Table Of Content

Bel I EIGHTH EDITION Scott.G ilbert SwarthCmoolrlee ge by Witahc hapotneP rL ANDTE VELOPMENT SusaSni nger R. CarleCtoolnl ege SinauAesrs ociaItnecPs.u,,b l ishers SunderlaMnads,s achuseUtStAs Deveo a Biology EIGHTH EDITION �• •••• t.,. Scot.tG ilbert SwarthCmoolrlee ge by Witahc hapotneP rL ANDTE VELOPMENT SusaR.n Singer CarleCtoolnl ege SinauAesrs ociaItnecPs.u,,b lishers SunderlMaansds,a chuUsSeAtt s About Ctohvee r Zebrfias h rerio L.-� subjefcottrsh es tudoyfc elmlo vementssi,n ce (Danio -Gl t thee mbryoi sl ram:'.,i lopsq uickalnydi, t dse velopmteanktep sl acoeu t- .&: :L �- sidaeu terus _ - _ t7lefiaotfit ohenz e bfrias nhe rvoussy stecme lilsss e en ort"i �.. ... int het riply embryToh.e m icrogrlaopcha ltihzreesge e nper oducts .;.: thahte lp <.;n - �--n_sIo ft heb raiTnh.e o tx2 genpe oduct( anterdiaorrpk u r ';'<'_ --¢ r ple ins pecifytihnefg o rebraanidmn i dbratihnek; ro x20g ene regi -;...lnt prod:.;� · _ purp"lset ripieses x"p)r esisne!:' dN op lacaensd i sc ritical for � -.t;nOi spoeoir.-: - =dbraina;n dt hep roducotfts h ee ngragielne(edg olhde)l fpo rmt he ",-_�=:-t..;ndb raibno rdearn da rae lssoe einn s omem usclcee lalJso ng midline the (seCeh apte1r1as n d1 2)M.o rphoge(ntehgseei nse ratoiffo onr mi)s �:-)O ··moviinew "h icshe vedriaffle resnttaliyn ed ofz ebrfias hec toder rcgioos ....1. jt ao f ormt hen eurtaulb eN.u meroumsu tatiionnz se brafcainsc ha usfea ulm ty ....e-,n tubfeo rmatiporno,d ucidnigs eases to manb itrh defecSttsa.i ned . Tal similarII I embryiom agkei ndplryo vidbeyDd r .G .H aupunann and Verlamgo;v ie Springer- imagegse neroupsrloyv idbye dD r. K.ffu .ra. DEVELOPMEBNITOALLO GEYi,gh Etdhi tion C<lryright2 00b6y S inauAesrs ociIantce.s righrtess ervTehdib.so okm ayn ot © All ber eproduicnew dh oloer i np arwti thotuhtep ermissoifto hne p ublisSbienra.u er AssociaItnecs2.,3,P lumtrReoea dS,u nderlaMnAd O,l 37U5S A. emaiplu:b [email protected]@sinauer.com www.smauer.com LibraorfyC <lngreCsast aloging-in-PubDlaitcaa tion GilbeSrcto,t t1 949- E, Developmebnitoallo IgS yc ottG ilb;ew rittahc haptoenrp lant E developmbeynS tu saRn. S inger8-the d. pcm. . Includbeisb liograrpehfiecraelan ncdei sn dex. ISBN9 78-0-87893-(2h5a0r-d4c over) 1.E mbryolo2g.Dy e.v elopmebnitoallo 1gy..T itle. QL955.G240806 571.8-dc22 2006004780 PrintienCd h ina 6 5 4 3 DaniSealr,a ahn,dD avid To BriCeofn tents Principdleevse loofp mebnitoallo gy PAR T 1 Chapt1e rD evelopmebnitoallo Tghyea: n atomitcraald it3i on Chapt2e rL ifcey claensdt heev olutoifdo env elopmepnattatle rn2s5 Chapt3e rP rinciopfel