Repeated reunions and splits feature the highly dynamic evolution of 5S and 35S ribosomal RNA genes (rDNA) in the Asteraceae family 英文参考文献.docVIP

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Repeated reunions and splits feature the highly dynamic evolution of 5S and 35S ribosomal RNA genes (rDNA) in the Asteraceae family 英文参考文献.doc

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Repeated reunions and splits feature the highly dynamic evolution of 5S and 35S ribosomal RNA genes (rDNA) in the Asteraceae family 英文参考文献

Garciaetal.BMCPlantBiology2010,10:176 /1471-2229/10/176 RESEARCH ARTICLE OpenAccess Repeatedreunionsandsplitsfeaturethehighly dynamicevolutionof5Sand35SribosomalRNA genes(rDNA)intheAsteraceaefamily SòniaGarcia1,2,JoséLPanero3,JiriSiroky2,AlesKovarik2* Abstract Background:InfloweringplantsandanimalsthemostcommonribosomalRNAgenes(rDNA)organisationisthat inwhich35S(encoding18S-5.8S-26SrRNA)and5Sgenesarephysicallyseparatedoccupyingdifferent chromosomalloci.However,recentobservationsestablishedthatbothgeneshavebeenunifiedtoasingle35S-5S unitinthegenusArtemisia(Asteraceae),agenomicarrangementtypicalofprimitiveeukaryotessuchasyeast, amongothers.Hereweaimtorevealtheorigin,distributionandmechanismsleadingtothelinkedorganisationof rDNAintheAsteraceaebyanalysingunitstructure(PCR,Southernblot,sequencing),genecopynumber (quantitativePCR)andchromosomalposition(FISH)of5Sand35SrRNAgenesin~200speciesrepresentingthe familydiversityandothercloselyrelatedgroups. Results:DominantlinkedrDNAgenotypewasfoundwithinthreelargegroupsinsubfamilyAsteroideae:tribe Anthemideae(93%ofthestudiedcases),tribeGnaphalieae(100%)andinthe“Heliantheaealliance”(23%).The remainingfivetribesoftheAsteroideaedisplayedcanonicalnonlinkedarrangementofrDNA,asdidtheother groupsintheAsteraceae.Nevertheless,lowcopylinkedgeneswereidentifiedamongseveralspeciesthat amplifiedunlinkedunits.Theconservedpositionoffunctional5Sinsertionsdownstreamfromthe26Sgene suggestsaunique,perhapsretrotransposon-mediatedintegrationeventatthebaseofsubfamilyAsteroideae. Furtherevolutionlikelyinvolveddivergenceof26S-5Sintergenicspacers,amplificationandhomogenisationof unitsacrossthechromosomesandconcomitanteliminationofunlinkedarrays.However,theoppositetrend,from linkedtowardsunlinkedarrangementwasalsosurmisedinfewspeciesindicatingpossiblereversibilityofthese processes. Conclusions:Ourresultsindicatethatnearly25%ofAsteraceaespeciesmayhaveevolvedunusuallinked arrangementofrRNAgenes.Thus,inplants,fundamentalchangesinintrinsicstructureofrDNAuni