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RNABiosynthesis(Transcription)RNABiosynthesis（transcription）RNA的生物合成課件11．復(fù)制和轉(zhuǎn)錄過程有什么相似之處？又各有什么特點(diǎn)？
2.試述真核細(xì)胞RNA轉(zhuǎn)錄后的加工修飾1．復(fù)制和轉(zhuǎn)錄過程有什么相似之處？又各有什么特點(diǎn)？
2.2Concept
TheprocessofcopyingDNAtoRNAbyanenzymecalledRNApolymerase(RNAP)ThetransferofgeneticinformationfromDNAintoRNAtranscriptionRNADNA
ConcepttranscriptionRNA3ContentTemplatesandEnzymesTheProcessofTranscriptionPost-transcriptionalModificationContentTemplatesandEnzymes4ThedifferencebetweenreplicationandtranscriptionThedifferencebetweenreplica5SubstancesfortranscriptionSubstrates:
NTP(ATP,UTP,GTP,CTP)
Template:
DNAEnzyme:
RNApolymerase(RNA-pol)OtherproteinfactorsSubstancesfortranscriptionSu6TemplatesandEnzymesSectionISectionI7TemplatefortranscriptionStructuralgene:aDNAfragmentcodingforanyRNAorproteinproductotherthanaregulatoryelementTemplatestrand(sensestrandorWatsonstrand)
:thestrandofDNAthattheRNApolymeraseusesasatemplatetoproducecomplementarymRNACodingstrand(antisensestrandorCrickstrand)
:thestrandthatisnotusedasthetemplateithasthesamebasesequenceastheRNAtranscriptproduced(althoughwiththyminereplacedbyuracil)TemplatefortranscriptionStru85′···GCAGTACATGTC···3′3′···cgtgatgtacag···5′5′···GCAGUACAUGUC···3′N······Ala·Val·His·Val······CCodingstandTemplatestrandmRNAProteintranscriptiontranslation5′···GCAGTACATGTC···3′3′···c95335TemplatestrandCodingstrandCodingstrandTemplatestrandStructuralgeneDirectionoftranscriptionDirectionoftranscription53TemplatestrandCodingstra10Asymmetrictranscription
certainregionofonlyonestrandofDNAservesasatemplatefortranscriptionThetemplatestandisnotalwaysrestrictedtothesamesingleDNAstrandAsymmetrictranscription11RNApolymeraseProkaryoticRNApolymeraseRNApolymeraseProkaryoticRNA12coreenzymeholoenzymecoreenzymeholoenzyme13ThebindingsiteofRNApolholoenzymeonthetranscriptioninitiationregionThebindingsiteofRNApolhol14EukaryoticRNApolymeraseEukaryoticRNApolymerase15AsiteforRNApoltoidentifyandbindonthetemplateOperon:aunitoftranscriptioninprokaryotesincludingoneormorestructuralgenesandupstreamregulon
5335StructuralgeneregulonRNA-polpromoter:
thesiteonthetemplateDNAforRNApolbindingAsiteforRNApoltoidentify16RNApolymeraseprotectionassay目錄RNApolymeraseprotectionassay17TranscriptioninitiationTTGACAAACTGT-35region(Pribnowbox)TATAATPuATATTAPy-10region1-30-5010-10-40-205335Prokaryoticpromoteconservativesequencerecognitionsite
55RNApolprotectionregionStructuralgene33TranscriptioninitiationTTG18
TATAbaxCAATbaxGCbaxenhancerCis-actinelementStructuralgene-GCGC---CAAT---TATATranscriptioninitiationEukaryoticpromoteconservativesequenceTATAbaxCAATbaxGCbaxenhan19TheProcessofTranscriptionSectionIISectionII20InitiationTwoissuesrequiredtobeaddressed
