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cranlum bifidum thus involves the establ ishment of a regular serles of speclmens between pre-neurulation and Iate fetal stages. 1.8 PROGNOS tS The current emphasis on early surgical correct¡on, facilitated by the development of antibiotics and the control led drainage of hydrocephal ic fluîd, l^:s greatly increased the survivar of dysraphic infants (Sharrard et al., 1967) . However where neural tissue ís involved the damage is irreversible, so that anencephaly is always fatal vrhí le myelcmenlngocele invariably produces some degree of neurological impairment, The survival of an increasing number of affected infants poses serrous social and economic problems (Tizard, l968; Lîghtowler, 1971). These wîll only be avoíded when the dysraphic states themselves are preventable through a fuller understanding of their etio¡ogy.
Page 1 and 2: E}4BRYOGENES IS OF EXPERIHENTALLY I
Page 3 and 4: MY PARENTS c.J.H. G. M.
Page 5 and 6: lt Congenital malformêtlons of the
Page 7 and 8: TABLE OF CONTENTS SECT ION PAGE 1 I
Page 9 and 10: },I.ATE R I ALS 3.1 THE CHICK EI4BR
Page 11 and 12: vt I I 6.3.2 Stage 10 Experímentel
Page 13 and 14: 9 APPENDTCES 378 APPENDIX A . 379 1
Page 15 and 16: 1.1 TERATOLOGY Birth defects have a
Page 17 and 18: 4 of the complexity of normal devel
Page 19 and 20: 6 flat, exposed plaque of neural ti
Page 21: 8 However, several early embryos in
Page 25 and 26: REV IEW OF L ITERATURE
Page 27 and 28: t3 The value of such col lected ser
Page 29 and 30: t5 ln general the lncldence of dysr
Page 31 and 32: t7 factors êre suggested by the co
Page 33: 9 TABLE l._ EXPERTHENTAL pySRApH_t
Page 36 and 37: 22 suggest¡ng that dysreph¡sm res
Page 38 and 39: 24 2,4.10 lnfect ion FInally, some
Page 40 and 41: 26 3.1 THE CHTCK EMBRYo The chlck e
Page 42 and 43: 2B To expose each embryo a wlndow w
Page 45 and 46: GENERAL I.,IETHODS
Page 47 and 48: 7.2 untreated embryos showed almost
Page 49 and 50: Ëigs. t4. Human neurcr*sp ina I d)
Page 51 and 52: JO ,\.3 REIN!UBATION AFTER. li,INDO
Page 53: IÆ 6
Page 56 and 57: 4ì ,'ä: 1 Æ(o) \/u I
Page 58 and 59: 43 \,\.3 Effect of Artìficial Aìr
Page 60 and 61: 4S or 22 KOH. Three types of skelet
Page 62 and 63: 47 I'leurulation showed a comparabl
Page 64 and 65: 49 TABLE 4. . .STAGE. t 1:l2. .Etrt
Page 66 and 67: 5l TABLE 6, % STAGE lT-20 EHBRyos l
Page 68 and 69: 53 Because of the difficulty in sep
Page 70 and 71: 55 5.1 TERAToGEN!! !FFECT 0F,WtNp0I
Page 72 and 73:
Numbers of Eggs 31 73 31 135 47 Eâ
Page 74 and 75:
14 HRS" 2ó HRS. 38 HRS. CONTROLS :
Page 76 and 77:
6t 0n examlning. the survivi.ng emb
Page 78 and 79:
63 5.2 MALFoRMAT I 0NS .P¡OpUCEp B
Page 80 and 81:
Fis. Percentages of neural arrd non
Page 82 and 83:
67 5.3 INVFSïIGATtON 0F THE TERATo
Page 84 and 85:
69 ÏABLE 1 I. MALFORMATIONS AFTER'
