneratedformsofnitrogeninprimaryproductivity[J].LimnologyandOceanogra2phy,1967,12:1962206.[3]DugdaleRC.Nutrientlimitationinthesea:dynamics,identifica2tionandsignificance[J].LimnologyandOceanography,1967,12:6852695.[4]BrandesJA,DevolAH.Aglobalmarine2fixednitrogenisotopicbudget:ImplicationsforHolocenenitrogencycling[J].GlobalBi2ogeochemicalCycles,2002,16(4):112021132.[5]CaponeDG.Marinenitrogenfixation:what’sthefuss?[J].E2cologyandIndustrialMicrobiology,2001,29(2):3412348.[6]CaponeDG.Trichodesmium:Agloballysignificantmarinecya2nobacterium[J].Science,1997,276:122121229.[7]ReddyKJ.Unicellular,aerobicnitrogen2fixingcyanobacteriaofthegenusCyanothece[J].JournalofBacteriology,1993,175:128421292.[8]SullivanMB,WaterburyJB,ChisholmSW.Cyanophagesinfec2tingtheoceaniccyanobacteriumProchlorococcus(vol424,pg1047,2003)[J].Nature,2003,426(6966):584.[9]NeveuxJ,LantoineF,VaulotD,etal.PhycoerythrinsinthesoutherntropicalandequatorialPacificOcean:Evidencefornewcyanobacterialtypes[J].JournalofGeophysicalResearch,1999,104:331123321.[10]GruberN,SarmientoJL.Globalpatternsofmarinenitrogenfixa2tionanddenitrification[J].GlobalBiogeochemistryCycles,1997,11:2352266.[11]ZehrJP,WaterburyJB,TurnerPJ,etal.Unicellularcya2nobacteriafixN2inthesubtropicalnorthPacificOcean[J].Na2ture,2001,412:6352638.[12]CampbellL,LiuH,NollaHA,etal.Annualvariabilityofphy2toplanktonandbacteriainthesubtropicalNorthPacificOceanatstationALOHAduringthe199121994ENSOevent[J].DeepSeaResearch,1997,144:1672192.[13]KarlDM.TheroleofnitrogenfixationinbiogeochemicalcyclinginthesubtropicalnorthPacificOcean[J].Nature,1997,388:5332538.[14]KarlDM.Anewsourceofnewnitrogeninthesea[J].TrendsinMicrobiology,2000,8(7):301.[15]ZehrJP,CarpenterEJ,VillarealTA.Newperspectivesonni2trogen2fixingmicroorganismsintropicalandsubtropicaloceans[J].TrendsinMicrobiology,2000,8(2):68273.[16]ZehrJP,BraunS,ChenY2B,etal.Nitrogenfixationinthemarineenvironment:Relatinggeneticpotentialtonitrogenaseac2tivity[J].JournalofExperimentalMarineBiologyandEcology,1996,203:61273.[17]ZehrJP.Newnitrogenfixingmicroorganismsdetectedinoligo2trophicoceansbytheamplificationofnitrogenase(nifH)genes[J].Appli