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1、REVIEWSTheevolutionofgeneregulationbytranscriptionfactorsandmicroRNAsKevinChen*andNikolausRajewsky*‡Abstract
2、Changesinthepatternsofgeneexpressionarewidelybelievedtounderliemanyofthephenotypicdifferenceswithinandbetweenspecies.Althoughmuchemphasishasbeenplacedoncha
3、ngesintranscriptionalregulation,geneexpressionisregulatedatmanylevels,allofwhichmustultimatelybestudiedtogethertoobtainacompletepictureoftheevolutionofgeneexpression.Herewecomparetheevolutionoftranscriptionalregulationandpost-transcriptionalregulationthatismediate
4、dbymicroRNAs,alargeclassofsmall,non-codingRNAsinplantsandanimals,focusingontheevolutionoftheindividualregulatorsandtheirbindingsites.Asaninitialsteptowardsintegratingthesemechanismsintoaunifiedframework,weproposeasimplemodelthatdescribesthetranscriptionalregulatio
5、nofnewmicroRNAgenes.Theemergenceofcomplex,multicellularorganismsprimacyoftranscriptionasthenecessaryfirststepinwasaccompanied,andperhapsfacilitated,bydramaticgeneexpressionisundeniable,thisdoesnotimplythatincreasesinthecomplexityofgeneregulatorymecha-transcription
6、alregulationhasthelargesteffectonthenisms1,2.Attheleveloftranscriptionalregulation,thisfinalconcentrationoftheactivegeneproduct,whichiscanbeclearlyseeninthemassiveexpansionsofthemostrelevantquantitytothephenotype.transcription-factorfamiliesandthepervasivecom-Anim
7、portantgoalforfutureresearchistoeluci-binatorialcontrolofgenesbymultipletranscriptiondatehowcomplexgeneregulatorynetworksevolveandfactorsinhigherorganisms1,3(BOX1).Atthelevelofpost-howtheirevolutionresultsinphenotypicchangetranscriptionalcontrol,entirelynewmechani
8、smsofgeneandspeciation.However,anecessaryfirststeptowardsregulationarose,typifiedbyalargeandgrowingclassthisgoalistounderstandthebasicprinciplesthatof~22-nucleotide-longnon-codingRNAs,knownasunderlietheevolutionoftheindividualregulatorsandmicroRNAs(miRNAs),whichfu
9、nctionasrepressorsintheirregulatoryinteractionswiththeirtargetgenes.allknownanimalandplantgenomes4,5(BOX1).AlthoughRecentcomputationalandexperimentalwor
