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55 5.1 TERAToGEN!! !FFECT 0F,WtNp0I{1!Nq Tt¡e effect of windowîng 999s in the early developmental period ls closely related to embryonic age. To învestigate this effect, the standard wlndowing. technîc was performed on eggs lncubated for l\, 26 and 38 hours. After windowi.ng, enbryos were reincubated to a tot¡l of /2 hours. Eggs ln the control group were not opened, but were removed from the lncubator for a simîlar length of time to the experimentâl groups. None of the eggs were turned. 4t.72 hours all the embryos were fixed and examined under a dissectlng mlcroscope. Table 7 and Fîg. l0 show the mortal lty and overall malformation rates for experrmentar embryos windowed at r4, 26, and l8 hours, together wlth the control embryos. Hany experlmental embryos, especially those windowed at l4 hours, showed early death and such complete degeneratìon that embryonic structures were.unrecognizab,le within the_smal I nodule of necrotic tissue. For this reason, individual malformation rates are not given as percentages of the number of embryos whìch continued to develop after windowing. survivî.ng embryos deveroped to stages 13-20 and were recovered dead or al ive, w¡th or without malformations, at 72 hours. Fig.l0 ¿s¡q¡strâtes a decrease in early deaths betv,reen the four groups, and.an increãse in survival wlthout defects âmong the embryos which contínued to develop. Statlstical analysls reveals slgnificant dlfferences in: (a) early deaths and survivrng embryos between the combrned experimenta¡ groups and the control group (p < O.Ol) (b) early deaths and survlving embryos in each experimentêl group (p < o.ot)
(c) deaths and defects in the surviving embryos of each experimental group (p < o.o5) . 56
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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
Page 19 and 20: 6 flat, exposed plaque of neural ti
Page 21 and 22: 8 However, several early embryos in
Page 24 and 25: .1- 5
Page 26 and 27: l2 2.1 ErI0L0rL!! fïE qYs34t!1!!jr
Page 28 and 29: 4 which has rema¡ned fairly consta
Page 30 and 31: 6 and about 112 after two (Laure,nc
Page 32 and 33: l8 ln some very early human embryos
Page 35 and 36: 2l and kinky-ta¡l as disorders of
Page 37 and 38: 23 Vogel and McClenahan (1952) tnus
Page 39 and 40: I'IATE R IALS 25
Page 41 and 42: 27 lf necessary, eggs were stored a
Page 43: FÍg. 5. lncubator allowing precise
Page 46 and 47: 3l 4. r sELEcr I oN !F EGGq 4.1,1 .
Page 48 and 49: uler. Accurate Staging (Hamburger a
Page 50 and 51: .\lz '?. t
Page 52 and 53: Fis. I AsB, Windowìng followed by
Page 55 and 56: Fis. 9 Aã8. t/indow?ng followed by
Page 57 and 58: 42 ( \--l /) ^ ) t0
Page 59 and 60: 44 serîal sectlonlng a camera luci
Page 61 and 62: 46 Thus the embryos selected for se
Page 63 and 64: 48 TABLE 3. srAGE t0 El,tBRyos (cno
Page 65 and 66: 50 TABLE 5 , Reg i ons srAGE 13:16'
Page 67 and 68: 52 4.8.4 HistologÌcal_De:plþtÍon
Page 69: 5 RESULTS OF TERAÎOLOGICAL PROCEDU
Page 73 and 74: Fi g. Percentagcs of early deaths a
Page 75 and 76: 6o MoRTALtTY, AFTER ì,'tNpowtNq,AT
Page 77 and 78: TABLE B.'DEVEIOPI{ENT'AFTER'WINDOIi
Page 79 and 80: TABLE 9. ¡4ALFORMATIONS PRODUCED'B
Page 81 and 82: N=231 ffil e ooyi ffi s oays [il t2
Page 83 and 84: 6B TABLEt loj lloRTALtTy AFTER VtBR
Page 85 and 86: 70 5,3.2 Pa-rôf i Idì ¿irid PIè
Page 87 and 88: TABLE 13; MALFORI4ATIONS AFTER PLAS
Page 89 and 90: 41 13 0 23 No l^l i ndow Control s
Page 91 and 92: Fíg. Percentages of early deaths a
Page 93 and 94: 78 ALBUMEN INTRODUCTION Analysís o
Page 95 and 96: 0 1 0 0 a o TABLE 17. I'lALF0RÌ1AT
Page 97 and 98: N Eorly Deothi N=10ó N a ú, o d,
Page 99 and 100: 103 51 52 28 14 4 6 No W î ndow Co
Page 101 and 102: Fi g. 14. Percentages of early deat
Page 103 and 104: 88 AtR qELL, REEXPANSI0N Analysls o
Page 105 and 106: 90 5.4 BACTERt4L CUTTURE 0F UlNpohr
Page 107 and 108: 92 TABLE 22. ¡¡UI'ISELTS OF PLATE
Page 109 and 110: 94 6.1 EMBRY0ûIIES I s 0L 0!!\- NE
Page 111 and 112: Fis. I5 A¿8, Range of s izes (mm.)
Page 113 and 114: Figs. 16 AaB.. Range of sizes (mm.
Page 115 and 116: 100 6.1.2 f4ortal í!y with Vsrying
Page 117 and 118: 421.. 26 25 I (3.85) 20 (76 .92) 1
Page 119 and 120: 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
Page 167 and 168:
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
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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
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TABLE 38D. STAOE 17.20 CONTROL AND
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Reglon Embryo Stage Condition Neura
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428 72 20 closed covered none conta
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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
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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
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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
Page 306 and 307:
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
Page 314 and 315:
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
Page 326 and 327:
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
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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
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
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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
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
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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
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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
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363 APPENDIX A Preparatlon of Early
Page 394 and 395:
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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