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39B<br />

REPROOUCTIVE BIOLOGY ANO EMBRYOLOGY OF CROCOOILIAN9<br />

evident <strong>and</strong> may be invaginating. The optic <strong>and</strong> otic vesicles are also obvious,<br />

but <strong>the</strong> brain is not yet regionalized. The optic pIacodes <strong>and</strong> vesicles<br />

are more obvious than <strong>the</strong> otic.<br />

Flexures <strong>and</strong> Rotation. The embryo lies at right angles to <strong>the</strong> yolk surface,<br />

that is, body torsion has not yet commenced.<br />

Blastopore <strong>and</strong> Primitive Streak. Obvious.<br />

Gut. The gut is incomplete caudally <strong>and</strong> open ventrally along its entire<br />

length.<br />

Somitomeres. Three pairs <strong>of</strong> cranial somitomeres are visible anterior to<br />

<strong>the</strong> otic vesicle.<br />

Notochord. The notochord is evident.<br />

Stage 2 [Fig. 1 S)<br />

Blastoderm. The dorsal surface <strong>of</strong> <strong>the</strong> blastoderm is attached to <strong>the</strong><br />

overlying shell membrane, <strong>and</strong> <strong>the</strong> embryo remains so te<strong>the</strong>red throughout<br />

subsequent development. Blood vessels are now visible; one pair<br />

emerges from <strong>the</strong> embryo at <strong>the</strong> caudal level <strong>of</strong> <strong>the</strong> heart, whereas ano<strong>the</strong>r<br />

(larger) pair runs down <strong>the</strong> lateral wall <strong>of</strong> <strong>the</strong> embryo to emerge at approximately<br />

<strong>the</strong> level <strong>of</strong> <strong>the</strong> 20th somite.<br />

Somites. 21-25 pairs (decreasing markedly in size caudally).<br />

Delineation <strong>of</strong> <strong>the</strong> Embryo. Dorsally <strong>the</strong> embryo is almost completely<br />

delineated except for a very small circular area at <strong>the</strong> extreme caudal tip.<br />

Ventrally <strong>the</strong> caudal <strong>and</strong> caudo-Iateral boundaries <strong>of</strong> <strong>the</strong> body wall have<br />

formed.<br />

Branchial Arches.<br />

visible.<br />

The 1st <strong>and</strong> 2nd arches <strong>and</strong> <strong>the</strong> 1st branchial cleft are<br />

Heart. An extra vertical loop has developed making a total <strong>of</strong> three<br />

loops. The heart lies in <strong>the</strong> midline <strong>of</strong> <strong>the</strong> embryo.<br />

Sensory Placodes, Pits, etc. The lens placode <strong>and</strong> optic cup are defined,<br />

<strong>and</strong> <strong>the</strong> otic pit is distinctly patent.<br />

Brain. Hindbrain discernible as a clear transparent region.<br />

Flexures <strong>and</strong> Rotation. The cranial end <strong>of</strong> <strong>the</strong> embryo is flexed at approximately<br />

<strong>the</strong> level <strong>of</strong> <strong>the</strong> heart with <strong>the</strong> head lying at approximately 45° to<br />

<strong>the</strong> plane <strong>of</strong> <strong>the</strong> body. No body torsion has occurred.<br />

Stage 3 [Fig. 1 S)<br />

Somites. 26-30 pairs.<br />

Delineation <strong>of</strong> <strong>the</strong> Embryo. Complete.<br />

Branchial Arches. Three branchial arches, <strong>the</strong> 1st branchial cleft, <strong>the</strong> 2nd<br />

branchial groove, <strong>and</strong> <strong>the</strong> branchial sinus are all present.<br />