thebindingofRNApolymerasetotheinitialregionofthetranscriptiontemplatepreciselyonlyonestrandofDNA,thetemplatestrand,isusedtoproducemRNAProkaryotictranscriptionInitiationTwoissuesrequired21TheDNAisunwoundandbecomessingle-strandedinthevicinityoftheinitiationsite(definedas+1).RNApolholoenzyme(2)specificallybindingtothepromoterregiononDNA1stpolyreactioncatalyzedbyRNApolandtranscriptioninitiationcomplexisformedRNApol(2)-DNA-pppGpN-OH3transcriptioninitiationcomplex5-pppG-OH+
NTP
5-pppGpN
-OH3+ppiInitiationTheDNAisunwoundandbecome22Elongationsubunitdissociatesfromtheholoenzyme，andelongationproceeds
withtheeffectofthecoreenzyme，NTPmaintainpolymerization，RNAstrandelongated(NMP)n
+
NTP(NMP)n+1
+PPiElongationsubunitdissociate23transcriptionbubble：RNA-pol（Coreenzyme）
····DA
····RNA目錄transcriptionbubble：RNA-pol（24目錄目錄2553DNAFeatherphenomenaduringprokaryotictranscriptionribosomeRNARNApol53DNAFeatherphenomenadurin26Rho(ρ)factordependenttranscriptionterminationRho-independenttranscriptionterminationTerminationTheprocessoftheRNApolymerasestoppingonDNAtemplateandthedissociationoftheRNAfromthetranscriptioncomplexClassificationRho(ρ)factordependenttrans27ATPRho(ρ)factordependenttranscriptionterminationATPRho(ρ)factordependent28TerminationindependentonRhofactorTheterminatorsequencewithintheRNAforthestopofRNApolymerase.Theterminatorsequenceusuallyformsastem-loophairpinstructureleadingtothedissociationoftheRNAPfromtheDNAtemplateTerminationindependentonRho295`UUGCAGCCUGACAAAUCAGGCUGAUGGCUGGUGACUUUUUAGUCACCAGCCUUUUU...3`5`UUGCAGCCUGACAAAUCAGGCUGAUGGCUGGUGACUUUUUAGUCACCAGCCUUUUU...3`
RNA5TTGCAGCCTGACAAATCAGGCTGATGGCTGGTGACTTTTTAGTCACCAGCCTTTTT...3DNA
UUUU...…
UUUU...…5`UUGCAGCCUGACAAAUCAGGCUGAUGGCUGGUGACUUUUUAGUCACCAGCCUUUUU...3`stem-loop/hairpinstructure5`UUGCAGCCUGACAAAUCAGGCUGAUGGC30MechanisminvolvedintheterminationbystemloopallostericchangeofRNApolyforthestopoftranscriptiondissociationoftranscriptioncomplexforthereleaseofRNA5′pppG5335RNA-polMechanisminvolvedintheterm31EukaryotictranscriptioninitiationinitiationAvarietyofupstreamcontrolelementsexistRNA-poldoesnotdirectlybindtothetemplateintheinitiationMuchmorecomplicatedthanprokaryotictranscriptionEukaryotictranscriptioniniti32TranscriptionInitiationnsiteTATAboxCAATboxGCboxenhancercis-actingelementTheupstreamregionfromthetranscriptioninitiationAATAAAExcisionandaddingtailTranscriptionTerminationsiteModificationsiteexonTranslationinitiationsiteintronOCT-1
OCT-1：ATTTGCAToctamerTranscriptionTATAboxCAATboxG33TranscriptionfactorTrans-actingfactors:TheproteinswithabilitytorecognizeandbinddirectlyorindirectlytotheupstreamcontrolelementofDNAHundredsfactorshavebeenreportedTranscriptionalfactors(TF):
Amongthetrans-actingfactors,theproteinswithabilityto
binddirectlyorindirectlytoRNApolTranscriptionfactorTrans-acti34參與RNA-polⅡ轉(zhuǎn)錄的TFⅡ
參與RNA-polⅡ轉(zhuǎn)錄的TFⅡ35Pre-initiationcomplex(PIC)Eukaryotictranscriptioninitiationrequireavarietyoftranscriptionfactor
