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G (õ--==: \z-- HEMTMYELIA: STAGE tA (+ZESO) ç--æ -@ é @ @ e @ @ e @a @o eo e€ @€ @ @e l9B øe - ç. ee Pv ps Øs q a g ø g , , t'e-, q 9 g o
6.4w t99 The hlstologlcal study of neuruiation (Section 6.3.9) revealed certaln dlfferences between experímentar and contror embryos. The changes in the representatlve appearance of each region in every embryo are shown ln Tables 35 -38,. The assessment of the separation of neural tìssue from adjacent nobchord or somites was complicated by artifactual splittíng in some embryos. However, when experîmental (g) and control (C) embryos were compared in groups, some consîstent differences were evident. Each regíon of each embryo is presented cnanío-caudal ly, so that more than one description recorded for a region indicates a change in the appearânce of that region from above downwards (Tables 35 _ 3g ). Regions A,B,C,D and E of each embryo are described in terms of: (a) cond¡tion of embryo after processing (b) progress of neural closure (c) number of accessory canals (d) morphology of neural defects (") cover of neural tissue by ectoderm (f) cover of neural tissue by mesenchyme (neurar crest or somitic mesoderm) (s) contact.of notochord with neural tissue (h) contact of somitic mesoderm with neural tissue (l) abnormal ities of somitîc nesoderm.
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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
Page 157 and 158:
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
Page 173 and 174: ---l r r'I * 40
Page 175 and 176: t59 a) spîna bif¡da occulta is se
Page 177 and 178: :al . '**Ç- 4 ff, À(,) ftø 9-' o
Page 179 and 180: 162 lmnediately after closure, sepa
Page 181 and 182: 164 Hensenrs node and the primitive
Page 183 and 184: -l :: l 48 I 49 i :. t: .lì 50 ì:
Page 185 and 186: t67 somite area (through asymmetry
Page 187 and 188: t69 ln both embryos the volume of n
Page 189 and 190: 171 The rhomblc roof became membran
Page 191 and 192: 173 At areas of myeloschisîs conta
Page 193 and 194: 175 neurêl tîssue wíth mesoderm
Page 195 and 196: t77 e) encroachment of somitîc mes
Page 197 and 198: ôt rf) , ,,. ì _ ^iiir¡:ù\. I r
Page 199 and 200: l 55 56 57 58 59 ó0
Page 202 and 203: Fiss. 65 Early myeloschisis in a St
Page 204 and 205: fiSr. 71 - 76. Later myeloschisis i
Page 206 and 207: Fiss. 77 -82. Early myelodysplesia
Page 208 and 209: FÌgs. B3 - BB. Later myelodysplas
Page 210 and 211: Figs, 89 - 91. Processíng artifact
Page 212 and 213: ì86 6.3.10 SequenJ¡¿il ll lúSti
Page 214 and 215: Fígs. 95 - 102, Sequentiêl drawin
Page 216 and 217: l9o EVERsToN: srAGE rf (o r o+) o @
Page 218 and 219: @ @@ "@ @ e q q a a 192 CoNTROL: ST
Page 220 and 221: MYELOSCHTSTS: STAGE tZ (tZeïl OOG
Page 222 and 223: coNTRO[: STAGE tó (ro c zz) . a@ Q
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
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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
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0 0 0 0 0 0 0 0 0 0 o 0 0 0 0 0 0 N
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42c 21 zo B c D E 0000 00 00 00 1\.
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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
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l\) \l N D E 00 00 0 0 13.79 0 100
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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
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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
Page 344 and 345:
315 NEURAL TUBE -NOTOCHORD RATIOS I
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317 NEURAL TUBE-NOTOCHORD RATIOS IN
Page 348 and 349:
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
Page 364 and 365:
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
Page 380 and 381:
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
Page 386 and 387:
357 Uindowîlg at ear¡y stages of
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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
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365 APPENDIX B Stainl.ng of Carti l
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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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