Tail. Bud present, but lacks somites.<br />

Brain. Forebrain, midbrain, <strong>and</strong> hindbrain are now discernible, <strong>the</strong><br />

latter appearing distinctly transparent.<br />

Sensory Placodes, Pits, etc. The optic cup has an elongated horseshoe<br />

BTAGEB OF EMBRYONIC OEVELOPMENT [AFTER EGG LAYING)<br />

399<br />

shape, which extends below <strong>the</strong> lens vesicle onto <strong>the</strong> ro<strong>of</strong> <strong>of</strong> <strong>the</strong> primitive<br />

oronasal cavity.<br />

Blastopore <strong>and</strong> Primitive Streak. Not visible.<br />

Extra-embryonic Membranes. The amnion is attached ventrally to <strong>the</strong> lateral<br />

body walls, cranially to <strong>the</strong> borders <strong>of</strong> <strong>the</strong> pericardium about <strong>the</strong> level<br />

<strong>of</strong> <strong>the</strong> 7th to 8th somite <strong>and</strong> caudally to <strong>the</strong> cranial margin <strong>of</strong> <strong>the</strong> fold <strong>of</strong> tail<br />

bud.<br />

Flexures <strong>and</strong> Rotation. Head at approximately right angles to <strong>the</strong> body.<br />

No body torsion.<br />

Stage 4 [Fig. 1 S)<br />

Somiles. 31-35 pairs. The first is beginning to disappear.<br />

Tail. Distinct, straight, <strong>and</strong> contains 3-5 somites in its base; <strong>the</strong> tip is<br />

unsegmented.<br />

Flexures <strong>and</strong> Rotation. Body torsion has commenced. The cranial half <strong>of</strong><br />

<strong>the</strong> embryo is rotated so that its right surface is in contact with <strong>the</strong> shell<br />

membrane <strong>and</strong> its left is parallel to <strong>the</strong> underlying yolk. The caudal half <strong>of</strong><br />

<strong>the</strong> embryo is not rotated <strong>and</strong> lies at right angles to <strong>the</strong> shell <strong>and</strong> yolk.<br />

Heart. Displaced to <strong>the</strong> left side <strong>of</strong> <strong>the</strong> embryo <strong>and</strong> large.<br />

Allantois. Small elevation is just visible immediately caudal to <strong>the</strong> craniallimit<br />

<strong>of</strong> <strong>the</strong> ventral tail fold, at approximately somite 27.<br />

Branchial Arches. Three branchial arches, <strong>the</strong> 1st branchial cleft, <strong>the</strong> 2nd<br />

<strong>and</strong> 3rd branchial grooves, <strong>and</strong> <strong>the</strong> branchial sinus are present. The cranial<br />

nerves to branchial arches 1-3 are discernible using oblique or transmitted<br />

illumination.<br />

Stage 5<br />

[Fig. 1 gJ<br />

Somites. 36-40 pairs. Only traces <strong>of</strong> <strong>the</strong> first somite can be detected,<br />

although it is included in this count.<br />

Tail. The tail tip bends ventrally at right angles to <strong>the</strong> body <strong>of</strong> <strong>the</strong><br />

embryo; 6-10 somites in its base, tip unsegmented.<br />

Flexures <strong>and</strong> Rotation. Body torsion is complete except for <strong>the</strong> tail. The<br />

head is fur<strong>the</strong>r flexed with <strong>the</strong> ro<strong>of</strong> <strong>of</strong> <strong>the</strong> brain at approximately 25° to<br />

<strong>the</strong> plane <strong>of</strong> <strong>the</strong> body.<br />

Allantois. The allantoic bud is distinctly swollen, smaller in height than<br />

<strong>the</strong> tail.<br />

Sensory Placodes, Pits, etc. The otic pit lies dorsal to <strong>the</strong> junction <strong>of</strong> <strong>the</strong><br />

2nd <strong>and</strong> 3rd branchial arches, <strong>and</strong> its external opening is closing.<br />

Stage S<br />

[Figs. 1 S. 1 g. <strong>and</strong> 23J<br />

Sensory Placodes, Pits, etc. Nasal placodes present. Otic pit closed.<br />

Limbs. Hindlimb buds are just visible on each side, <strong>the</strong> right hindlimb<br />

~ bud being marginally advanced over <strong>the</strong> left, but no forelimb buds are<br />

present (Fig. 23).

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