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Blastula period begins when the cell reaches it 128-cell stage or after the 8th zygotic division.

In
this stage, the cell enters midblastula transition, forms yolk syncytial layer and begins epiboly.

Epiboly begins during the late blastula and it is the thinning and spreading of both the yolk
syncytial layer and the blastodisc over the cell.. During epiboly in the zebrafish, the margin
moves down to cover the yolk and at the end of gastrulation the cell becomes engulfed
completely.

Stages of zebrafish development are identified as percentage of epiboly which is the amount of
yolk surrounded at that time. Epiboly depends on the activity of microtubules within the yolk
syncytial layer and is disrupted by microtubule disrupting agents like nocodazole.

In the 50% epiboly, hypoblast, germ ring, and embryonic shield can be seen. While in the 75%
epiboly epiblast, hypoblast and evacuation zone can be ssen and the dorsal side of the cell is
thicker than the ventral side. On the other hand, in the 90% epiboly the tailbud can be seen.

During epiboly the blastodisc becomes a cup-shaped multilayer cell of almost uniform thickness
from being a cell mound. This occurs by the streaming of the deepest blastomeres outward and
toward the surface.

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During gastrulaton epiboly (A) continues while involution and convergent extension (B) occurs
to produce the primary germ layers. The term gastrulation is associated with the involution of
the mesoderm as distinct from the epibolic spreading process.

Involution is only carried out by the deep cells and outer enveloping layer cells do not
participate. The lack of mixing among the marginal cells may have significance in the
establishment of the pattern during early development because mesoderm will form from the
nonmixing marginal cells and is important to maintain regions with different cellular
specifications. Involution takes place around the blastoderm region but the dorsal involution is
more pronounced. Involution results in the formation of germ ring, a thickened marginal region
which can be observed during the onset of gastrulation at 50% epiboly.

The presence of the germ ring creates two germ layers, the epiblast, which is the upper layer
and the hypoblast, the lower layer.

Th epiblast continues to feed the cells into the hypoblast throughout gastrulation. The cells left
in the epiblast at the end of gastrulation correspond to the ectoderm which will give rise to the
epidermis, and other structures. The hypoblast then gives rise to both mesoderm and
endoderm.
Simultaneous with involution the convergence movement then produces a thickening of the
dorsal region of the germ ring which is call the embryonic shield. Epiboly temporarily arrest
until the shield is formed. The embryonic shield is comparable to the dorsal lip of Xenopus or
the node of amniotes.

As the involution proceeds, the shield elongates in an anteroposterior manner by convergent


extension. The driving force for convergent extension is an active process of cell intercalation in
a mediolateral direction. The Wnt-PCP pathway is necessary for convergent extension to
proceed.

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This figure shows the fate maps of zebrafish at different stages.

The fatemap for the zebrafish has been constructed by injection of blastomeres with
fluorescent dextrans. Fate maps from the cleavage stage are only statistical as there is
considerable mixing of deep cells. On the other hand, during gastrulation the range of random
cell movement is much reduced and a fate map can be produced. Cells may populate more than
one tissue type or germ layer if labelled in the early stage sof epiboly but by the start of
involution, all clones have been constricted to tissue types.