xepse rimeenmtbarly olo4g9y Chapt4e rT heg eneticocfdo ervee lopm7e7nt Chapt5e rT hep aradiogfdm i fferengteinaeelx press1i0o1n Chapt6e rC ell-ccoemlmlu nicaitndi eovne lopm1e3nt9 Earleym bryondiecv elopment PAR T 2 Chapt7e rF ertilizationa:n eBweo grignanniin1sg7m 5 Chapt8e rE arldye velopmiensn etl ecitnevde rteb2ra1t1e s Chapt9e rT heg enetoifca sx issp ecificianDt rioosno ph2i5la3 Chapt1e0rE arldye velopmaenndat x ifso rmatiinao mnp hibia2n9s1 Chapt1e1rT hee ardleyv elopmoefvn etr tebrFaitsebhsi,:r dasn,dm ammals3 25 Later embdryeovneicl opment PAR T 3 Chapt1e2rT hee mergenocfte h ee ctodeCremn:t ral nseyrsvtoaeunmsd e pider3mi7s3 Chopt1e3rN eurcarle csetl alnsda xonsaple cific4i0ty7 Chapt1e4rP araxainadil n termedmieastoed er4m4 3 Chapt1e5rL aterpalla mtees odearnmd e ndoder4m7 1 Chapt1e6rD evelopmoeftn htte e trolpiomd5b 05 Chapt1e7rS exd etermina5t2i9o n Chapt1e8r P ostembryodneivce lopmMeentta:m orphroesgiesn,e raatnidoo gni,n 5g5 5 Chapt1e9rT hes agoaf tgheer ml in5e9 3 Ramificatoifdo envse lopmebinotlaolg y PAR T 4 Chapt2e0rA no vervioefpw l andte velopm6e2nt7 Chapt2e1rM ediciamlp licaotfid oenvse lopmebnitoallo 6g5y5 Chapt2e2rE nvironmernetgaull aotfio onni madle velopm6e9nt3 Chapt2e3rD evelopmemnetcahla niosfme sv olutiocnhaarnyg7 e2 1 Contents Princisop fl e PAR T 'De1vel opamle nBtiology CHAPTE1R DeveloBpimoelnotgayl: 3 TehA natoTmriacdailt ion TheQ uestioofnD se velopmeBnitoallo 4gy EvolutioEnmabrryy ology 15 AppraochetsoD evelopmellBn itoalogy EmbrYOl'lhiocm ologies 5 16 TheA natomiAcpaplr oach MedicEamlb ryolaongdyT eratol1o7g y 5 Comparateimvber yolo5g y Genetmiacl formoantsai nds yndromes 18 Epigeneasnidps r eformation Disruptainodnt se ratogens 6 18 Namintgh ep artTsh:ep rimagreyr ml ayearnsde arly MathematMiocdaell ionfDg e velopme1n9t organ8s Them athematoifoc rsg anismal growth 19 Thef ouprr inciopflK earslE rnsvto nB aer8 Them athematoifpc ast tern2i1n g Fatmea ppintgh ee mbryo 10 Coda2 3 TheC elluBlaasrio sfM orphogene1s3i s Celmli gration 13 CHAPTE2R 25 LiCfyec alnetdshE ev olouDfte ivoenl oPpamtetnetranls TheC ircolfLe i fTeh:eS tagoefsA nimal .SIDWGNTS£.SP(CULATIROuNlSeos:fE videnc3e9 I Developme2n5t DevelopmiennF tl owerPilnagn t4s0 Thefr ogl ifcey cle 26 DevelopmePnattatle armnosn gt heM etazo4a2 MulticellTuhleaE rviotlyu:to ifDo inf ferentainadt ion TheD iploblParsottso,s toamnedDs e,u terosto4m4e s Morphogenesis 31 .SIDUlGHT£.SS P{CULAHTS:IT Ootipotency tahmeo ng Thev olvocaceans 31 Flatworms £.IPKllLATlOH:IS ex 44 .llDUlGIITI anIdn diviidnu aVloiltyv ox 33 Differentiaantdim oonr phogenine