Page 86 and 87:
71 TABLE I2;},IORTALITY AFTE-R PLAS
Page 88 and 89:
73 5.3.3 .o¡literation,lof lt¡tro
Page 90 and 91:
No t'i i ndow Controls 0 0 0 1 0 0
Page 92 and 93:
N Eorly Deoths N=100 .n 9óo d co =
Page 94 and 95:
F 12 I'IEDIUH AT VARIOUS INTERVALS
Page 96 and 97:
Fis. 13. Percentages of early death
Page 98 and 99:
B3 Fl2 INTR0DUCTtqN Analysis of ear
Page 100 and 101:
No Wi ndow Control s 0 0 0 0 0 1 0
Page 102 and 103:
NOT REEXP. 50 HRS. 38 ¡-IRS. 2ó H
Page 104 and 105:
Ro Comparison of Figs. 12, lJ and 1
Page 106 and 107:
9l TABLE 2P" MORTALITY AFTER Ii'IND
Page 108 and 109:
6 RESULTS OF EMBRYOLOG I cAL STUD I
Page 110 and 111:
95 TABLE 2?, SI,ZES AND STAGES OF D
Page 112 and 113:
97 N=234 ttt o d, ao ã t¡¡ llo b
Page 114 and 115:
N=52 56789r0 STAGES (H.&H.) rN GROU
Page 116 and 117:
\28 73 20(27 .40) 53 21 (28.77') 25
Page 118 and 119:
Fig. 17. Percenteges of early death
Page 120 and 121:
05 I,IORTALITY O; 6; IB: 30 Ê.42'I
Page 122 and 123:
107 6.1.3 l:legral Cþ;ure and Negl
Page 124 and 125:
Fig' 18. Percentages of experimenta
Page 126 and 127:
Fi g' 19. Percentages of experiment
Page 128 and 129:
lig. 20. Development of open brain
Page 130 and 131:
I l5 l'/hen neural closure was asse
Page 132 and 133:
117 lmm AT WINDOWING 2ó-30 HOURS S
Page 134 and 135:
lmm OPEN ANTERIOR NEUROPORE TRUNK C
Page 136 and 137:
Fis. 2l+ . Normal oval shape of the
Page 138 and 139:
123 t1 lmm TRIANGUTAR RHOMBOID SINU
Page 140 and 141:
t--J lmm EARLY OPEN CORD DEFECT REG
Page 142 and 143:
Fìs. 28. lrregular neural folds wi
Page 144 and 145:
129 lmm LA,TEROPEN CORDDEFECT ¡RRE
Page 146 and 147:
3ì lmm CAUDAL CYST óO HOURS STAGE
Page 148 and 149:
133 TABLE 27, ANTERIOR NEUROPORE CL
Page 150 and 151:
ffi Opàn Anterior Neuropore 135 I
Page 152 and 153:
Fì s. 32. Percentages of experImen
Page 154 and 155:
t39 DEFECTS BY STAGES Experimentals
Page 156 and 157:
4l N=38o ffi! Regulcr Cord ffil lrr
Page 158 and 159:
143 TR IBUTION OF D DEFECT5 N umbe
Page 160 and 161:
145 ffit Regulor Cord Defects ñ=2
Page 162 and 163:
Fis. 35. Percentage d¡stribution o
Page 164 and 165:
149 TABLE 32. SOI4ITE LEVELS OF l4I
Page 166 and 167:
SOMITE LEVELS OF MID-POINTS OF OPEN
Page 168 and 169:
t53 TABLE- 33,.VER,TE-BRAL qEFqcTs
Page 170 and 171:
CERVICAL THORACIC LUMBAR FUSED CAUD
Page 172 and 173:
Figs. 39 - 42. Malformations ¡n ex
Page 174 and 175:
ì58 One embryo shows ðn open brai
Page 176 and 177:
FÌgs. 43 - 46, Vertebral defects i
Page 178 and 179:
t6l 6.3 DESCRIPTION OF HISTOLOGICAL
Page 180 and 181:
t63 j) ênd prom¡nence of Hensenrs
Page 182 and 183:
Figs. l+7 ^ 5A, Ëversic¡n of the
Page 184 and 185:
166 6.3.5 staEe 13-16 córitról Em
Page 186 and 187:
t68 with lack of contact in severar
Page 188 and 189:
170 These two lesions were clearly
Page 190 and 191:
172 The centraì part of each defec
Page 192 and 193:
t74 contour and an Irreguìar centr
Page 194 and 195:
t76 the lesions. The interpretation
Page 196 and 197:
Figs. 51 - 54, Normai de'¡elopment
Page 198 and 199:
Fí9s" 55 - 60. llcrmaI development
Page 200:
F;gs 51 - 64. Normal devcìopn¿¡i
Page 203 and 204:
: ó5 66 67 69 70
Page 205 and 206:
\l\) ìr,jtr, r: : :;r:,irrliirqn ^
Page 207 and 208:
: F- 77
Page 209 and 210:
''i l.l
Page 211 and 212:
B9 90
Page 213 and 214:
187 overl îes three accessory cana
Page 215 and 216:
189 GoNTROL: STAGE ll+ (te c a) O G
Page 217 and 218:
Fïgs. 97 " 99. Development of m)/e
Page 219 and 220:
193 MYELOSCHTSIS:STAGE ra.(re rea)
Page 221 and 222:
Figs. 100 - 102, Development of mye
Page 223 and 224:
HEMTMYELTA: STAGE ló (SO r ZA) @ q
Page 225 and 226:
6.4w t99 The hlstologlcal study of
Page 227 and 228:
contact none contact none contact n
Page 229 and 230:
none none N) o 6c 21 0c 46 10 10 go
Page 231 and 232:
Som i te Defects contact none cont
Page 233 and 234:
contáct contact con tâc t contêc
Page 235 and 236:
Son i te Con têct sepa ra t ¡on s
Page 237 and 238:
none none none none none none none
Page 239 and 240:
contact con tac t contact con tact
Page 241 and 242:
Som i te Con tact Som i te Defects
Page 243 and 244:
sepa ra t lon none contact/ cys ts
Page 245 and 246:
N) N) o r8E 28 14 very poor 18E 36
Page 247 and 248:
Som i te Som i te Contact Defects s
Page 249 and 250:
sePârat Îon sepa ra t ¡on separa
Page 251 and 252:
none none none none none none none
Page 253 and 254:
Region Embryo Stage Condltîon Neur
Page 255 and 256:
none none 428 17 zo c I osed covere
Page 257 and 258:
contact none contact/ none separat
Page 259 and 260:
contact cys ts separation none sepa
Page 261 and 262:
contact cysts sepãration none sepa
Page 263 and 264:
238 This revlew of the histologlcal
Page 265 and 266:
240 The histological features selec
Page 267 and 268:
242 (d) an lrregular neural defect
Page 269 and 270:
c I osed closed closed c I osed NJ
Page 271 and 272:
TABLE 41. APPEARANCE OF ì^IHOLE EM
Page 273 and 274:
TABLE 42. APPEARANCE OF I,'HOLE EMB
Page 275 and 276:
RHOMB IC ROOF DEl/ELOPI,IENT N' o E
Page 277 and 278:
Emb ryo Stage Rhombic Roof Neural D
Page 279 and 280:
B C DE BC 1..) 30E 76 16 very thin
Page 281 and 282:
CD BCD.E CD BCDE CD CD BCDE c NJ \
Page 283 and 284:
7- 4. 103 r05
Page 285 and 286:
.t l l]] 80t ¿ot rfÞ'. !-d...---
Page 287 and 288:
'Ì 1ú 111
Page 289 and 290:
261 ln these tables the contrors an
Page 291 and 292:
0 0 0 .0 0 0 0 0 0 0 0 a 0 0 0 0 0
Page 293 and 294:
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Page 295 and 296:
00 0 0 0 0 0 0 0 0 0 0 0 0 0 N) o\
Page 297 and 298:
N o\ \o DO 0 0 65.38 0 EO 0 0 100 0
Page 299 and 300:
0 2.67 0. N' ! 18E 36 14+ DE 22.22
Page 301 and 302:
N) .{ \28 1 t8 42Ê 44 18 \zE 54 t8
Page 303 and 304:
16.76 13.52 0 29.57 100 0 14.04 49.