becauseof
theindirectlybindingofEukaryoticRNA-poltoDNAPre-initiationcomplex(PIC)36POL-ⅡTFⅡFⅡAⅡBPICtranscriptioncatalyzedbyRNA-PolⅡPOL-ⅡTFⅡFⅡHⅡETBPTAFTFⅡD-ⅡA-ⅡB-DNAcomplexTATAⅡAⅡBTBPTAFTATAⅡHⅡECTD-PAfterPICassemble，CTDphosphorylationbyTFⅡH
POL-ⅡTFⅡFⅡAⅡBPICtranscription37Piecingtheory
Theeukaryotictranscriptionrequires3to5TFbindingtogetherandbecomingthecomplexfortheactivationofspecificgenetranscriptionPiecingtheoryTheeukaryotic38ElongationSimilartoprokaryotictranscriptionelongation.synchronoustranscriptionandtranslationisnotobservedbecauseofsegregationofnuclearmembraneTheforwardmovingprocessofRNA-polencounteringnucleosomeThetranslocationanddissociationofnucleosomesobservedduringthetranscriptionElongationSimilartoprokaryot39RNA-PolRNA-PolRNA-PolnucleosomeNucleosometranslocationduringelongationDirectionoftranscriptionRNA-PolRNA-PolRNA-Polnucleosom405------AAUAAA-5------AAUAAA--nuclease-GUGUGUGRNA-polAATAAAGTGTGTGModificationsitefortranscriptiontermination55333addingtailAAAAAAA······3mRNATermination——associatedwithpost-transcriptionalmodification5------AAUAAA-5------AAUAAA41Post-transcriptionalmodificationSctionIIISctionIII42ConceptIneukaryoticcells,ageneticprocessfortheconversionofaprimarytranscriptRNAtomatureRNAConcept43MajormodificationssplicingcleavagemodificationadditionMajormodificationssplicingcle44EukaryoticmRNAprocessingModificationstothe“head"(5')and“tail"(3')endsofthemRNA5capping
(m7GpppGp—)3polyadenylation
(polyAtail)EukaryoticmRNAprocessingModi455
capstructure5capstructure465pppGp…5GpppGp…pppGppiGMPtransferase5
m7GpppGp…methyltransferase
SAMTheformationof5
cap
5ppGp…phosphatasePi5pppGp…5GpppGp…pppGppiGMP47mRNAsplicinghetero-nuclearRNAa(hnRNA):theprimarymRNAwithinnucleussnRNA:aclassofsmallRNAmoleculeswithinthenucleus
ProteinswithinNuclear
smallnuclearribonucleoproteins
（splicesome）snRNAmRNAsplicinghetero-nuclearR48astrandofDNAcontainingbothintronsandexons.Mostadvancedeukaryoteshaveinterruptedgenesandintronsarelongerthanexons.TheresultinggeneismuchlongerthanitscodingregionInterruptedgene
CABDCodingregionA、B、C、DNon-codingregionastrandofDNAcontainingbot49ExonandintronexontheregionsofagenethatarerepresentedinmatureformofthemessengerRNA(mRNA)intronThesectionsofagenethatarepresentinprecursormRNA(pre-mRNA)andremovedbyaprocesscalledsplicingduringtheprocessingtomatureRNAExonandintronexon50ChickenOvalbuminGene
hnRNAModificationstothe"front"and"back"endshnRNAsplicingMaturemRNAChickenOvalbuminGeneanditsTranscriptionandPost-transcriptionalModification
目錄ChickenOvalbuminGenehnRNAMo51Electron-microscopicpictureofthematuremRNAandDNAhybridizationofChickenOvalbuminDNAmRNA目錄Electron-microscopicpictureo52Classificationofintrons