sDiisc tyostelium 36 viii CONTENTS PrinciopfEl xepse rimeEnmtbarly ology CHAPTE3R 49 EnvironmeDnetvaell opmeBnitoallo 4gy9 Morphoggenr adients r6e3v isited Environmesnetxda elt ermina5t0i on .SIRDlGIm{' I:I SPHULATIOSH:R uloefs EvidCeonncet r6o5 ls Adaptatoifoe nm bryoasn dl arvateo t heir Stemc elalnsd c ommitme6n6t environme5n0t s MorphogenaensdiC se ll Adhe6s7io n TheD evelopmeDnytnaalm iocfsC elSlp ecific5at3i on Differencteilaallffi nit6y8 Autonomosupse cifica5t3i on Thet hermodynammoidce olf c eliln teract7i0o ns Syncytsipaelc ifica5t6i on Cadheriannsdc elald hesi7o1n Conditiosnpaelc ifica5t7i on Coda7 4 TheG enectC ioroefD evelopment CHAPT4E R 77 TheE mbryologOirciaglio nfts h eG eneT heor7y 7 RNA LocaliTzeacthinoinq u8e7s Nucleoursc ytoplaWshmi:c hc ontrhoelrse di7t7y ' Detectsipencgi mfiRcN AsR:T -PCR8 7 Thes plbiett weeemnb ryolaongdyg eneti7c9s !\1irocarray8s9 Earlayt tempattds e velopmegnetnaelt i7c9s iVlcaroraarys8 9 EvidenfcoeGr e nomiEcqu ivalen8c0e LocatimnRgN Asi ns pacaen dt imeI:ns it u hybridiza9t0i on Amphibicalno niTnhge:r estriocftn iuocnl epaort en8c1y Amphibianc loniTnhget: o tipootefsn ocmyat ciecl ls DetermintihneFg u nctioofGn e nedsu ring 82 Developme9n2t Cloning als 83 m mm SIDUIGaImS{.P HULATISO:WH hyC lonMea mma?l 8s5 Transgecneilacl nsd o rganis9m2s • Determintihnefg u nctioanm eosfs aAgnet:i sense9 R6N A DifferenGteinaeEl x press8io6n Coda9 8 TheP aradiogfDm i fferenGteinaEelx pression CHAPT5E R 101 DifferenGteinaeTl r anscrip1t0i1on SIDEUGHTSSP ECULANTS:IT OheM echaniosfXm s • £. ChromosIonmcatei vation Anatomoyft heg eneA:c tiavned repressed 1c0h1r omatin 123 Anatomoyf t he gene: Exons 1a0n2d introns DifferenRtNiAaP lr ocess1in2g5 Anatomoyft he gene: Promeonthearnsc e1ar0ns3d Controofle ardleyv elopmbeynn tu cleRaNrA select1i2o5n Transcrifpatcitoon1r 0s8 Creatifnagm iloifep sr toeinst hroudgifhfre entiln RNA SIIlUlGIm{'S P(CULATSI:PO oHeeTrnascriptniF oacto1r1s2 a • in r splic1i2n6g Transcrifpatcitoconar s cad1e1s3 Spliceinnhga ncaernsdr ecogniftacitoon1r2 s8 SII)(UG�TSSP (CULATI:SO tHuSdyDiNnAgR egulatory • f, SII)(UGr.IS mP ((ULAll:DD HSiffereniatln RNPAr ocesasning Elmeent1s1 3 • d SeDxe termination Silenc1e1r6s Drosophila 129 ControofGl e neE xpressaitto hneL eveolf MethylatPiaotnt earnndt heC ontroofl . Translat1i3o0n Transcrip1t1io6n Differentiall omnRgNeAv 1i3t0y DNA methylatainodgn e naec tiv1i1t6y Selectiinvhei biotfim oRnN A translSattoirooenod:c yte SII)(UG�{.T SIP ECULAT:IG OeHSnomIimcp rinting mRNAs 130 • 117 Chromatmuold ificiantdiuocnbe yDd N A ControoflR NA expressbiyco