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369 Braun, l'{. (1882). entwicklungsvo.gägn" am Schi^¡anzende bei einigen Så.ugethÌeren mit Berücks¡chtìgung der Verhältn¡sse bein menschen. Arch. Anat. Physiol i Anat. Abt. 207"1 Bremer; J. L. (1952). Dorsal ¡ntestinal fîstula; accessory neurenteric canal; dlastematomyel ia. Arch. Path. 54:132. Brocklehurst, G. (1969). l'he development of ih. hur"n cerebrosplnal fluid pathr^ray with particuiar reference to the roof of the fourth ventr¡cle. J. Anat. 105:467. Brouwer, B. (1916). Klinische. - anatomîsche Untersuchung 'tiber partielle. Anencephal ie. Z. Ges. Neur. Psychíat. 32t16\. Brown, M.G., Hamburger, V. and Schmltt, F.0. (1941). Density studies on amphlbían embryos wíth special reference to the mechanism of organizer actîon. J.. Exp. Zool. 88:353. Browne, D. (1934). Tal ipes equino-varus. Lancet. ll:969. Browne, D. (1967). A mechanist¡c înterpretion of certain m_alfermatlons. Adv. ln Teratol.2:11. Bryden, H. 14., Perry, C. and Keeler, R.F. (1973). Effects of alkaloids of Veratrum cal ifornic.um on.chîck embryos. Teralol. 8:19. Burda, D. J. (1968). Studies on the experimental inductíon of over- . grov,rth in chick embryos. Anat. Rec. 161:419. Burda-Wi lson, D. (1971). Distríbution of thymidine - H3 ln the overgrown brain of the chlck embryo. J. Comp. Neurol. 141t37. Burnside, B. (.l971). |,licrotubules and mlcrofllaments ¡n ne.wt neurulation Devel . Biol. 26:416. , Butros, J.(1962). Studîes on the ¡nductive action of the early chick axls on the fsolated post-nodal fragments. J. Exp. Zool. 149¡'l.
Butros' J. (1965). Action of hearr and liver R.N.A. on the differentiation of segmen.ts of. chick blastoderms. J. Enb. Exp. llorphol . 13:1.19. Caffey, J, .(19721, ln, Pediatrlc X-Ray Diagnosis. 6th Edirlon. year Book Hedical Pub. Ch i cago. Caneron, A.H. (1956). The spînal cord leslon in spiria biflda cystica. Lancet ll:1/1 . Carter, C.0. (1969). Spina bifida and anencephaly: a problem in genetÌcenvlronmental interactíon. J. Biosoc. Sci. l:71 Carter, T.C. (1956). Genetics of the Little and Bagg X-rayed mouse stock. J. Genet. 54:311. Carter, T- C. (1959). Embryology of the Little and Bagg X-rayed mouse stock. J. Genet. 56 :401 . Catlzone, 0. and Gray, P. (1941). Experiments wíth chemicêl interference of early morphogenesis of the chick. ll. Effects of lead on C.N.'S. J. Exp. Zool . 87:71 . Coffey, V.P., and Jessop, tr.J.E. (,|957). A study of 137 cases of anencephaly. Br¡t. J. Prev,Soc. l4ed. lt:174. Cohlan, S.q. (1954) . Congenital anomalies ìn the rat produced by excess intake of vitam¡n A ín pregnancy. paediatrics 132556. Corl iss, C.E.,' Fedinec, A.A. and Robertson, c.G. (,|966). . Teratogenlc effect of teta¡ìus toxin on c.N.S. of early chick embryos. Anat. Rec. 15\..221 - Corliss, C.E. and Robertson, c.c. (1959). Cephalocaudal and alar-basal variations in the neural ep¡thel ium of 36 hour chick ernbryos. Anat. Rec. 1 33:446. Corliss, C.E. and Robertson, G.G.(1963). The pattern of. mitotic densi ty in the early chlck neural epithel ium. J. Exp. Zool . 15j¿125. 370
Page 1 and 2:
E}4BRYOGENES IS OF EXPERIHENTALLY I
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MY PARENTS c.J.H. G. M.