Fourclasses:accordingtogenetypesandsplicingmethodsI：mainlyfoundinmitochondria,chloroplast,andcertainlow-leveleukaryoticrRNAgenesII：foundinmitochondria,chloroplast,inmRNAgenesIII：frequentlyfoundinthesplicingafterloopformation，inmostmRNAgenesIV：tRNAgenesandprimarytranscriptionproductionintron，splicingrequiringenzymeandATPClassificationofintronsFour53mRNAsplicingtoremovetheintronsofhnRNAandligatetheexonsSplicesomeisformedbyligationofsnRNPandhnRNA目錄①mRNAsplicingtoremovethein54②③UACUACA-AGUGU4U5U6E1E2U1U2UACUACA-AGUGU6E1E2U1、U4、U5②③UACUACA-AGUGU4U5U6E1E2U1U255pG-OH(ppG-OH,pppG-OH)U-OHGpUpGpA1STtransesterificationreaction
2nd
transesterificationreaction
UpAGpUExon1IntronExon2G-OHUpUpGpA剪接過程的二次轉(zhuǎn)酯反應(yīng)
pG-OHU-OHGpUpGpA1STtransester56DifferentialRNAprocessing:
RNAeditingshowsthatthecodingsequenceofgeneshasavarietyofdifferentialfunctionafterpost-transcriptionprocessingmRNAeditingHumanapoBgene
mRNA（14500Nucleotides）liverapoB100（MW500000）intestinalcellsapoB48（MW240000）mRNAeditingDifferentialRNAprocessing:57Post-transcriptionalprocessoftRNAtRNAprecursorRNApolⅢTGGCNNAGTGCGGTTCGANNCCDNA目錄Post-transcriptionalprocesso58RNAaseP、endonuclease目錄RNAaseP、endonuclease目錄59tRNAnucleotidetransferase
、ligaseATPADP目錄tRNAnucleotidetransferase、l60BasemodificationsReducedreactione.g.：UDHUTranslocationwithinnucleotidese.g.：UψDeaminationreaction
e.g.：AIe.g.：AAmMethylation（1）（1）（3）（2）（4）目錄BasemodificationsReducedreac61transcription45S-rRNAsplice18S-rRNA5.8Sand28S-rRNArDNAintronintron28S5.8S18SPost-transcriptionalprocessofrRNAtranscription45S-rRNAsplice162anRNAmoleculethatcatalyzesachemicalreactionRibozymeanRNAmoleculethatcatalyzes63SecondarystructureofTetrahymena
rRNAintronSelf-splicingrRNAofTetrahymena5′-terminalsequenceSecondarystructureofTetrahy64hammerheadstructure——ThemostsimplesecondarystructureofribozymesubstrateAround60nucleotidesWithcatalyticpartandsubstrateinthesamemolecularhammerheadstructureconsistedofcatalyticpartandsubstrateApartfromrRNA，theprocessoftRNA、mRNAalsoexistsSelf-splicinghammerheadstructure——Themo65SignificanceofribozymeresearchTheimportantsupplementarytothecentraldogmaThechallengetotraditionalenzymologyDesignaribozymeaccordingtothenativeribozymestructureSignificanceofribozymeresea66Thedesignedribozyme
SyntheticnucleicacidsshownwiththicklinesNativenucleicacidsshownwiththinlinesconsensussequenceshownwithSplicedsiteshownbyarrow目錄ThedesignedribozymeSyntheti67AppendixAppendix68GOH3′G5′OH5′3′GOH414G3995′3′3955′3′E1E2ISelf-splicingRNAofTetrahymena5′3′L19RNAFragmentwithcatalyticactivityGOH3′G5′OH5′3′GOH414G3995′3′369RNABiosynthesis(Transcription)RNABiosynthesis（transcription）RNA的生物合成課件701．復(fù)制和轉(zhuǎn)錄過程有什么相似之處？又各有什么特點(diǎn)？
2.試述真核細(xì)胞RNA轉(zhuǎn)錄后的加工修飾1．復(fù)制和轉(zhuǎn)錄過程有什么相似之處？又各有什么特點(diǎn)？
2.71Concept
TheprocessofcopyingDNAtoRNAbyanenzymecalledRNApolymerase(RNAP)ThetransferofgeneticinformationfromDNAintoRNAtranscriptionRNADNA
