Early development Microlecithal egg
Unfertilized egg
The development of an organism involves an increase in mass and
elaboration of structures. Both these changes can be understood in
terms of cell activity, like an increase in cell number (due to mitosis),
enlargement of cells, migration of cells, as well as differentiation and
specialization of cells.
Cleavage is a mechanism for cloning the zygote genome generated
at fertilization following male and female pronuclear fusion. This
occurs right after fertilization, and it includes the zygote undergoing
a series of mitotic cell division known as segmentation or cleavage,
which leads to the formation of blastula. In animals where cleavage
(cell division) involves the whole egg, the blastula usually consists of
a hollow ball of cells. Gastrulation is the first time many organisms
begin to express the new embryonic genome that was formed at
fertilization.
Eggs are classified according to the relative amount of yolk which
they contain.
Those with little yolk are designated microlecithal or
oligolecithal Ex. Echinoderms, coelenterates, amphioxus
and mammals except monotremes
Those with moderate amount of yolk are called
medialecithal or mesolecithal. Ex. Eggs of annelid,
mollusks, lampreys, lungfishes and ampbihibians.
Those with large amount of yolk are called megalecithal or
macrolecithal. Ex. Arthropods, hag fishes, bony fishes,
reptiles, birds and monotremes
Those with even distribution of egg are called isolecithal.
Therefore, a microlecithal egg is isolecithal.
A telocithal egg on the other hand, has most of its yolk
concentrated on one pole the vegetal pole. Therefore,
amphibians are moderately telolecithal, and birds are
highly telolecithal. In arthropods, especially insects, the
yolk is concentrated in the interior of the egg, and the
cytoplasm is distributed as a thin coat on the external
surface. Eggs with this type of yolk distribution are called
centrolecithal eggs.
Cleavage
Cleavage is not simply multiplication of cells but rather it produces
cells which, eventually, become arranged distinctly. These cells are
called cleavage cells or blastomeres. This is due to the
predetermined pattern influenced by the amount and distribution of
the yolk among vertebrates. The yolk is very important factor in the
cleavage formation because its presence may inhibit cell division.
In eggs with small amount of egg or none at all (alecithal) and is
equally distributed throughout the embryo will be completely
divided into blastomeres of equal size. Those with medium amount
of yolk (mesolecithal or medialecithal), the yolk are pulled to its
lower position (the vegetal pole). The pole with more yolk will have
bigger and fewer blastomeres (macromeres) because given time, the
cells have not divided as often while the other pole with less yolk,
will have many small blastomeres (micromeres).
- The inactivated ovum of the starfish exhibits
prominent prematuration nucleus, the germinal
vesicle, with a distinct nucleolus. It also possess a
viteline membrane which is difficult to identify
because of its close adherence to the egg. This is
formed by the egg and is, therefore, a primary
membrane.
Fertilized egg
- After fertilization, the zygote shows an elevated
vitelline membrane, appropriately termed the
fertilization membrane. Because of the separation of
this structure from the egg, a space is formed, the
perivitelline space. The germinal vesicle of the
fertilized egg has broken down in preparation for the
maturation process. The sperm head may appear as a
densely staining body within the egg cortex.
Cleavage division stage
- The starfish egg exhibits holobastic radial cleavage.
Medialecithal egg
Amphibians typically exhibit mesolecithal eggs, having an
intermediate amount of yolk. The amphibian egg is larger
than that of the starfish due to the larger quantity of yolk.
It is distinctly divided into a dark pigmented hemisphere
(the animal hemisphere) and a lightly or unpigmented
hemisphere (the vegetal hemisphere). The pigement
granules reside in the cortical cytoplasm of the egg within
the plasma membrane and are more abundant in the
animal hemisphere. The yolk is distributed throughout the
cytoplasm of the egg but is much more concentrated
toward the vegetal pole end of the egg. Hence, the yolk
offers resistance to cleavage in the vegetal hemisphere.
The amount of yolk is still sufficiently small to allow
complete cleavage of the egg cell.
Blastula
Microlecithal egg
Blastula
- A hollow ball of cells resulting from the successive