Page 305 and 306:
Figs. 112-119. Percentage lengths o
Page 307 and 308:
ECTODERM DISCONTINUITY IN EXPERIMEN
Page 309 and 310:
SOMITE SEPARATION IN EXPERIMENTAL E
Page 311 and 312:
NgTOCHORD SEPARATTON rN EXPERTMENTA
Page 313 and 314:
REGION E N=|9 SOMITE DEFECTS IN EXP
Page 315 and 316:
286 6.8. EXTENT OF THE OVERLAP ZONE
Page 317 and 318:
100 o 20.79 47.50 100 ,N' oo 100 æ
Page 319 and 320:
t4.09 13.94 0 1).59 100 100 88. 89
Page 321 and 322:
13.85 10. 84 14'5 t 15.97 11.06 1.5
Page 323 and 324:
0 12.18 62.07 100 77.78 o t3.72 78.
Page 325 and 326:
Embryo Stage Type of Regîons of Re
Page 327 and 328:
298 OVERLAP ZONE IN CONTROL EMBRYOS
Page 329 and 330:
OVERLAP ZONE IN EXPERIMENTAL EMBRYO
Page 331 and 332:
302 6.9 ANALYSIS OF NEURAT VOTUHES
Page 333 and 334:
Rat io NT/N 6.53 6.zB 6.78 5.02 8.2
Page 335 and 336:
7.30 6,5\ o C'\ \2c 21 29763.8\ 258
Page 337 and 338:
3.97 4. B0 8. 3g 5.81 6.37 5.53 8.8
Page 339 and 340:
5.33 q.40 3.14 3.88 4.50 4.75 \.65
Page 341 and 342:
(u2 ) Ratio NT/N 1.72 2,33 2.19 2.7
Page 343 and 344:
314 NEURAL. TUBE -NOTOCHORD RATIOS
Page 345 and 346:
316 NEURAL TUBE_NOTOCHORD RAÏIOS I
Page 347 and 348:
3t8 Figures fl:4-127 ' demons t ra
Page 349 and 350:
adjustment for nominal alpha level
Page 351 and 352:
322 TABLE .64. ¡leunn|ilssue/noroc
Page 353 and 354:
Dlscusst0N 324
Page 355 and 356:
326 Spontaneous neural defects have
Page 357 and 358:
328 caudal regions. rrregurar or de
Page 359 and 360:
330 defects seen ln fowls, human sa
Page 361 and 362:
332 lrnplicated in the pathogenesls
Page 363 and 364:
334 et al. 1961). tríth the single
Page 365 and 366:
336 The inpresslon of somlte separ
Page 367 and 368:
338 Embryos wîth myelodysplasia ho
Page 369 and 370:
340 ectoderm and neuraì tlssue ln
Page 371 and 372:
3\2 marked reduction in myerodyspra
Page 373 and 374:
344 and .the rhombo¡d s f nus at t
Page 375 and 376:
346 the braln wall were.due to mark
Page 377 and 378:
348 and Lassman, 1972). ln the sirn
Page 379 and 380:
350 of rapid ectodermal mitosis. Ho
Page 381 and 382:
352 liver, and muscle (but not gut
Page 383 and 384:
354 (Rose,nqu i s t,l !66) , and re
Page 385 and 386:
356 resu¡t ¡n only minor defects
Page 387 and 388:
358 braln and cord volume, and ofte
Page 389 and 390:
360 I sutlt'lARY ANp coNcLUsto]s 1.
Page 391 and 392:
APPEND I CES 362
Page 393 and 394:
364 1i. Dehydrate with changes of 8
Page 395 and 396:
366 t0 BIBLIOGRAPHY
Page 397 and 398:
368 Barson, A.J. (1920). Spina blfi
Page 399 and 400:
Butros' J. (1965). Action of hearr
Page 401 and 402:
372 Feeney., J. F. and Watterson, R
Page 403 and 404:
374 Grobstein, C. (1953) . orphogen
Page 405 and 406:
376 Anat. Rec. t 00 :745. Hughes, A
Page 407 and 408:
378 Karfunkel, P. .(1974). The mech
Page 409 and 410:
380 Llghtowler, C.D.R. .(1971). Men
Page 411 and 412:
382 Hoseley, H.R. (.l947). lnsul in
Page 413 and 414:
384 Robertson, G.G., I.,illiamson,
Page 415 and 416:
386 Schroeder, T.E. (1970) " Neurul
Page 417 and 418:
vJêdd¡ngton, C.H. (1932).. Exper
Page 419:
390 Zwllli.ng, E..(1942). Restituti
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