ConcepttranscriptionRNA72ContentTemplatesandEnzymesTheProcessofTranscriptionPost-transcriptionalModificationContentTemplatesandEnzymes73ThedifferencebetweenreplicationandtranscriptionThedifferencebetweenreplica74SubstancesfortranscriptionSubstrates:
NTP(ATP,UTP,GTP,CTP)
Template:
DNAEnzyme:
RNApolymerase(RNA-pol)OtherproteinfactorsSubstancesfortranscriptionSu75TemplatesandEnzymesSectionISectionI76TemplatefortranscriptionStructuralgene:aDNAfragmentcodingforanyRNAorproteinproductotherthanaregulatoryelementTemplatestrand(sensestrandorWatsonstrand)
:thestrandofDNAthattheRNApolymeraseusesasatemplatetoproducecomplementarymRNACodingstrand(antisensestrandorCrickstrand)
:thestrandthatisnotusedasthetemplateithasthesamebasesequenceastheRNAtranscriptproduced(althoughwiththyminereplacedbyuracil)TemplatefortranscriptionStru775′···GCAGTACATGTC···3′3′···cgtgatgtacag···5′5′···GCAGUACAUGUC···3′N······Ala·Val·His·Val······CCodingstandTemplatestrandmRNAProteintranscriptiontranslation5′···GCAGTACATGTC···3′3′···c785335TemplatestrandCodingstrandCodingstrandTemplatestrandStructuralgeneDirectionoftranscriptionDirectionoftranscription53TemplatestrandCodingstra79Asymmetrictranscription
certainregionofonlyonestrandofDNAservesasatemplatefortranscriptionThetemplatestandisnotalwaysrestrictedtothesamesingleDNAstrandAsymmetrictranscription80RNApolymeraseProkaryoticRNApolymeraseRNApolymeraseProkaryoticRNA81coreenzymeholoenzymecoreenzymeholoenzyme82ThebindingsiteofRNApolholoenzymeonthetranscriptioninitiationregionThebindingsiteofRNApolhol83EukaryoticRNApolymeraseEukaryoticRNApolymerase84AsiteforRNApoltoidentifyandbindonthetemplateOperon:aunitoftranscriptioninprokaryotesincludingoneormorestructuralgenesandupstreamregulon
5335StructuralgeneregulonRNA-polpromoter:
thesiteonthetemplateDNAforRNApolbindingAsiteforRNApoltoidentify85RNApolymeraseprotectionassay目錄RNApolymeraseprotectionassay86TranscriptioninitiationTTGACAAACTGT-35region(Pribnowbox)TATAATPuATATTAPy-10region1-30-5010-10-40-205335Prokaryoticpromoteconservativesequencerecognitionsite
55RNApolprotectionregionStructuralgene33TranscriptioninitiationTTG87
TATAbaxCAATbaxGCbaxenhancerCis-actinelementStructuralgene-GCGC---CAAT---TATATranscriptioninitiationEukaryoticpromoteconservativesequenceTATAbaxCAATbaxGCbaxenhan88TheProcessofTranscriptionSectionIISectionII89InitiationTwoissuesrequiredtobeaddressed
thebindingofRNApolymerasetotheinitialregionofthetranscriptiontemplatepreciselyonlyonestrandofDNA,thetemplatestrand,isusedtoproducemRNAProkaryotictranscriptionInitiationTwoissuesrequired90TheDNAisunwoundandbecomessingle-strandedinthevicinityoftheinitiationsite(definedas+1).RNApolholoenzyme(2)specificallybindingtothepromoterregiononDNA1stpolyreactioncatalyzedbyRNApolandtranscriptioninitiationcomplexisformedRNApol(2)-DNA-pppGpN-OH3transcriptioninitiationcomplex5-pppG-OH+