cleavages.
o Blastocoel
- The centrally located cavity, the walls of which consist
an epithelial layer of cells. This is not readily
identifiable because the cells at the back part of the
section is still visible but the organization of the
blastomeres at the external surface of the embryo is
prominent.
o Blastoderm
- The layer of the blastomeres arranged at the side of
the embryo and enclosing the blastocoel.
o Fertilization membrane
- The thin membrane surrounding the blastula. It is
considered as a secondary membrane because it is
not secreted by the egg.
The animal and vegetal will not be identified for this specimen
because it is not easy to identify these in oligolecithal eggs because
the cells are almost of the same size.
Medialecithal egg
Blastula
- A hollow sphere resulting from the segmentation of
the zygote
o Animal pole
- The region where numerous smaller cells are found
o Vegetal pole
- The region where fewer but bigger cells are found.
o Blastocoel
- The eccentrically located cavity. It will be obliterated
in the future.
o Micromeres
- The cells at the animal pole.
o Macromeres
- The cells are the vegetal pole.
o Blastoderm
- Same description as in microlecithal egg.
o Fertilization Membrane
- The thin membrane surrounding the blastula. It is
considered as a secondary membrane because it is
not secreted by the egg but by the follicle cells in the
ovary. The chorion plus the vitelline membrane, if it
exists at all, makes up the fertilization membrane.
(This term is used to refer also to one of the fetal
membranes of the amniotes)
Gastrula
Gastrulation is a stage characterized by morphogenic movement
and well-ordered rearrangement of cells after the cleavage stage
and blastula stages. The gastrula that is formed is the embryo. This
formation occurs after invagination happens at the vegetal pole. It
involves 3 kinds of movements: epiboly, involution and
invagination. Groupings of cells of the same developmental
potentialities occur here. All these changes lead to the formation of
the 3 germ layers, namely the ectoderm, mesoderm and the
endoderm.
The start of gastrulation with the different kinds of embryo of
different cleavage patterns just studied also differs from each other.
In animals with microlecithal egg, gastrulation is simple the in
pocketing of the blastula with the formation of a double-layered cup.
In the mesolecithal egg, it starts with the formation of a dorsal
blastoporal lip within the grey crescent area, while that of the chick
and the placental mammals follow the same general pattern that is
the formation of a primitive streak.
Microlecithal egg
After the formation of a relatively large blastocoel, there is
plenty of space to move some of the surface cells to the
inside to form new layers. Gastrulation can now start. The
first sign of gastrulation in microlecithal eggs is first seen
as the starfish embryo becomes flattened at the vegetal
pole (the initial migration and differentiation of cells). The
cells will then invaginate within the blastocoel. When this
occurs, the internal layers of the cells are formed. This will
give rise to the endoderm and mesoderm germ layers. The
cells, which remain on the outer surface of the gastrula
become the ectoderm. The cavity created by the
invagination is called the gastrocoel or the archenteron
(future primitive gut), whose opening at the vegetal pole is
called the blastopore.
o Gastrocoel
- The new cavity that is formed by the invaginated
cells. It will become the digestive tract of the embryo.
The length of the gastrocoel indicates the specific stage of gastrula
whether early, middle or late. Early gastrula exhibits a short
gastrocoel whereas in late gastrula, the gastrocoel extends over half
the length of the embryo and its apex may be seen a thin-walled
vesicle.
o Vesicles
- Diverticula or lateral pouches found at the apical
region of the gastrocoel.
- They become separated from the gastrocoel and give
rise to the many mesodermal structures of the
animal.
- Syn: coelomic sacs
o Enterocoel
- The cavity formed from the outpocketings of the
gastrocoel
o Blastopore
- The opening of the gastrocoel to the outside. It marks
the posterior end of the embryo and becomes the
future anus.
o Mesenchyme
- Large, stellate cells which are mesodermal in origin.
- Scattered between the ectoderm and gastrocoel
- These cells arise by budding off from the outer walls
of the gastrocoel
o Bipinnaria larva
- The larva of the starfish that is formed at the end of
the gastrula stage.
