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249 6.6 pEVEL0PMENT 0F rH!_¡Ho|4glc R00F Changes in the structure of the rhornbic roof were also evaluated for Groups I - lV. They are presented, together w¡th the norphology and levels of neurâl defects (from Section 6.2), in Tables 43 - 46. and Flgs. 103- I11. Tables 43 - 46 show that the rhor¡rbic roof undergoes progressive th i nn îng after neural closure: Stages l0 - 11+ Stages 12 - 15 Stâges 16 - 17 Stages 18 - 20 thick thin very th¡n membranous The choroid plexus of the fourth ventricle is not present before Stage 18 (f¡S. lO9 ) considerably later than the first appeêrance of myeloschîsis and myelodysplasia. ìlîthin each g¡-oup there is no difference in rhombic development between experimental and control embryos. ln this ser¡es of chick embryos, the formatíon of open neural defects cannot be secondary to excessive pressure within the cerebro-spinal fluid system.
RHOMB IC ROOF DEl/ELOPI,IENT N' o Emb ryo Stage Rhombic Roof Neural Defects Regions of Defects 6c 20 t0- èlosíng none oc 49 t0- clos i ng none 0c 52 10- closing none 6c 21 t0 thick none 0c 46 10 closlng none 6E 15 6E8 t0- t0 closing thick none none 6E 30 6E \s t0 10 th ick thick none early myeloschisis E 6E 18 10+ thick none 6E 4r 10+ thlck early myeloschlsís E
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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
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142 6.1.5 Distríbution of 0pen Cor
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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
Page 224 and 225: G (õ--==: \z-- HEMTMYELIA: STAGE t
Page 226 and 227: Som i te Defects contact none sePa
Page 228 and 229: Somite Som i te Contact Defects N'
Page 230 and 231: Som i te Contact Som i te Defects c
Page 232 and 233: none none N) o o\ B r8c 23 12 good
Page 234 and 235: separat ion none contact/ none sepa
Page 236 and 237: Regíon Embryo Stage Conditîon Neu
Page 238 and 239: none none none none none none l'.J
Page 240 and 241: 18E 10 13- vëry poor closed 0 cove
Page 242 and 243: separãtion none contact none conta
Page 244 and 245: Regíon Embryo Stage Condition Neur
Page 246 and 247: separation none sepâ ra t îon non
Page 248 and 249: separâtion none sepa ra t ion sepa
Page 250 and 251: Reglon Embryo Stagd cond¡t¡on Neu
Page 252 and 253: none none none none N) NJ .{ A A A
Page 254 and 255: 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
Page 262 and 263: 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
Page 272 and 273: closed closed c I osed c I osed c I
Page 276 and 277: Emb ryo Stage Rhombîc Roóf Neural
Page 278 and 279: CDE D'E DE CDE DE DE DE D BCD BC BC
Page 280 and 281: TABLE q6,. STAGE 17-20 CoNTRoI AND
Page 282 and 283: Figs- 10J - i11. Development of the
Page 284 and 285: Fî s. tub. Thick rhombic roof in 5
Page 286 and 287: Fig. 109 Hernbranous rhombic roof w
Page 288 and 289: 260 6.7 HrqroLoctcAL, cHANgË Assoc
Page 290 and 291: 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
Page 298 and 299: 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
Page 304 and 305: 46. 15 76.92 0 20.51 14.83 2i,92 53
Page 306 and 307: ECTODERM DISCONTINUITY IN EXPERIMEN
Page 308 and 309: 279 ECTODERM DISCONTINUITY IN EXPER
Page 310 and 311: SOMITE SEPARATION IN EXPERIMENTAL E
Page 312 and 313: 283 NOTOCHORD SEPARATION IN EXPERIM
Page 314 and 315: 285 SOMITE DEFECTS IN EXPERIMENTAL
Page 316 and 317: 287 and distr¡bfrtíon in the cont
Page 318 and 319: 19.09 r8.95 r 1.84 14.82 15.20 0 17
Page 320 and 321: 0 15. 89 81 .08 100 100 ' 76.92 0 1
Page 322 and 323: 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
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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
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Ratio NT/N 4 .90 4.11 6.17 7.14 6.9
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Rêtlo NT/N 4.go \.36 3 .44 3. 18 3
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6.7t 6.lt 4.63 4.60 4.13 3.48 hzE 5
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Fi9s, 124-127, Neurál tube-norocho
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315 NEURAL TUBE -NOTOCHORD RATIOS I
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317 NEURAL TUBE-NOTOCHORD RATIOS IN
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319 TABLE 598 ' ANALYSIS OF VARIANC
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321 TABLE 61., .NEURAL TISSUE/NOTOC
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323 Apart f.rom the sîgnîficant d
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t25 7. Dlscusst0N 1'/i th progressi
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327 C¡osure of the anterior neurop
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329 the mid-polnts of the defects w
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331 'c¡osure of the neural folds w
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333 . These findings s.ugges r that
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335 Lendon (1968, 1975) and notos (
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337 Ferm, 1965; l"larin-Padi r ¡a,
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339 Kôplan (1965) and Kaplan and G
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341 reach the floor-plate; asymmetr
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343 took up the label, shovri.ng th
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345 The defects in early human embr
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347 after pred¡cted closure (Lemî
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349 hours could provide evidence fo
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351 t971), Gal lus and Xenopus (l'l
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353 1962i 1965). Barth and Barth.(1
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The result of any teratogen¡c insu
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357 Uindowîlg at ear¡y stages of
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SUHHARY AND CONCLUS I ONS
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361 separatlon between the neural p
Page 392 and 393:
363 APPENDIX A Preparatlon of Early
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365 APPENDIX B Stainl.ng of Carti l
Page 396 and 397:
367 10, BIBLIOGRAPHY, Alter, l.,l.
Page 398 and 399:
369 Braun, l'{. (1882). entwicklung
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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