NTP
5-pppGpN
-OH3+ppiInitiationTheDNAisunwoundandbecome91Elongationsubunitdissociatesfromtheholoenzyme，andelongationproceeds
withtheeffectofthecoreenzyme，NTPmaintainpolymerization，RNAstrandelongated(NMP)n
+
NTP(NMP)n+1
+PPiElongationsubunitdissociate92transcriptionbubble：RNA-pol（Coreenzyme）
····DA
····RNA目錄transcriptionbubble：RNA-pol（93目錄目錄9453DNAFeatherphenomenaduringprokaryotictranscriptionribosomeRNARNApol53DNAFeatherphenomenadurin95Rho(ρ)factordependenttranscriptionterminationRho-independenttranscriptionterminationTerminationTheprocessoftheRNApolymerasestoppingonDNAtemplateandthedissociationoftheRNAfromthetranscriptioncomplexClassificationRho(ρ)factordependenttrans96ATPRho(ρ)factordependenttranscriptionterminationATPRho(ρ)factordependent97TerminationindependentonRhofactorTheterminatorsequencewithintheRNAforthestopofRNApolymerase.Theterminatorsequenceusuallyformsastem-loophairpinstructureleadingtothedissociationoftheRNAPfromtheDNAtemplateTerminationindependentonRho985`UUGCAGCCUGACAAAUCAGGCUGAUGGCUGGUGACUUUUUAGUCACCAGCCUUUUU...3`5`UUGCAGCCUGACAAAUCAGGCUGAUGGCUGGUGACUUUUUAGUCACCAGCCUUUUU...3`
RNA5TTGCAGCCTGACAAATCAGGCTGATGGCTGGTGACTTTTTAGTCACCAGCCTTTTT...3DNA
UUUU...…
UUUU...…5`UUGCAGCCUGACAAAUCAGGCUGAUGGCUGGUGACUUUUUAGUCACCAGCCUUUUU...3`stem-loop/hairpinstructure5`UUGCAGCCUGACAAAUCAGGCUGAUGGC99MechanisminvolvedintheterminationbystemloopallostericchangeofRNApolyforthestopoftranscriptiondissociationoftranscriptioncomplexforthereleaseofRNA5′pppG5335RNA-polMechanisminvolvedintheterm100EukaryotictranscriptioninitiationinitiationAvarietyofupstreamcontrolelementsexistRNA-poldoesnotdirectlybindtothetemplateintheinitiationMuchmorecomplicatedthanprokaryotictranscriptionEukaryotictranscriptioniniti101TranscriptionInitiationnsiteTATAboxCAATboxGCboxenhancercis-actingelementTheupstreamregionfromthetranscriptioninitiationAATAAAExcisionandaddingtailTranscriptionTerminationsiteModificationsiteexonTranslationinitiationsiteintronOCT-1
OCT-1：ATTTGCAToctamerTranscriptionTATAboxCAATboxG102TranscriptionfactorTrans-actingfactors:TheproteinswithabilitytorecognizeandbinddirectlyorindirectlytotheupstreamcontrolelementofDNAHundredsfactorshavebeenreportedTranscriptionalfactors(TF):
Amongthetrans-actingfactors,theproteinswithabilityto
binddirectlyorindirectlytoRNApolTranscriptionfactorTrans-acti103參與RNA-polⅡ轉(zhuǎn)錄的TFⅡ
參與RNA-polⅡ轉(zhuǎn)錄的TFⅡ104Pre-initiationcomplex(PIC)Eukaryotictranscriptioninitiationrequireavarietyoftranscriptionfactor
becauseof
theindirectlybindingofEukaryoticRNA-poltoDNAPre-initiationcomplex(PIC)105POL-ⅡTFⅡFⅡAⅡBPICtranscriptioncatalyzedbyRNA-PolⅡPOL-ⅡTFⅡFⅡHⅡETBPTAFTFⅡD-ⅡA-ⅡB-DNAcomplexTATAⅡAⅡBTBPTAFTATAⅡHⅡECTD-PAfterPICassemble，CTDphosphorylationbyTFⅡH
POL-ⅡTFⅡFⅡAⅡBPICtranscription106Piecingtheory
