Developmental Biology | |
A Regulatory Gene Network That Directs Micromere Specification in the Sea Urchin Embryo | |
Eric H. Davidson1  Paola Oliveri1  Deanna M. Carrick1  | |
[1] Division of Biology 156-29, California Institute of Technology, Pasadena, California, 91125 | |
关键词: sea urchin embryo; homeodomain; pmar1; tbrain; delta; gene regulatory network; | |
DOI : 10.1006/dbio.2002.0627 | |
学科分类:生物科学(综合) | |
来源: Academic Press | |
【 摘 要 】
Micromeresandtheirimmediatedescendantshavethreeknowndevelopmentalfunctionsinregularlydevelopingseaurchins:immediatelyaftertheirinitialsegregation,theyarethesourceofanunidentifiedsignaltotheadjacentveg2cellsthatisrequiredfornormalendomesodermalspecification;afewcleavageslater,theyexpressDelta,aNotchligandwhichtriggerstheconditionalspecificationofthecentralmesodermaldomainofthevegetalplate;andtheyexclusivelygiverisetotheskeletogenicmesenchymeofthepostgastrularembryo.Wedemonstratethekeycomponentsofthezygoticregulatorygenenetworkthataccountsformicromerespecificity.ThisnetworkisasubelementoftheoverallendomesodermspecificationnetworkoftheStrongylocentrotuspurpuratusembryo.Acentralroleisplayedbyanewlydiscoveredgeneencodingapairedclasshomeodomaintranscriptionfactorwhichinmicromeresactsasarepressorofarepressor:thegeneisnamedpmar1(paired-classmicromereanti-repressor).pmar1isexpressedonlyduringcleavageandearlyblastulastages,andexclusivelyinmicromeres.Itisinitiallyactivatedassoonasthemicromeresareformed,inresponsetoOtxandβ-Catenin/Tcfinputs.Therepressivenatureoftheinteractionsmediatedbythepmar1geneproductwasshownbytheidenticaleffectofintroducingmRNAencodingthePmar1factor,andmRNAencodinganEngrailed-Pmar1(En-Pmar1)repressordomainfusion.Inbothcases,theeffectsarederepression:ofthedeltagene;andofskeletogenicgenes,includingseveraltranscriptionfactorsnormallyexpressedonlyinmicromeredescendants,andalsoasetofdownstreamskeletogenicdifferentiationgenes.ThespatialphenotypeofembryosbearingexogenousmRNAencodingPmar1factororEn-Pmar1isexpansionofthedomainsofexpressionofthedownstreamgenesovermostoralloftheembryo.Thisresultsintransformationofmuchoftheembryointoskeletogenicmesenchymecellsthatexpressskeletogenicmarkers.Thenormalroleofpmarlistoprevent,exclusivelyinthemicromeres,theexpressionofarepressorthatisotherwiseoperativethroughouttheembryo.Thisfunctionaccountsforthelocalizationofdeltatranscriptioninmicromeres,andtherebyfortheconditionalspecificationofthevegetalplatemesoderm.Italsoexplainswhyskeletogenicdifferentiationgenebatteriesnormallyfunctiononlyinmicromeredescendants.Moregenerally,theregulatorynetworksubelementemergingfromthisworkshowshowthespecificityofmicromerefunctiondependsoncontinuingglobalregulatoryinteractions,aswellasonearlylocalizedinputs.
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