ytno plasmic methylat1i1o8n localiza1t3i5o n Insulat1o1r9s SIDRIGIm {. SPICULATSI;mO iRHNAi nT ransipctroinalG ene • Transcrirpetgiluaoionna olfa ne nticrher omosome: Regulation t 135 Dosagceo mpensa1t1i9o n Posttranslraetgiuolnaatolif go enn eexp ressi1o3n7 CONTENTSi x Cel-ClelClo mmunicaitniD oenv elopment CHAPTE6R 139 InductainodCn o mpetence SDIUIIfGfi&SPHULA,T telOHS 157 139 • S m Cell Niches Cascadoef.si nduction: aRnedcs iepqruoecnatli al Othepra racrfiancet1 o5r8s inductive1 4e1v ents CelDle ath Pathways 158 Instruacntdpi evrem isisnitveer ac1t4i2o ns JuxtacrSiingen aling Epithelial-meisnetnecrhayc1mt4ai3lo ns 160 TheN otcpha thwjauyx:t aploisgeadan nddrs ecept1o60r s ParacrFiancet oTrhseI: n ducer Mol1e4c5u les SlDUIflfiG&S PUULAT:IJ OuHxStacirngailinngea ndC ell • S SignTarla nsducCtaisocna dTehse:R espontsoe Patteml1n6g2 Inducers Thee xtracemlalturlaiasaxr s ourocfde e velopmental 147 sign1a6l4s Fibrobglraoswtft ahc taonrdts h ReT K path1w4a7y Diretcrta nsmisosfsi iognn tahlrso uggahpj uncti1o6n7s TheJ AK-STAT pa1t50h way TheH edgehog f1a51m ily Cross-Talk between Pathways 168 TheW ntf ami1l5y3 Maintenantchee D oiff ferentSitaatteed 169 The TGsFu-p�e rfa1m5i5l y Coda 171 P2AE aRrElmyTb ryocn i Development FertilizBaetgiiotnl:ll ing CHAPTE7R aN ewO rganism 175 StructouftrheeG ametes Latree sponses: Reosfpu rmopttaeiniodnnD NA 175 synthe1s9i4s Sperm17 5 SlDWG&HTS SP HULAT:IT OhKSAec tivation Thee gg1 78 • ofGa mete Recogniotfei goagnn ds per1m8 1 Metabo1l9i5s m Fusioofng enemtaitce r1i9a8l External FeritniS leiaUz racthiionns 181 MammaliFaenr tilization Speramt tracAtcitoinao:tna d ista1n8c1e 199 Getttihngega metienstt oh oev iduTcrta:n slocaaDtdi oo Thea crosormeea cti8o3n capacit1a9t9j on t Species-srpeeccoigfinci1 t8i4o n Int hvei cionfit thyoe o cyHtyep:e ractivation, Fusiooftn h eeg agn ds percme mlelm bran1e8s6 thermotaanxdci hse,m ot�2x0i1s TheP reventoifPo onl yspermy Recognittihozeno napate lluc2i02d a 187 Thef asbtlo ctkop olyspe1r8m7y Gametfeu siaonndt hper evenotfip oonl yspe2r0m4y Thes lobwl octkop olyspe1r8m9y Fusioofng enemtaitce r2i0a5l Calciaustm h ien itioaftt hocero rtigcraaln ule Coda 207 react1i9o0n {,SP{( ULATIOKS: The Nonequivaloefn ce • SlDUIGIiTS ActivaotfiE ogngM etaboliinsS me aU rchins MammaliParno nuclei . 