Theeukaryotictranscriptionrequires3to5TFbindingtogetherandbecomingthecomplexfortheactivationofspecificgenetranscriptionPiecingtheoryTheeukaryotic107ElongationSimilartoprokaryotictranscriptionelongation.synchronoustranscriptionandtranslationisnotobservedbecauseofsegregationofnuclearmembraneTheforwardmovingprocessofRNA-polencounteringnucleosomeThetranslocationanddissociationofnucleosomesobservedduringthetranscriptionElongationSimilartoprokaryot108RNA-PolRNA-PolRNA-PolnucleosomeNucleosometranslocationduringelongationDirectionoftranscriptionRNA-PolRNA-PolRNA-Polnucleosom1095------AAUAAA-5------AAUAAA--nuclease-GUGUGUGRNA-polAATAAAGTGTGTGModificationsitefortranscriptiontermination55333addingtailAAAAAAA······3mRNATermination——associatedwithpost-transcriptionalmodification5------AAUAAA-5------AAUAAA110Post-transcriptionalmodificationSctionIIISctionIII111ConceptIneukaryoticcells,ageneticprocessfortheconversionofaprimarytranscriptRNAtomatureRNAConcept112MajormodificationssplicingcleavagemodificationadditionMajormodificationssplicingcle113EukaryoticmRNAprocessingModificationstothe“head"(5')and“tail"(3')endsofthemRNA5capping
(m7GpppGp—)3polyadenylation
(polyAtail)EukaryoticmRNAprocessingModi1145
capstructure5capstructure1155pppGp…5GpppGp…pppGppiGMPtransferase5
m7GpppGp…methyltransferase
SAMTheformationof5
cap
5ppGp…phosphatasePi5pppGp…5GpppGp…pppGppiGMP116mRNAsplicinghetero-nuclearRNAa(hnRNA):theprimarymRNAwithinnucleussnRNA:aclassofsmallRNAmoleculeswithinthenucleus
ProteinswithinNuclear
smallnuclearribonucleoproteins
（splicesome）snRNAmRNAsplicinghetero-nuclearR117astrandofDNAcontainingbothintronsandexons.Mostadvancedeukaryoteshaveinterruptedgenesandintronsarelongerthanexons.TheresultinggeneismuchlongerthanitscodingregionInterruptedgene
CABDCodingregionA、B、C、DNon-codingregionastrandofDNAcontainingbot118ExonandintronexontheregionsofagenethatarerepresentedinmatureformofthemessengerRNA(mRNA)intronThesectionsofagenethatarepresentinprecursormRNA(pre-mRNA)andremovedbyaprocesscalledsplicingduringtheprocessingtomatureRNAExonandintronexon119ChickenOvalbuminGene
hnRNAModificationstothe"front"and"back"endshnRNAsplicingMaturemRNAChickenOvalbuminGeneanditsTranscriptionandPost-transcriptionalModification
目錄ChickenOvalbuminGenehnRNAMo120Electron-microscopicpictureofthematuremRNAandDNAhybridizationofChickenOvalbuminDNAmRNA目錄Electron-microscopicpictureo121Classificationofintrons
Fourclasses:accordingtogenetypesandsplicingmethodsI：mainlyfoundinmitochondria,chloroplast,andcertainlow-leveleukaryoticrRNAgenesII：foundinmitochondria,chloroplast,inmRNAgenesIII：frequentlyfoundinthesplicingafterloopformation，inmostmRNAgenesIV：tRNAgenesandprimarytranscriptionproductionintron，splicingrequiringenzymeandATPClassificationofintronsFour122mRNAsplicingtoremovethein
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