R=Sox6-A Versatile Regulator invertebrate development2011Hagiwara

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cimanyD latnempoleveDDEVELOPMENTALDYNAMICS240:1311–1321,2011

REVIEWS–APEERREVIEWEDFORUM

Sox6,JackofAllTrades:AVersatileRegulatoryProteininVertebrateDevelopment

NobukoHagiwara*

Approximately20,000genesareencodedinourgenome,onetenthofwhicharethoughttobetranscriptionfactors.Consideringthecomplexityandvarietyofcelltypesgeneratedduringdevelopment,manytranscrip-tionfactorslikelyplaymultipleroles.UncoveringtheversatilerolesofSox6invertebratedevelopmentshedssomelightonhowanorganismef cientlyutilizesthelimitedresourcesoftranscriptionfactors.Thestruc-tureoftheSox6geneitselfmaydictateitsfunctionalversatility.First,Sox6containsnoknownregulatorydomains;instead,itutilizesvariouscofactors.Second,Sox6hasalong30-UTRthatcontainsmultiplemicro-RNAtargets,thusitsproteinlevelisdulyadjustedbycelltype–speci cmicroRNAs.JustcombiningthesetwocharacteristicsalonemakesSox6extremelyversatile.Todate,Sox6hasbeenreportedtoregulatediffer-entiationoftissuesofmesoderm,ectoderm,andendodermorigins,makingSox6atrulymultifacetedtran-scriptionfactor.DevelopmentalDynamics240:1311–1321,2011.VC

2011Wiley-Liss,Inc.

Keywords:Sox6;Soxfamilyoftranscriptionfactors;terminaldifferentiation;vertebratedevelopmentAccepted21March2011

INTRODUCTION

bodyplan(Bowlesetal.,2000;identi edinmodelorganismsofgna-Whenweembarkonassigningspeci cSchepersetal.,2002;Phochanukulandthostoma(vertebrateswithjaws)rolestoindividualgenes,genefamiliesRussell,2010).VertebrateSoxgenes(Bowlesetal.,2000),exceptforfugupresentachallengingtaskbecauseofhavebeengroupedintoeightgroups(puffer sh),whichcontainstwoSox6theiroftenhard-to-de nefunctional(A–H)basedonsimilaritiesintheirgenes(Sox6AandSox6B)(Koopmanredundancies.Teasingoutspeci crolesaminoacidsequences(Schepersetal.,etal.,2004).Re ectingtheevolution-forindividualmembersofagenefamily2002).Sox6(alongwithSox5andaryproximityofthevertebrateSoxDismadedif cultbecausegenefamiliesSox13)belongstotheSoxDfamily.Thisgenes,manyoverlappingdevelopmen-evolvethroughtheprocessofrepeatedfamilyappearstohaveevolvedfromatalfunctionshavebeenreportedforgeneduplicationsandsubsequentgenesingleancestralSoxDgene,appearingSox5andSox6inmultipletissues(Le-loss.Therefore,themorerecentthe rstinBilatera(Bowlesetal.,2000;febvreetal.,1998;Stoltetal.,2006;Le-duplicationevent(i.e.,youngerfamilyLarrouxetal.,2008;Phochanukulandfebvre,2010).Expressionofthethreemembers),thegreaterthelikelihoodRussell,2010).InvertebratessuchasSoxDgeneshasbeendetectedinawideDrosophila,Caenorhabditiselegans,varietyofvertebratetissues(RoosethatfunctionalredundancywillbeandseaurchinpossessonlyoneSoxDetal.,1998;Hagiwaraetal.,2000;seen.IntheSoxgenefamily,Sox6isgene(Bowlesetal.,2000;PhochanukulKasimiotisetal.,2000;Cohen-Barakoneoftheseyoungbloods.TheSoxfam-andRussell,2010),asdoesthelampreyetal.,2001;Ikedaetal.,2002),anditisilyoftranscriptionfactorscurrentlyofthetaxoncyclostomata(jawlesslikelythatthereareagreatdealofconsistsofaboutadozengenesin shes)(Ohtanietal.,2008).Asingleunknownfunctionssharedbytheseinvertebratesand20genesinverte-copyofthreemembersoftheSoxDfam-SoxDgenes.Oneofthemainchal-brates,relativetothecomplexityoftheily(Sox5,Sox6,andSox13)hasbeenlengesthenliesindiscerningbetween

DivisionofCardiovascularMedicine,DepartmentofInternalMedicine,UniversityofCalifornia,Davis,CaliforniaGrantsponsor:MDA;Grantnumber:4135;Grantsponsor:NIH;Grantnumber:R01AR055209.

*Correspondenceto:NobukoHagiwara,DivisionofCardiovascularMedicine,DepartmentofInternalMedicine,UniversityofCalifornia,Davis,OneShieldsAve,Davis,CA95616.E-mail:nhagiwara@ucdavis.eduDOI10.1002/dvdy.22639

Publishedonline14April2011inWileyOnlineLibrary(http://www.wendangwang.com).

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C2011Wiley-Liss,Inc.

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