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1、Chapter6MobileGeneticElementsIntroductionMuchofthediscussionthroughoutthisbookdealswiththefunctionsofthegeneproductsexpressedfromproteincodinggenesandRNAgenes.However,noncodingrepetitiousDNA,consistingofsimple-sequenceDNAandmoderatelyrepeatedDNA,constitu
2、tesasignificantfractionofthegenomicDNAinhighereukaryotes.Inthissection,wefocusonmoderatelyrepeatedDNAsequences,ormobileDNAelements,whichareinterspersedthroughoutthegenomesofhigherplantsandanimals.AlthoughmobileDNAelements,rangingfromhundredstoafewthousan
3、dbasepairsinlength,originallywerediscoveredineukaryotes,theyalsoarefoundinprokaryotes.Theprocessbywhichthesesequencesarecopiedandinsertedintoanewsiteinthegenomeiscalledtransposition.MobileDNAelements(orsimplymobileelements)areessentiallymolecularparasite
4、s,whichappeartohavenospecificfunctioninthebiologyoftheirhostorganisms,butexistonlytomaintainthemselves.Forthisreason,FrancisCrickreferredtothesesequencesas“selfishDNA.”ThetranspositionofmobileDNAelementsisbelievedtohaveresultedintheirslowaccumulationineu
5、karyoticgenomesoverevolutionarytime.TheseelementsalsoarelostataveryslowratebydeletionofsegmentsofDNAcontainingthemandbyaccumulationofmutationsuntiltheycannolongerberecognizedtoberelatedtotheoriginalmobileDNAelement.Sincemobileelementsareeliminatedfromeuk
6、aryoticgenomessoslowly,theyhaveaccumulatedtothepointwheretheynowconstituteasignificantportionofthegenomesofmanyeukaryotes.Asresearchonmobileelementsprogressed,theywerefoundtofallintotwocategories:(1)thosethattransposedirectlyasDNAand(2)thosethattranspose
7、viaanRNAintermediatetranscribedfromthemobileelementbyanRNApolymeraseandthenconvertedbackintodouble-strandedDNAbyareversetranscriptase.MobileelementsthattransposethroughaDNAintermediatearegenerallyreferredtoastransposons.(Asdiscussedbelow,thistermhasamore
8、specificmeaninginreferencetobacterialmobileelements.)MobileelementsthattransposetonewsitesinthegenomeviaanRNAintermediatearecalledretrotransposonsbecausetheirmovementisanalogoustotheinfectiousprocessofretroviruses.Indeed,r