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lt Congenital malformêtlons of the
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TABLE OF CONTENTS SECT ION PAGE 1 I
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},I.ATE R I ALS 3.1 THE CHICK EI4BR
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vt I I 6.3.2 Stage 10 Experímentel
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9 APPENDTCES 378 APPENDIX A . 379 1
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1.1 TERATOLOGY Birth defects have a
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4 of the complexity of normal devel
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6 flat, exposed plaque of neural ti
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8 However, several early embryos in
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.1- 5
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l2 2.1 ErI0L0rL!! fïE qYs34t!1!!jr
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4 which has rema¡ned fairly consta
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6 and about 112 after two (Laure,nc
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l8 ln some very early human embryos
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2l and kinky-ta¡l as disorders of
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23 Vogel and McClenahan (1952) tnus
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I'IATE R IALS 25
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27 lf necessary, eggs were stored a
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FÍg. 5. lncubator allowing precise
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3l 4. r sELEcr I oN !F EGGq 4.1,1 .
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uler. Accurate Staging (Hamburger a
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.\lz '?. t
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Fis. I AsB, Windowìng followed by
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Fis. 9 Aã8. t/indow?ng followed by
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42 ( \--l /) ^ ) t0
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44 serîal sectlonlng a camera luci
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46 Thus the embryos selected for se
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48 TABLE 3. srAGE t0 El,tBRyos (cno
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50 TABLE 5 , Reg i ons srAGE 13:16'
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52 4.8.4 HistologÌcal_De:plþtÍon
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5 RESULTS OF TERAÎOLOGICAL PROCEDU
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(c) deaths and defects in the survi
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Fi g. Percentagcs of early deaths a
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6o MoRTALtTY, AFTER ì,'tNpowtNq,AT
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TABLE B.'DEVEIOPI{ENT'AFTER'WINDOIi
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TABLE 9. ¡4ALFORMATIONS PRODUCED'B
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N=231 ffil e ooyi ffi s oays [il t2
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6B TABLEt loj lloRTALtTy AFTER VtBR
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70 5,3.2 Pa-rôf i Idì ¿irid PIè
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TABLE 13; MALFORI4ATIONS AFTER PLAS
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41 13 0 23 No l^l i ndow Control s
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Fíg. Percentages of early deaths a
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78 ALBUMEN INTRODUCTION Analysís o
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0 1 0 0 a o TABLE 17. I'lALF0RÌ1AT
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N Eorly Deothi N=10ó N a ú, o d,
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103 51 52 28 14 4 6 No W î ndow Co
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Fi g. 14. Percentages of early deat
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88 AtR qELL, REEXPANSI0N Analysls o
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90 5.4 BACTERt4L CUTTURE 0F UlNpohr
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92 TABLE 22. ¡¡UI'ISELTS OF PLATE
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94 6.1 EMBRY0ûIIES I s 0L 0!!\- NE
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Fis. I5 A¿8, Range of s izes (mm.)
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Figs. 16 AaB.. Range of sizes (mm.
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100 6.1.2 f4ortal í!y with Vsrying
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421.. 26 25 I (3.85) 20 (76 .92) 1
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00 N Eorly Deoths ,n -¡ F- ô- xu.