193 207 Earrleys pon1s9e3s x CONTENTS CHAPTERE arlDye velopmienSn etl ecbtweedr tebr2a1t1e s 8 SlDUIGIITSSf.P HULAn:OA dNaSptatiboyMn o difying EARLDYE VELOPMENTPARLO CESSES: • EmbryoniCcI .avage ANO VERVIE2W1 1 233 GastrulaitnSi noani 2l3s7 Cleava2g1e1 EARLDYE VELOPMEINNTT UNICATE2S3 7 Fromf ertilizactlieoanv a2tg1oe2 Thec ytoskemleecthaaln isomfms i tos2i1s3 Tunicate Cl2e3a7v age Patteorfne sm bryoncilce ava2g1e5 Thet unicfaatteme a p 237 Gastrula2t1i5o n Autonomoaunsdc onditisopneacli ficoaftt uinoinc ate hlastome2r3e9, CelSlp ecificaantdAi xoins Forma2t1i6o n Specificoaftt hieeomn b ryonaixce 2s4 1 EARlYD EVELOPMENSTEIANU RCHINS2 17 GastrulaitnTi uonni cat2e4s1 SeaU rchCilne ava2g1e7 THEN EMATODCEA ENORHABOEmLSE GAN2S4 3 Blastfuolram ati2o1n7 WhyC .e lega2n4s3? Fatmea psa ndt hed eterminaotfsi eouanr chin blastome2r1e7 s CleavaagnedA xiFso rmatiionCn .e lega2n4s3 SeaU rchGians trula2t2i3o n Rottaioncalle avoafgt eh C.e elegeagng2 ,4 3 ingressoifto hne primary mes2e2n3c hyme Anterior-poasxtifesor rimoart i2o4n3 Firsstta gofea rchentienrvoang ina2t2i6o n Formatiooftn h ed orsal-vaenndtr riaglh t-alxeeft2s 4 7 Seconadn dt hirsdt agoefsa rchentienrvoang ina2t2i7o n Controoflb l astomiedreen t2i4t7y Integraotfai uotno nomoaunsdc onditisopneacli fication: THEE ARLDYE VELOPMEONFTS NAILnS9 Differentioaftt ihoeC.n e iegpahnarsy n2x4 9 Cleavage iEnm bSrnyaoi2sl2 9 GastrulaitnCi .eo lne ga2n4s9 A fatmea po fI lyanoabsssoal2 3e2t a Coda2 50 Thep olalro bCee:l dle terminaatnidao xnif so rmati2o3n2 CHAPTERT he GenoefAxt i icsSs p ecificaitnDi roons ophil2a5 3 9 EARLDYR OSOPHDIELAV ELOPMEN2T5 3 Thea nteroirogra nizcienngt eTrh:eB icoaindd Hunchbagcrka die2n7t1s Fertiliza2t5i4o n Thet ermigneanlge r ou2p7 4 Cleava2g5e4 Anterior-poasxtisespr eicoirf icSautmimoanr:y2 7 5 Them id-blasttrualnas it25i6o n SegmentatGieonne s2 75 Gastrula2t5i6o n StOUlGIITS f.SP ICULAT:IS eOgNmSenatns?d a rasegments • 276 GENEST HAT PATTTEHREN DROSOPHBOIDLYAP LAN Theg apg ene2s7 7 258 Thep air-rguelnee2 s7 8 PrimaArxyi Fso rmatidounr iOnogg enes2i5s9 Thes egmepnotl argietnye 2s8 1 Anterior-popsotlearriiionttr yh eo ocyt2e5 9 TheH omeotSiecl ecGteonre s2 83 Dorsal-vepnattrtaelr niintn hgeo ocyt2e5 9 Patteorfnh so meotgiecn eex press2i8o3n GeneratDionrgs al-VePnattrtaeilrnt n h eE mbryo26 3 Initiatthien gp attehronmseo toigfce neex press2i8o5n Dorsalth,ev entrm.olr phoge2n6 3 M<lintaintihnpega tteorfnh so meotgiecn eex press2i8u5n Establiasn huicnlge daorr sgarla die2n6t3 Realisgaetnoer2s 8 5 E1f.:cts oft hed orsparlo tegirna die2n6t4 • SIDGlWlTSf.S P[(ULlTIOTHhSHeo :m eodomaPirno te2in86s Sejjmentaatnidto hneA nterior-PoBsotdeyr iPol2ra6 n6 Axeasn dO rgaPnr imordTihaeC: a rtesCioaonr dinate .Ia:<gmrala diePnotlsa:r rietgyu labtyio oonc yte Model2 88 _ �nplasm2 66 Coda2 88 :t;., molecumloadre lPr:o tegirna dieinntt hsee arly em rm 266