植物萜类化合物的天然合成途径及其相关合酶

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43420078779*310029PlantTerpenoidNaturalMetabolismPathwaysandTheirSynthasesZHANGChang-Bo,SUNHong-Xia,GONGZhong-Jun,ZHUZeng-Rong*StateKeylaboratoryofRiceBiology,InstituteofInsectSciences,ZhejiangUniversity,Hangzhou310029,China2007-05-082007-07-09(3037093530571259)(2004E60055)*(E-mailzrzhu@zju.edu.cnTel0571-86971623)(terpenoids)(isopreneC5)(C10)(C15)(C20)(C30)(1)(2)-6-(3)(4)(Taxusbrevifolia)β-1(mevalonatepathwayMVA)(Chappell1995)-5-(1-deoxy-D-xylulose-5-phos-phatepathwayDXP)(Rohmer1993)4-(methylerythritol4-phosphatepathwayMEP)3(isopentenylpyrophosphateIPPC5)(dimethylallylpyrophosphateDMAPPC5)MVA(NewmanChappell1999)2CoA(acetyl-CoA)CoA(acetoacetyl-CoAthiolaseATOT)CoA(acetoacetyl-CoA)CoA(hydroxymethylglutaryl-CoAsynthaseHMGS)3--3-CoA(3-hydroxy-3-methylglutarylcoenzymeAHMG-CoA)CoA(hydroxymethylglutaryl-CoAreductaseHMGR)2NADPHMVAMVAMVA(mevalonatekinaseMK)-5-(mevalonate-434200787805-phosphateMVAP)MVA(phosphomevalonatekinasePMK)MVA-5-(mevalonate-5-diphosphateMVAPP)MVA(mevalonatepyrophosphatedecarboxylaseMPD)IPP(1)DXP-3--5-(1-deoxy-D-xylulose5-phosphatesynthaseDXS)1---5-(1-deoxy-D-xyulose5-phosphateDXP)(1-deoxy-D-lxylulose5-phosphatereductoisomeraseDXR)DXP2C--D--4-(2C-methyl-D-erythritol4-phosphateMEP)(LangeCroteau1999)MEP4--2-C--D-(4-diphosphocytidyl-2-C-methyl-D-erythritolsynthaseCMS)(Rohdich2001)4--2-C-(4-diphosphocytidyl-2-C-methylerythritolCDP-ME)4--2-C-(4-diphosphocytidyl-2-C-methyl-D-erythritolkinaseCMK)4--2-C--D--2-(4-diphosphocytidyl-2-C-me-thyl-D-erythritol2-phosphateCDP-ME2P)2--2,4-(2-C-methyl-D-erythritol2,4-cyclodiphosphatesynthaseMCS)2C--D-2,4-(2C-methyl-D-erythritol2,4-cyclodiphosphateME-2,4cPP)1--2--2-(E)-4-[1-hydroxy-2-methyl-2-(E)-butenyl4-diphosphatesynthaseHDS]1--2--2-(E)-4-[1-hydroxy-2-me-thyl-2-(E)-butenyl4-diphosphateHMBPP]IPP(isopentenylmonophosphatekinaseIPK)(Schwender2001)(1)IPPIPP(Laule2003)IPPMVADXP(Rohdich2001Rodríguez-ConcepciónBoronat2002)IPP(isopentenyldiphosphateisomeraseIDI)DMAPPIPPDMAPP1(geranyldiphosphatesynthaseGPPS)C10(geranyldiphosphateGPP)2IPP1DMAPP(farnesyldiphosphatesynthaseFPPS)C15(farnesyldiphosphateFPP)3IPP1DMAPP(geranylgeranyldiphosphatesynthaseGGPPS)C20(geranylgeranyldiphosphateGGPP)3GPPFPPGGPP(terpenesynthase,TPS)GPP(monoter-penesynthase)(monoterpeneC10)FPP(sesquiterpenesynthase)(sesquiterpeneC15)GGPP(diterpenesynthase)(diterpeneC20)TPS22.1CoA(HMGR)HMGRMVAHMG-CoAMVAMVA11986HMGR180kDa445kDa5.9(Bach1986)HMGR3(hmgr-1hmgr-434200787811(HsiehGoodman2005WithersKeasling2007)ATOTCoAHMGSCoAHMGRCoAMKMVAPMKMVAMPDMVADXS-5