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ì06 Degrees of Freedom 12 Chi Squa
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hzE 53 42C z5 51(96.4)2\(96) 2(3.77
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ffil Op"n Anterior Neuropore I I0 N
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N=380 El Ovol Rhonnboid 'Sinus ì l
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Øl Open Broin Defecls N=384 ffi F"
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ligr. 21 - 30. Camera lucida drawin
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Fi s. 22. lrregular open anteríor
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lmm OPEN BRAIN DEFECT Á,NOPHTHALMI
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lmm OVAL RHOMBOID S¡NUS 48 HOURS S
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FiS. 26, Open neural folds extendin
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t-l lmm IATER OPEN CORD DEFECT . RE
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l2B t--l lmm EARLY OPEN CORD DEFECT
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Fis. 30. Large caudal cyst Ìn an o
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132 6.1 .4 DeveloÞñent of oÞen N
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Fig. 31 . Percentêges of experinen
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36 TABLE ?8. RH0I4B0lD stNUs ctôs
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N=38o H ovol Rhomboid sinus l^ r38
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F¡ S. 33. Percentages Õf open cor
Page 157 and 158:
142 6.1.5 Distríbution of 0pen Cor
Page 159 and 160:
Fi s. "l! Percentage distrìbution
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lrregul ar (Z) somi te be low level
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ì48 N=2ó4 ffi m Regulor Cord Defe
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Fis. 36. Percentage distrîbution o
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152 6.2 sPtIA! LFVELS 0F oPEN CORp
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Fìgs. 37'38, Vertebral defecrs in
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CERVICAT THORACIC LUMBAR FUSED CAUD
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---l r r'I * 40
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t59 a) spîna bif¡da occulta is se
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:al . '**Ç- 4 ff, À(,) ftø 9-' o
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162 lmnediately after closure, sepa
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164 Hensenrs node and the primitive
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-l :: l 48 I 49 i :. t: .lì 50 ì:
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t67 somite area (through asymmetry
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t69 ln both embryos the volume of n
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171 The rhomblc roof became membran
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173 At areas of myeloschisîs conta
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175 neurêl tîssue wíth mesoderm
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t77 e) encroachment of somitîc mes
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ôt rf) , ,,. ì _ ^iiir¡:ù\. I r
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l 55 56 57 58 59 ó0
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Fiss. 65 Early myeloschisis in a St
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fiSr. 71 - 76. Later myeloschisis i
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Fiss. 77 -82. Early myelodysplesia
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FÌgs. B3 - BB. Later myelodysplas
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Figs, 89 - 91. Processíng artifact
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ì86 6.3.10 SequenJ¡¿il ll lúSti
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Fígs. 95 - 102, Sequentiêl drawin
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l9o EVERsToN: srAGE rf (o r o+) o @
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@ @@ "@ @ e q q a a 192 CoNTROL: ST
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MYELOSCHTSTS: STAGE tZ (tZeïl OOG
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coNTRO[: STAGE tó (ro c zz) . a@ Q
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G (õ--==: \z-- HEMTMYELIA: STAGE t
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Som i te Defects contact none sePa
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Somite Som i te Contact Defects N'
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Som i te Contact Som i te Defects c
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none none N) o o\ B r8c 23 12 good
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separat ion none contact/ none sepa
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Regíon Embryo Stage Conditîon Neu
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none none none none none none l'.J
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18E 10 13- vëry poor closed 0 cove
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separãtion none contact none conta
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Regíon Embryo Stage Condition Neur
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separation none sepâ ra t îon non
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separâtion none sepa ra t ion sepa
Page 250 and 251:
Reglon Embryo Stagd cond¡t¡on Neu
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none none none none N) NJ .{ A A A
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contact/ none separatlon contact no
Page 256 and 257:
Region Embryo Stage Conditlon Neura
Page 258 and 259:
TABLE 38D. STAOE 17.20 CONTROL AND
Page 260 and 261:
Reglon Embryo Stage Condition Neura
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428 72 20 closed covered none conta
Page 264 and 265:
239 neural tissue ât the later sta
Page 266 and 267:
241 The control group (Table 99 ) s
Page 268 and 269:
TABLE 39. APPEARANCE OF I,'HOLE EI4
Page 270 and 271:
TABLE 40. APPEARANCE OF }/HOLE EI"I
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closed closed c I osed c I osed c I
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249 6.6 pEVEL0PMENT 0F rH!_¡Ho|4gl
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Emb ryo Stage Rhombîc Roóf Neural
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CDE D'E DE CDE DE DE DE D BCD BC BC
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TABLE q6,. STAGE 17-20 CoNTRoI AND
Page 282 and 283:
Figs- 10J - i11. Development of the
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Fî s. tub. Thick rhombic roof in 5
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Fig. 109 Hernbranous rhombic roof w
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260 6.7 HrqroLoctcAL, cHANgË Assoc
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TABLE 47. CONTROL EMBRYoS.HISTOLOGI
Page 292 and 293:
0 0 0 0 0 0 0 0 0 0 o 0 0 0 0 0 0 N
Page 294 and 295:
42c 21 zo B c D E 0000 00 00 00 1\.
