产油微生物油脂生物合成与代谢调控研究进展

45120052ActaMicrobiologicaSinicaVol.45FebruaryNo.120053TelPFax:862411284379211;E2mail:[e-mail]:(1969-),(),,,,:2004205224,:20042112083(116023):20%,,,,,,:,,,:Q93919:A:000126209(2005)0120153204,(Endomyces)(Fusarium),,,,,,,,,,,,,,1,20%,(Oleaginousmicroorganisms)[1],,,,70%,(Triacylglycerol,TAG),80%,10%TAG,,TAG,,,,,,[2],-(GLA)[3](ARA)[4](EPA)[5](DHA)[6]PapanikolaouS.M.isabellina(100gPL),1811gPL,[7],,,,(Bio2diesel)2211TAG,[8],TAG(PA)(DAG)(S.cerevisiae)PAPA,,232(G232P)(DHAP)232(GAT)sn21,(LPA),DHAP(DHAPAT)sn21,12acyl2DHAP(ADR)LPATAG,Gat1pGat2pG232PDHAP,sn21[9](AGAT)sn22,PAPAD(PAP)DAG,DAG,DAG©1994-2007ChinaAcademicJournalElectronicPublishingHouse.Allrightsreserved.[10]DAGTAGDga1pLro1p,(DAGAT),TAGCoA;,:(PDAT),sn22TAGsn23[11]CoA:(ACAT)Are1pAre2p,CoA,DAG[12],DAGTAG[13],TAG[14](R.glutinis)TAG,(ACP)2ACPAGATPAPDAGATLPACoATAG,,TAG,(FeSOD)TAG,()[15]EMSTAG,TAG[16],TAG,TAG,PA;TAG212,,,,CoATAG,,,TAG,1TAG,TAG,,(AMP),AMPAMP(IMP),[17](ICDH)AMP,AMP[18],22,(TCA),1Fig.1OutlineofthemetabolicregulationoflipidaccumulationinoleaginousyeastsACL:ATP:Citratelyase;CS:Citratesynthase;FAS:Fattyacidsynthase;ICDH:Isocitratedehydrogenase;MDH:Malatedehydrogenase;ME:Malicenzyme;PDH:Pyruvatedehydrogenase;PYC:Pyruvatecarboxylase.P[19],ATP:(ACL)CoA,CoA,(FAS)CoA,M.circinelloidesM.alpinaICDHAMP[20]AMP,ICDH,(Zygomycetes),ICDH,[21]ICDH,,,ICDH,AMP,AMP,,?,ACL(ME),CoAACL,TAG20%ACLACL,10%,,[22]ACLACL(MDH),ME,NADPH,,NADPH(PDH)CoA,45145©1994-2007ChinaAcademicJournalElectronicPublishingHouse.Allrightsreserved.(PYC)(CS),ACL,TAG,CoACoA,NADPHCoANADPH,ME,ME,NADPH,FASMENADPH(1)[23],FASME,(Lipogenicmetabolon)[24]M.circinelloidesME,[25]MEdosSantos[26],,ME,,,,ME,TAG,M.isabellinaTAG;C.echinulata,TAG[21]TAG,,,,,,[1]RatledgeC,WynnJ.Thebiochemistryandmolecularbiologyoflipidaccumulationinoleaginousmicroorganisms.AdvApplMicrobiol,2002,51:1-511[2]CertikM,ShimizuS.Biosynthesisandregulationofmicrobialpolyunsaturatedfattyacidproduction.JBiosciBioeng,1999,87:1-141[3]SomashekarD,VenkateshwaranG,SambaiahK,etal.Effectofcultureconditionsonlipidandgamma2linolenicacidproductionbymucoraceousfungi.ProcessBiochemistry,2002,38:1719-17241[4],,,..,1999,29:1-3.