RNA病毒基因组和转录复制多样性的分子基础

RNA刘青珍李凌云齐义鹏*杨复华(,430072):RNADNA,RNA:Myxobacteria(Retron):DNARNA/DNARNARNARNA/RNA3poly(A)tRNA,,5RNARNARNARNARNA,RNA,5UTR3UTRRNA:RNA,,,:Q939.4,Q753,Q522+.4:B:1005-0094(2001)03-0294-07MolecularmechanismfortranscriptionandreplicationofRNAvirusLIUQingZhen,LILingYun,QIYiPeng*,YANGFuHuaInstituteofVirology,WuhanUniversity,Wuhan430072Abstract:RNAvirusesaremuchmorethanDNAvirusesinquantityandspeciesinnature.Itcanbedividedintomanytypesofvirusesaccordingtogenomictype.Recently,manyresearchersindicatethattheretronexistinginMyxobacteria,whichcarriesonlyonereversetranscriptasegene,isthepossibleancestorforallkindsofviruses,withthefollowingevolutionmodel:RetronRetroposonRetransposonRetrovirusPararetrovirusDNAviruses.Itisverydifferentbetweentranscription/replicationofRNAvirusandDNAvirusgenomesinmanyaspects.RNAdependentRNApolymeraseisthemajorcatalystfortranscription/replication.InitiationsitesforRNAvirustranscription/replicationarespecificstructures[poly(A)ortRNA]at3terminalregion.Internalstopresultsintranscriptionbutreadingthroughto5endresultinginreplication.RNAvirusesusethefollowingfourkindsoftemplatesintranscription/replication:(+)ssRNAtemplate,(-)ssRNAtemplate,(+)aRNAtemplate,(-)aRNAtemplate.ThemechanismofselectiveregulationonRNAtemplatesisverycomplexandtheacknowledgementsoftheregulationmechanismarequitefewuptonow.SelectingtemplateandthebindingoftranscriptionfactorswithRNApoltoformrepliconaretwomajorregulatorymeans.Besidespromoterandtranscriptionfactors,5UTRand3UTRalsoregulatethetranscriptionofRNAviruses.Keywords:RNAvirus,replication,transcription,diversity,RNA,WalterKeller(1999)Science!TheRNAWorld∀(),!IntheBeginning,ThereWasRNA∀:RNARNA,RNA(vi#:(39970032):2000-09-11;:2001-01-15:,,1965,*Authorforcorrespondence:Email:[e-mail]2001,9(3):294300#BiodiversityScienceroid)DNARNA,,RNARNA,,RNA,;,RNA,,RNA,,,DNA,RNA(Liuetal.,2000)RNA,,1RNA,DNARNA,DNAdsDNA,,RNA(Bercoff,1987;,1992),:1)RNA[(+)ssRNA]:RNA,,mRNA,(Calicivirus)RNA(Picornavirus)(Togavirus)(Flavivirus)(Coronavirus)(Nodavirus)2)RNA[(-)ssRNA]:RNA,,mRNA,,mRNARNA:(Paramyxovirus)(Orthomyxovirus)(Rhabdovirus)(Bunyavirus)(Arenavirus),3)RNA(dsRNA):,(Reovirus)RNA(Cystovirs)HypovirusTotivirusBirnavirusPartitivirus,(+)/(-)RNA,10~12dsRNA(+),(+)/(-)4)RNA[(+)ssRNA]:RNA,(+)ssRNA,dimer(pol),(+)ssRNAPOL,RNAdsDNAdsDNADNA,DNA,mRNA(Retrovirus)(Oncovirus)(HIV)(Spumavirus)5)RNA[(+)ssRNA]:RNARNARNA,RNA,RNARNA(TMV)RNA(TRSV)(TRV);(CMV)(BMV)6)(Viroid,ssRNA):,,ssRNA,300nt,7)RNA(SatelliteRNA):ssRNA,,,,(Virosoid),RNA,RNARNA,DNARNA?10RNA(Lanpsonetal.,1991;Yechiel,1996),myxobacteria3:RNA295(Retron),RNADNAmsDNA,Retrongagint(Retroposon)intLTR(Retrotransposon)env(,1997),,RNA,(Pararetrovirus)DNA,RNADNARNA(Baron&Barrett,1995;Stecetal.,1991),RetronRNA/DNA(,1997;Stecetal.,1991;Pochetal.,1990)2RNA/RNA/,,,,,,(DNA)DNARNARNA/,(RNAPOLmotif),DNAPochetal.