- Noticeable is the presence of the cilia covering the
entire ectodermal surface of the embryo. These are
the principal locomotors organs of the larva.
Medialecithal egg
Gastrulation begins with the invagination of the advancing
germ ring cells and continues in a process called involution.
Involution is the in-turning or inward movement of the
outer layer of cells so that it spreads out under the internal
surface of the outer layer of cells. Cells are the margin of
the germ ring, well below the equator of the blastula,
change shape so that the surface layer turns inward and
then begins to migrate anteriorly along the root of the
blastocoel. This invagination of the migrating cells creates
the beginning of the archenteron. The mouth of the in-
pocketing is the blastopore and the upper margin of the
curved fold is known as the dorsal lip of the blastopore. As
the in-pocketing depends the opening extends
progressively into a crescent, then into a horseshow, and
finally into a complete circle. As the processes of
invagination and involution occur, epiboly continues as the
germ ring advances over the exposed yolk cells, so that by
 the time the circle is complete, the yolk can only be seen
within the confines of the blastopore, in an area known as
the yolk plug. The blastopore is now surrounded by dorsal,
lateral and ventral lips. Involution first occurred at the
dorsal lip near the more rapidly dividing animal pole. The
expanding layer of cells which forms the roof of the
archenteron becomes the endoderm and
chordamesoderm. For some time the floor of the
archenteron remains composed of large yolk cells. The cells
on the outer surface of the embryo now form the
ectoderm. As gastrulation continues involution begins to
occur at the ventral lip of the blastopore and gradually the
invaginated endoderm proliferates to enclose the yolk and
complete the archenteron. The third layer, the mesoderm,
soon develops between the ectoderm and the endoderm.
Gastrula
- Two-layered embryo resulting from the ectodermal
movements of the cell
o Gastrocoel/Archenteron, Blastopore
- As described in the previous section
o Lip of the blastopore
- The portion of the ectoderm where involution took
place
Dorsal lip the more defined margin of the
blastopore where involution is prominent
Ventral lip the less refined lower margin of the
blastopore marked by a deep cleft
o Yolk plug
- The portion of the vegetal pole that protrudes and
obstructs the blastoporal opening.
- endodermal in origin.
o Blastocoel
- Remnant of this may be seen.
o Fertilization membrane
- The thin membrane that envelopes the gastrula
toward the midline in the posterior half of the area
pellucida.
This movement of cells is homologous to the epiboly in amphibians.
The cells that sink inside this thickened area forms a layer of cells
between the epiblasts and the hypoblasts. At the end of the
immigration, the remaining cells on the surface ore the ectoderm.
The endoderm are the innermost layer of cells near the blastocoel
which has contributed some cells to the middle layer of sunken cells
to form the mesoderm.
Primitive groove the central furrow
Primitive ridges the thickened margins on both
sides of the primitive groove
Primitive plate thickening at the caudal end of
the streak
Primitive pit found at the cranial most end of
the streak. This appears in the slide as a light
area flanked by darkly-stained small streaks.
Hensens node elevation immediately anterior
to the primitive pit. This is represented by the
streaks that bound the primitive pit. This is
where the presumptive notochordal cells
migrate. Syn: Primitive knot

Megalecithal egg

Gastrulation starts with a thickening of the cells in the

posterior portion of the area pellucida. As this happens, the
cells inside the blastodisc split and proliferate into the
blastocoels. The cells that splitted off are the hypoblasts
while the remaining cells on top are the epiblasts.

Epiboly and involution holds true also in megalectihal eggs,

but the manner in which they are accomplished is
diffrerent from the medialecithal eggs.
o Area opaca
- The darker, outer, peripheral area seen in the
specimens (due to the presence of yolk beneath it)
o Area pellucida
- The clearer, central area in the specimen.
- This area is separated from the yolk by the
segmentation cavity, thus the appearance.
o Primitive streak
- The thickened part of the blastoderm. It appears as a
long streak in the center part of the specimen. This is
brought by the convergence of its surface layer