Left-Right Asymmetry in Vertebrate Development PDF
Preview Left-Right Asymmetry in Vertebrate Development
Reviews and critical articles covering the entire field of normal anatomy (cytology, histology,cyto-andhistochemistry,electronmicroscopy,macroscopy,experimental morphologyandembryologyandcomparativeanatomy)arepublishedinAdvances in Anatomy, Embryology and Cell Biology. Papers dealing with anthropology and clinicalmorphologythataimtoencouragecooperationbetweenanatomyandrelated disciplineswillalsobeaccepted.Papersarenormallycommissioned.Originalpapers andcommunicationsmaybesubmittedandwillbeconsideredforpublicationprovided theymeettherequirementsofareviewarticleandthusfitintothescopeof“Advances”. Englishlanguageispreferred. It is a fundamental condition that submitted manuscripts have not been and will not simultaneously be submitted or published elsewhere. With the acceptance of amanuscriptforpublication,thepublisheracquiresfullandexclusivecopyrightforall languagesandcountries. Twenty-fivecopiesofeachpaperaresuppliedfreeofcharge. Manuscriptsshouldbeaddressedto Prof.Dr.F.BECK,HowardFloreyInstitute,UniversityofMelbourne, Parkville,3000Melbourne,Victoria,Australia e-mail:[email protected] Prof.Dr.F.CLASCÁ,DepartmentofAnatomy,HistologyandNeurobiology, UniversidadAutónomadeMadrid,Ave.ArzobispoMorcillos/n,28029Madrid,Spain e-mail:[email protected] Prof.Dr.M.FROTSCHER,InstitutfürAnatomieundZellbiologie,AbteilungfürNeuroanatomie, Albert-Ludwigs-UniversitätFreiburg,Albertstr.17,79001Freiburg,Germany e-mail:[email protected] Prof.Dr.D.E.HAINES,Ph.D.,DepartmentofAnatomy,TheUniversityofMississippiMed.Ctr., 2500NorthStateStreet,Jackson,MS39216-4505,USA e-mail:[email protected] Prof.Dr.H.-W.KORF,ZentrumderMorphologie,UniversitätFrankfurt, Theodor-SternKai7,60595Frankfurt/Main,Germany e-mail:[email protected] Prof.Dr.E.MARANI,DepartmentBiomedicalSignalandSystems,UniversityTwente, P.O.Box217,7500AEEnschede,TheNetherlands e-mail:[email protected] Prof.Dr.R.PUTZ,AnatomischeAnstaltderUniversitätMünchen, LehrstuhlAnatomieI,Pettenkoferstr.11,80336München,Germany e-mail:[email protected] Prof.Dr.Dr.h.c.Y.SANO,DepartmentofAnatomy, KyotoPrefecturalUniversityofMedicine, Kawaramachi-Hirokoji,602Kyoto,Japan Prof.Dr.Dr.h.c.T.H.SCHIEBLER,AnatomischesInstitutderUniversität, Koellikerstraße6,97070Würzburg,Germany Prof.Dr.K.ZILLES,UniversitätDüsseldorf,MedizinischeEinrichtungen, C.u.O.Vogt-Institut,Postfach101007,40001Düsseldorf,Germany e-mail:[email protected] 188 Advances in Anatomy Embryology and Cell Biology Editors F.F.Beck,Melbourne·F.Clascá,Madrid M.Frotscher,Freiburg·D.E.Haines,Jackson H.-W.Korf,Frankfurt·E.Marani,Enschede R.Putz,München·Y.Sano,Kyoto T.H.Schiebler,Würzburg·K.Zilles,Düsseldorf M.L. López-Gracia and M.A. Ros Left–Right Asymmetry in Vertebrate Development With17Figuresand7Tables 123 Ph.D.MariaL.López-Gracia Prof.Dr.MarianRos DepartamentodeAnatomíay BiologíaCelular FaculdaddeMedicina UniversidaddeCantabria 39011Santander Spain e-mail:[email protected] ISSN 0301-5556 ISBN-10 3-540-36347-5 SpringerBerlinHeidelbergNewYork ISBN-13 978-3-540-36347-7 SpringerBerlinHeidelbergNewYork Thisworkissubjecttocopyright.Allrightsreserved,whetherthewholeorpartofthematerialis concerned,specificallytherightsoftranslation,reprinting,reuseofillustrations,recitation,broad- casting,reproductiononmicrofilmorinanyotherway,andstorageindatabanks.Duplicationof thispublicationorpartsthereofispermittedonlyundertheprovisionsoftheGermanCopyrightLaw