[5]BajpaiP,BajpaiPK,WardOP.Eicosapentaenoicacid(EPA)productionbyMortierellaalpinaATCC322221ApplBiochemBiotechnol,1991,31:267-2721[6]SinghA,WardO.Microbialproductionofdocosahexaenoicacid(DHA,C22:6).AdvApplMicrobiol,1997,45:271-3121[7]PapanikolaouS,KomaitisM,AggelisG.Singlecelloil(SCO)productionbyMortierellaisabellingrownonhigh2sugarcontentmedia.BioresourTechnol,2004,95:287-2911[8]SorgerD,DaumG.Triacylglycerolbiosynthesisinyeast.ApplMicrobiolBiotechnol,2003,61:289-2991[9]ZhengZ,ZouJ.Theinitialstepoftheglycerolipidpathway:identificationofglycerol32phosphatePdihydroxyacetonephosphatedualsubstrateacyltransferasesinSaccharomycescerevisiae.JBiolChem,2001,276:1710-17161[10]OelkersP,CromleyD,PadamseeM,etal.TheDAG1genedeterminesasecondtriglyceridesyntheticpathwayinyeast.JBiolChem,2002,277:8877-88811[11]DahlqvistA,StahlU,LanmanM,etal.Phospholipid:diacylglycerolacyl2transferase:anenzymethatcatalyzestheacyl2CoA2independentformationoftriacylglycerolinyeastandplants.ProcNatlAcadSciUSA,2000,12:6487-64921[12]SandagerL,GustavssonM,StahlU,etal.Storagelipidsynthesisisnon2essentialinyeast.JBiolChem,2002,277:6478-64821[13]SorgerD,DaumG.SynthesisoftriacylglycerolsbytheacylcoenzymeA:diacyl2glycerolacyltransferaseDga1pinlipidparticlesoftheyeastSaccharomycescerevisia.JBacteriol,2002,184:519-5241[14]GangarA,KarandeAA,RajasekharanR.Isolationandlocalizationofacytosolic10Striacylglycerolbiosyntheticmultienzymecomplexfromoleaginousyeast.JBiolChem,2001,276:10290-102981[15]RaychaudhuriS,ReddyM,RajkumarNR,etal.Cytosolicironsuperoxidedismutaseisapartofthetriacylglycerolbiosyntheticcomplexinoleaginousyeast.BiochemJ,2003,372:587-5941[16]GangarA,RaychaudhuriS,RajasekharanR.Alterationinthecytosolictriacylglycerolbiosyntheticmachineryleadstodecreasedcellgrowthandtriacylglycerolsynthesisinoleaginousyeast.BiochemJ,2002,365:577-5891[17]EvansCT,ScraggAH,RatledgeC.Regulationofcitrateeffluxfrommitochondriaofoleaginousandnon2oleaginousyeastsbyadeninenucleotides.EurJBiochem,1983,132:609-6151[18]BothamPA,RatledgeC.AbiochemicalexplanationforlipidaccumulationinCandida107andotheroleaginousmicroorganisms.JGenMicrobiol,1979,114:361-3751[19]PalmieriL,PalmieriF,RunswickMJ,etal.Identificationbybacterialexpressionandfunctionalreconstitutionoftheyeastgenomicsequenceencodingthemitochondrialdicarboxylatecarrierprotein.FEBSLett,1996,399:299-3021[20]WynnJP,HamidAA,LiY,etal.BiochemicaleventsleadingtothediversionofcarbonintostoragelipidsintheoleaginousfungiMucorcircinelloidesandMortierellaalpina.Microbiology,2001,147:2857-28641[21]PapanikolaouS,SarantouS,KomaitisM,etal.RepressionofreservelipidturnoverinCunninghamellaechinulataandMortierellaisabellinacultivatedinmultiple2limitedmedia.JApplMicrobiol,2004,97:867-8751[22]AdamsIP,DackS,DickinsonFM,etal.ThedistinctivenessofATP:citratelyasefromAspergillusnidulans.BiochimBiophysActa,2002,1597:36-411[23]Ra