-DXRCMS4--2-C--D-CMK4--2-C-MCS2--2,4-HDS1--2--2-(E)-4-IPKIDIFPPSGPPSGGPPSTPS434200787822hmgr-3)Enfissi(2005)(Arabidopsisthaliana)HMGRhmgr-1(CaMV)35SMVA2.4Jiang(2006)(Eucommiaulmoides)HMGREuHMGR2281bp1770bp5902(housekeepinggene)HMGRJRY2394EuHMGRMVA2.2-5-(DXS)DXSDXP-3-DXPDXSDXP(Estévez2001)2006DXS(Xiang2007)3(IIIIII)III(thiaminepyrophosphateTPP)(thiazoliumring)2CGong(2006)(Ginkgobiloba)DXS2154bpDXSTPP(transketolase)DXS1DXSDXSWalter(2002)(Medicagotruncatula)3DXScDNA2DXSEST()3DXS1DXS(Estévez2000Rodríguez-ConcepciónBoronat2002)Kim(2005)3DXS(dxs1dxs2dxs3)Munoz-Bertomeu(2006)DXS(Lavandulalatifolia)Morris(2006)DXS3824956~70d2.3(DXR)DXRDXPMEPDXPDXSDXR(fosmidomycin)DXRKuntz(2005)2DXR4-(hydroxyamino)-4-oxobutylphosphonicacid4-[hydroxy(methyl)amino]-4-oxobutylphosphonicacidDXRDXRDXRRohdich(2006)DXRN(Escherichiacoli)DXPMEP5.6µmol·min-1·mg-1()Mn2+Mg2+pH8.0NADPHNADPHNADH14%2.42.4.1(GPPS)GPPSIPPDMAPPGPPGPPSGPPSGGPPSFPPS2004Tholl(Antirrhimummajus)(Clarkiabreweri)GPPSGPPSBurke(1999)(Menthahaplocalyx)GPPS53%45%GPPSGPPS2.4.2(FPPS)FPPS243420078783IPP1DMAPPGPPFPP(Cunillera1997)(Sanmiya1999)(OliverKrisans2000)FPPS7(Koyama1993)2IIDDXX(XX)D(DX)FARM(firstAsp-richmotif)E-2VIDDXXDSARM(secondAsp-richmotif)Liu(1999)(Gossypiumarboreum)FPPSgarfps(Verticilliumdahliae)FPPSFPPS(Cunillera1997)(Sanmiya1999)(Artemisiatridentata)(Hemmerlin2003)2FPPSHemmerlin(2003)cDNA3FDS-1FDS-2FDS-589%2.4.3(GGPPS)GGPPSIPPDMAPPGPPFPPGGPPGGPPSFARMKojima(2000)PCR(Scopariadulcis)(Crotonsublyratus)2GGPPSGGPPSFPPIPPGGPPGGPPS60%~70%FARMFARMFARM2DMAPPFPPGGPPFARMTakaya(2003)(Heveabrasiliensis)cDNAGGPPSGGPP40pH8.52.5(TPS)TPSGPPFPPGGPPDDXXD(Lesburg1997)(Bohlmann1998)2.5.1(K+)Mn2+Fe2+pH(Bohlmann1997)Mn2+Mg2+pH(AlonsoCroteau1991)Yang(2005)(Cinnamomumtenuipilum)CtGESGPPMg2+Mn2+CtGESCtGES2.5.2Picaud(2005)(Artemisiaannua)(E)-β-[(E)-beta-farnesenesynthase]1746bp574(66.9kDa)30%~50%FPPβ-pH6.5Mg2+Mn2+Co2+Green(2007)(Malus×domestica)α-GPPIPPα-43420078784FPP1%GPPK+2.5.3GGPP(Hamano2002)DDXXDRoBohlmann(2006)(Pinustaeda)PtTPS-LASGGPP(levopimaradiene)(abietadiene)(neoabietadiene)palu-stradiene3(1)(Tholl2006)(2)(Lange2000)(3)40~100mg(Sato2001)(Wu2006)3(2005)(1)cDNAPCRcDNA(Kojima2000Prosser2006)(2)cDNA(3)“”(Medicagotruncatula)(Bede2006)DNA,(2005)..,2743420078785(1):1~10AlonsoWR,CroteauR(1991).Purificationandcharacterizationofthemonoterpenecyclasegammar-terpinenesynthasefromThymusvulgaris.ArchBiochemBiophys,286:511~517BachTJ,RogersDH,RudneyH(1986).Detergent-solubilization,purification,andcharacterizationofmembrane-bound3-hydroxy-3-methylglutaryl-coe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