ofSeptember,9,1965,initscurrentversion,andpermissionforusemustalwaysbeobtainedfrom Springer-Verlag.ViolationsareliableforprosecutionundertheGermanCopyrightLaw. SpringerisapartofSpringerScience+BusinessMedia springer.com ©Springer-VerlagBerlinHeidelberg2007 Theuseofgeneraldescriptivenames,registerednames,trademarks,etc.inthispublicationdoesnot imply, even in the absence of a specific statement, that suchnames are exempt from the relevant protectivelawsandregulationsandthereforefreeforgeneraluse. Productliability:Thepublishercannotguaranteetheaccuracyofanyinformationaboutdosageand applicationcontainedinthisbook.Ineveryindividualcasetheusermustchecksuchinformationby consultingtherelevantliterature. Editor:SimonRallison,Heidelberg Deskeditor:AnneClauss,Heidelberg Productioneditor:NadjaKroke,Leipzig Coverdesign:WMXDesignHeidelberg Typesetting:LE-TEXJelonek,Schmidt&VöcklerGbR,Leipzig Printedonacid-freepaper SPIN11795179 27/3150/YL – 5 4 3 2 1 0 ListofContents 1 Introduction.............................................. 1 1.1 TheFirstMorphologicalAsymmetriesintheLeft–RightAxis ........... 1 1.2 Terminology.............................................. 2 1.3 TheAsymmetriesintheLeft–RightAxisAreHighlyConserved .......... 4 1.4 TheSitusofIndividualOrgansIsIndependentlyEstablished ............ 5 1.5 Left–RightMorphogenesis .................................... 5 1.5.1 VertebrateHeartDevelopment . ................................ 7 2 EstablishmentofLeft–RightAsymmetry.......................... 13 2.1 BreakingoftheInitialSymmetryintheEmbryo..................... 13 2.1.1 TheNodal-FlowModel ...................................... 14 2.1.2 TheTwo-CiliaModel ........................................ 17 2.1.3 TheGap-JunctionModel ..................................... 18 2.2 AsymmetricGeneExpressionintheNode ......................... 20 2.3 AsymmetricGeneExpressionintheLateralPlateMesoderm............ 24 2.4 RoleoftheMidlineinEstablishingLeft–RightAsymmetry ............. 29 2.5 LateralityDefectsinConjoinedTwins ............................ 33 2.6 RetinoicAcidandLeft–RightPatterning .......................... 34 3 SignalingPathwayswithaPredominantRoleinLeft–RightPatterning .... 35 3.1 SonicHedgehogSignalingandLeft–RightAsymmetry . ............... 35 3.1.1 IntraflagellarTransportProteinsandShhSignaling .................. 37 3.2 NodalSignalingandLeft–RightAsymmetry........................ 38 3.3 BmpSignalingandLeft–RightAsymmetry......................... 41 4 MouseModelsofLaterality ................................... 44 5 GeneticsofHumanAlterationsofOrganSitus ...................... 70 5.1 ThePrimaryCiliaryDyskinesiaSyndrome......................... 77 6 ConcludingRemarks........................................ 80 References..................................................... 96 1 Appendix ................................................ 115 A.1 CiliaryStructureandFunction . ................................ 115 2 Glossary................................................. 119 SubjectIndex................................................... 