Page 296 and 297:
l\t q\ 1oE 26 16 30E 59 16 3oE 77 1
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0 0 0 0 0 0 0 0 0 0 0 0 N) .{ Embry
Page 300 and 301:
l\) \l N D E 00 00 0 0 13.79 0 100
Page 302 and 303:
N) \¡ \2E 72 20 E B c D' E 00 0 15
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46. 15 76.92 0 20.51 14.83 2i,92 53
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ECTODERM DISCONTINUITY IN EXPERIMEN
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279 ECTODERM DISCONTINUITY IN EXPER
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SOMITE SEPARATION IN EXPERIMENTAL E
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283 NOTOCHORD SEPARATION IN EXPERIM
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285 SOMITE DEFECTS IN EXPERIMENTAL
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287 and distr¡bfrtíon in the cont
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19.09 r8.95 r 1.84 14.82 15.20 0 17
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0 15. 89 81 .08 100 100 ' 76.92 0 1
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43. rB 100 100 0 12.53 74.07 r00 't
Page 324 and 325:
7.t6 16.03 100 100 160 7.90 18.22 r
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Figs. 120'123. Percentage ie.ngths
Page 328 and 329:
299 O\TERLAP ZONE IN EXPERIMENTAL E
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301 OVERLAP ZONE IN EXPERIMENTAL EM
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303 ând every tenth sectlon was ne
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30c 12 16 c 17707,61 2463.3\ 7.19 D
Page 336 and 337:
Ratio NT/N 4 .90 4.11 6.17 7.14 6.9
Page 338 and 339:
Rêtlo NT/N 4.go \.36 3 .44 3. 18 3
Page 340 and 341:
6.7t 6.lt 4.63 4.60 4.13 3.48 hzE 5
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Fi9s, 124-127, Neurál tube-norocho
Page 344 and 345:
315 NEURAL TUBE -NOTOCHORD RATIOS I
Page 346 and 347:
317 NEURAL TUBE-NOTOCHORD RATIOS IN
Page 348 and 349: 319 TABLE 598 ' ANALYSIS OF VARIANC
Page 350 and 351: 321 TABLE 61., .NEURAL TISSUE/NOTOC
Page 352 and 353: 323 Apart f.rom the sîgnîficant d
Page 354 and 355: t25 7. Dlscusst0N 1'/i th progressi
Page 356 and 357: 327 C¡osure of the anterior neurop
Page 358 and 359: 329 the mid-polnts of the defects w
Page 360 and 361: 331 'c¡osure of the neural folds w
Page 362 and 363: 333 . These findings s.ugges r that
Page 364 and 365: 335 Lendon (1968, 1975) and notos (
Page 366 and 367: 337 Ferm, 1965; l"larin-Padi r ¡a,
Page 368 and 369: 339 Kôplan (1965) and Kaplan and G
Page 370 and 371: 341 reach the floor-plate; asymmetr
Page 372 and 373: 343 took up the label, shovri.ng th
Page 374 and 375: 345 The defects in early human embr
Page 376 and 377: 347 after pred¡cted closure (Lemî
Page 378 and 379: 349 hours could provide evidence fo
Page 380 and 381: 351 t971), Gal lus and Xenopus (l'l
Page 382 and 383: 353 1962i 1965). Barth and Barth.(1
Page 384 and 385: The result of any teratogen¡c insu
Page 386 and 387: 357 Uindowîlg at ear¡y stages of
Page 388 and 389: SUHHARY AND CONCLUS I ONS
Page 390 and 391: 361 separatlon between the neural p
Page 392 and 393: 363 APPENDIX A Preparatlon of Early
Page 394 and 395: 365 APPENDIX B Stainl.ng of Carti l
Page 396 and 397: 367 10, BIBLIOGRAPHY, Alter, l.,l.
Page 400 and 401: 371 Crlley, B,B.(1969). Analysîs o
Page 402 and 403: 373 dqs anomal ies de lrorganizatio
Page 404 and 405: 375 Heath, H.D., Shear, H.H., lm€
Page 406 and 407: 377 Jel inek, R. (1960). Developmen
Page 408 and 409: 379 Laurence, K.14. (1964). Thé na
Page 410 and 411: 381 McKeown, T. and Record, R.G.(19
Page 412 and 413: Penrose, L. S.(1957). Genetics of a
Page 414 and 415: 385 Russell, H.E. and A¡ tken' G.T
Page 416 and 417: 387 Spemann, H. (1938). Embryonic D
Page 418 and 419: 389 lJeed, ¡.H. (1937 -38'). M.nin
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