(1990),RNARNA(POL)RNA/RNAori,3poly(A)tRNA,,5,RNA(Howleyetal.,1999;Chandrikaetal.,1995)RNA/,DNAdsDNA,RNARNARNARNADNA/1(Liuetal.,2000)RNA,RNA1RNADNA/Table1TranscriptionalandreplicatingcharactersofRNAvirusandDNAvirusRNARNAvirusesDNADNAviruses/Relationshipbetweentranscriptionandreplication,,Transcriptioniscoupledwithreplication.Transcriptionisterminatedininsidesites,andreplicationisstoppedattheendTranscriptionandreplicationareindependentPolymeraseRNARNAOnlyonekindofRNAdependentRNApolymeraseisusedintranscriptionandreplicationDNA,RNAPOL;DNAPOLDNAistemplate.RNApolymeraseisusedfortranscription,DNApolymeraseforreplicationProducts:mRNA:+/-aRNA;+/-vRNATranscription:mRNAReplication:+/-aRNA;+/-vRNA:mRNA*:dsDNATranscription:mRNA*Replication:dsDNATemplates4:+/-vRNA;+/-aRNA+/-vRNA;+/-aRNA1:dsDNAdsDNAReplicationorigins3Poly(A)tRNAInitiatefrom3Poly(A)andtRNAstructureorotherstructuresori,oriOnlyoneisselectedforreplicationfrommanyoriginsOtherenzymesDon∃tneedhelicaseandligaseNeedhelicase,topoisomeraseandligaseReplicatingforms,Linearreplication,noOkazakifragments,Partialrollingcirclereplication,haveOkazakifragmentsReplicatingregulationThemajoristemplateselectiveregulationandantiterminationregulationPositiveandnegativeregulationofreplicationunits*DNAmRNAGeminivirus.SomekindsofDNAvirusdon∃ttranscribeintomRNA,suchasGeminivirus.296BiodiversityScience9,RNARNAHDV(HepatitisDeltaVirus)ss(+)RNA,RNA,RNARNA(Ribozyme)5-OH2,3-RNARibozyme(Rz),Rz,Rz,HDVRz,VSRz,3RNA/RNA/RNA,(Replicon),,Chandrikaetal.(1995)PRNAPOLN348~379aaLP,,,Howleyetal.(1999),(BMV);(tubulin)(Moyeretal.,1990);Q(+ssRNA)30sRibosomeS1EFTu/EFTs(Bercoff,1987),(ssRNA)PB1,PB2AP,PB1PB2RNARNA3/Q3RNA(3CCAOH)Litvaketal.(1994),tRNALys33HIV3(PBS)Kohlstaedtetal.(1992)tRNA-HIVI(RT)RTRNAdsDNA,HIVRT2(p66,p51)HIV-1RNA,H(RNaseH);tRNA,HIVPBSRT,tRNA,3;HIVp66RNAPOL(1)BanejeeBarik(1992),tRNA(2)1HIV1tRNA(Litvaketal.,1994)Fig.1HIV1replisomeandtRNAprimers(AdoptedfromLitvaketal.,1994)2tRNATable2tRNAprimersusedbyvariousRetrovirusRetrovirustRNAtRNAprimer(ASLV)Aviansarcomaleukosisvirus(ASLV)tRNATrp(AMV)Avianmyeloblastosisvirus(AMV)tRNATrp(MLV)Murineleukemiavirus(MLV)tRNAPro(SNV)Spleennecrosisvirus(SNV)tRNAProT(%,&)(HTLV)HumanTcellleukemiavirus(%,&)(HTLV)tRNAPro(BL