123 Acknowledgements This work was supported by grants FP12002–02946 from the Spanish Ministry of Education and Science and grant API 05/27 from the Instituto de Formación e Investigación Marqués de Valdecilla. We are very grateful to Dr. J.L. Ojeda for the SEM cilia pictures and Dr. J.M. Icardo for SEM pictures and critical reading ofsomepartsofthismanuscript.TheauthorsaregratefultoIanA.Williamsof theDepartmentofPhilology,UniversityofCantabria,forthelanguagerevisionof themanuscript.WealsoacknowledgetheexcellenttechnicalassistanceofMarisa Junco,SarayPereda,RaquelGarciaCeballos,yMiguelMier. Abbreviations ActRIIa ActivinreceptorIIa Acvr1 ActivinreceptorI Acvr2b ActivinreceptorIIb Aldh1a2 Aldehydedehydrogenasefamily1subfamily2 AP Antero-posterior ASE Asymmetricenhancer Bmp Bonemorphogeneticprotein BPI Bactericidal-permeability-increasingprotein Car Caronte Cer Cerberus CFC Crypto-FRL1-cryptic CHD Congenitalheartdefects Chrd Chordin CITED CBP/p300interactingtransactivatorswithED-richtermini co-Smad Common-partnerSmad CRELD1 Cysteine-richproteinwithEGF-LIKEdomain1 Cx43 Connexin43 Dand5 DANdomainfamilymember5 Dhh Deserthedgehog Disp Dispatched Dll1 Delta-like1 Dna1 Dyneinaxonemalintermediatechaintype1 Dnah7 Dyneinheavychain7 Dnah9 Dyneinheavychain9 Dnahc11 Dyneinaxonemalheavychain11 Dnahc2 Dyneinaxonemalheavychain2 Dnahc5 Dyneinaxonemalheavychain5 Dnai2 Dyneinintermediatechain2 Dnal1 Dyneinaxonemallightchain1 DORV Double-outletrightventricle DV Dorso-ventral Dync2h1 Dyneincytoplasmic2heavychain1 Dync2li1 Dyneincytoplasmic2lightintermediatechain1 E Embryonicday X Abbreviations EGF Epidermalgrowthfactor Fgf Fibroblastgrowthfactor8 flh Floatinghead Fltn Flectin Fts Fusedtoes Gdf1 Growthdifferentiationfactor1 GJA1 Gapjunctionalpha-1(Gapjunctionprotein43KD) Hfh4 Hepatocytenuclearfactorforkheadhomolog4 HH HamburgerHamilton hLAMP1 Heart-specificlectin-associatedmatrixprotein-1 β β HNF3- Hepatocytenuclearfactor3- ICS Immotile-ciliasyndrome IFT Intraflagellartransport Ift172 Intraflagellartransportprotein172 Ift88 Intraflagellartransportprotein88 Ihh Indianhedgehog Inv Inversionofembryonicturning Invs Inversin IP6 Inositolhexakisphosphate Ipk1 Inositol1,3,4,5,6-pentakisphosphate2-kinase I-Smad InhibitorySmad Iv Inversusviscerum Kif3a Kinesinfamilymember3A Kif3b Kinesinfamilymember3B Kif5C Kinesinfamilymember5C LEF1 Lymphoidenhancerbindingfactor1 Lefty1 Left–rightdeterminationfactor1 Lefty2 Left–rightdeterminationfactor2 Lfng Lunaticfringe Lgl Legless LPM Lateralplatemesoderm LR Left–right Lrd Left–rightdynein LSE Left-side-specificenhancer Mgat1 Mannosideacetylglucosaminyltransferase1 Mid1 Midline1 Mid2 Midline2 NCX1 Sodium–calciumexchanger Nkx3.2 Bagpipehomeoboxgene1homolog Nog Noggin NPHP2 Nefronophthisistype2 Nt Noturning Ntl Notail Abbreviations XI Ofd1 Oralfacialdigitalsyndrome1 OMIM OnlineMendelianinheritanceinman PC1 Polycystin1 PC2 Polycystin2 PCD Primaryciliarydyskinesia Pcl2 Polycomblike2 Pitx2 Paired-likehomeodomaintranscriptionfactor2 PKA ProteinkinaseA Pkd2 Polycystickidneydisease2 PKI ProteinkinaseAinhibitor α PKI PKAinhibitor PLUNC Palatelungnasalepitheliumclone λ λ Pol- DNApolymeraseXfamilymember Psck6 Proproteinconvertasesubtilisin/kexintype6 Ptc Patched RA Retinoicacid R-Smad Receptor-regulatedSmad Rttn Rotatin Shh Sonichedgehog Slc8a1 Solutecarrierfamily8 Smo Smoothened SSCP Single-strandconformationpolymorphism Stil Scl/Tal1interruptinglocus Sufu Suppressoroffused T Brachyury TCF TranscriptionfactorTcellspecific TFAP2 TranscriptionfactorAP2 TGA Transpositionofthegreatarteries β β TGF Transforminggrowthfactor THRAP2 Thyroid-hormone-receptorassociatedprotein2 TTTC8 Tetratricopeptiderepeatdomain8 Wnt Amalgamfromwingless(wg)andmammarytumorintegration(int) Zic3 Zinc-fingerproteinofthecerebellum3
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