EMBRYOLOGY

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Cleavage
refers to the stereotyped pattern of early mitotic divisions that
divides up the large volume egg cytoplasm. The early zygote is
unique in being so large. Most cells undergo a period of growth
between cycles of mitosis, but this is not true for early cleavage
stage blastomeres. With each division the cells get smaller.

PATTERNS OF EMBRYONIC CLEAVAGE

Pattern of embryonic cleavage is determined both by the

position of the mitotic spindles and by the amount and
distribution of yolk. Yolk tends to inhibit cleavage. It slows it
down or actually prevents complete cleavage. Yolk is an
adaptation of those animals that go through more or less of
embryogenesis isolated from any food supply. Some animals,
like sea urchin, have relatively little yolk because they rapidly
develop into a free swimming larval form that acquires nutrients
from their environment. Other animals such as marsupials are
born prematurely, but are provided nourishment in a parental
pouch. Placental mammals develop a specialized organ through
which the embryo is nourished throughout development and so
also have little yolk.

The types of eggs based on yolk characteristics are described as:

Isolecithal: sparse evenly distributed yolk, eg., sea urchin,
mouse
Mesolecthal: moderate amount of yolk, often unevenly
distributed, eg., frog

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Telolecithal: dense yolk concentrated at one end, eg., bird,
reptile
Centrolecithal:yolk concentrated at the middle of the egg,eg.fly

Many eggs are polarized with a yolk rich pole, termed the
vegetal pole and a yolk poor pole termed the animal pole, eg.,
frog. The zygotic nucleus is generally displaced towards the
animal pole.

A. Based on the amount and pattern of distribution of yolk in the

zygote, cleavage is of two types: holoblastic and meroblastic.

1. Holoblastic cleavage:

It divides the zygote and blastomeres completely into daughter

cells. It is of two types: equal and unequal.

(i) Equal Holoblastic Cleavage:

It forms equal blastomeres. It occurs in star fish.

(ii) Unequal Holoblastic Cleavage:

It forms unequal blastomeres. Blastomeres are micromeres

(smaller) and macromeres (larger). It is found in frog.

2. Meroblastic cleavage:

In this type of cleavage, the divisions are confined to the animal

pole or peripheral region of egg. The yolk remains undivided. It
is of two types: discoidal and superficial.

(i) Discoidal Cleavage:

The divisions are confined to the cytoplasmic disc located at the

animal pole. It occurs in reptiles, birds and egg laying mammals.

(ii) Superficial Cleavage:

The cleavage remains restricted to the peripheral portion of the

egg. It occurs in arthropods especially insects.
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B. Based on the potentiality of the blastomeres, cleavage is of
two types: determinate and indeterminate.

1. Determinate (Mosaic) Cleavage:

In this type of cleavage a complete embryo is formed only if all

the blastomeres remain together e.g., annelid eggs.

2. Indeterminate (Non-mosaic) Cleavage:

In this type of cleavage each early blastomere on separation

from other blasomeres may give rise to complete embryo e.g.,
chordate eggs.

There are several types of cleavage symmetry seen in nature:

radial (echinoderms, amphibians), spiral (mollusks, annelids),
Bilateral (ascidians,tunicates), Rotational (mammals). The two
figures below show examples of holoblastic and meroblastic
cleavage symmetries.

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Sea urchin cleavage
Sea urchins also have radial holoblastic cleavage, but with some
interesting differences. First cleavage is meridional. Second
cleavage is meridional. Third cleavage is equatorial Fourth
cleavage is meridional, but while the four animal pole cells split
equally to give rise to eight equal sized animal blastomeres
termed MESOMERES, the vegetal cells divide asymmetrically
along the equatorial plane to give 4 large MACROMERES and
4 much smaller MICROMERES at the vegetal pole. Fifth
division the MESOMERES divide equatorially to give two tiers
of eight MESOMERES an1 and an2 , the MACROMERES
divide meridionally forming a tier of eight cells below an2, the
MICROMERES divide to give a cluster of cells below the veg1
layer. The sixth divisions are all equatorial, giving a veg2 layer.
The seventh divisions are all meridional giving a 128 cell
blastula.

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The 128 cell blastula is a rather loose ball of cells surrounding a
hollow blastocoel. The ball is one cell layer thick with all cells
in contact with the external hyaline layer and the internal fluid
of the blastocoel. At this stage in development the cells begin to
form the tight junctions characteristic of an epithelium. The
central blastocoel is now isolated from the external
environment. The blastomeres continue to divide with their axis
parallel to the hyaline layer, remaining a epithelium one cell
thick. The blastocoel continues to enlarge.

Two theories attempt to account for the pattern of enlargement

of the blastocyst

1. The osmotic theory suggests that ions and proteins are

secreted into the blastocoel by the blastomeres and this results in
a pressure buildup due to the osmotic flow of water. This
pressure would then be responsible for aligning the axis mitosis
of the blastomeres and the enlargement of the blastocoel.

2. The alternate theory by Wolpert and his colleagues suggests

that it is really the adhesive interactions among the blastomeres
and between the blastomeres and the hyaline layer that aligns
the mitotic axis's. That is the adhesion to the hyaline is greatest,
the adhesion to other blastomeres is next, and finally the
interaction with the blastocoel wall is least. The dominant
adhesion with the hyaline layer forces the expansion of the
blastocyst and blastocoel.The cells of the blastula grow cilia on
their outer surface, secrete a “hatching enzyme” (hyalinase) and
become free swimming.

AMPHIBIAN CLEAVAGE

Cleavage in many amphibians is holoblastic with radial

symmetry, however the large volume of yolk (its mesolecithal)
interferes with cleavage. At the animal pole first cleavage
proceeds at about 1mm/min, while through the vegetal pole is
proceeds 50-100 times slower (.02mm/min). While the first
cleavage is still incomplete in the yolky vegetal region of the

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egg the second meridional cleavage begins to take place.

The third cleavage is equatorial, but because the nuclei and

asters are displaced “animal-ward” the cleavage plane although

perpendicular to the animal vegetal axis is also displaced
towards the animal pole and does not equally divide the
blastomeres. The result is four smaller animal blastomeres
(termed MICROMERES) and four large vegetal pole
blastomeres (termed MACROMERES). This unequal
holoblastic cleavage gives rise to a more rapidly dividing animal
pole made up of smaller micromeres and a slower dividing
vegetal pole made up of macromeres. The animal pole soon is
composed of many small micromeres and the vegetal pole a few
yolk filled large macromeres. Although the formation of the
blastocoel begins with the first cleavage, it does not become
obvious until the 128 cell stage.

WHAT FUNCTION DOES THE BLASTOCOEL SERVE?

The blastocel spatially separates cells so they do not touch one
another. Cells at the roof of the blastocoel normally become
ectoderm. If you transplant cells from the roof of the blastocoel
next to the yolky cells at the base of the blastocoel they will
differentiate as mesoderm. Mesodermal derivatives are normally
produced from cells adjacent to the endodermal precursors. One

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possibility that we will thoroughly explore is that the vegetal
cells “induce” via cell-cell interactions the adjacent cells to
become mesodermal. Thus the formation of the blastocoel may
be necessary to prevent inappropriate "inductive" interactions
among early cells of the blastocyst. The second obvious need for
the blastocoel may be during the subsequent stage of
development, GASTRULATION, where cells migrate into the
interior of the blastocoel.

MAMMALIAN CLEAVAGE

The mammalian egg is released from the ovary into the oviduct
where it is fertilized. First cleavage begins about a day after
fertilization within the oviduct. In sharp contrast to most
animals, cleavage in mammals can be very slow---1/day.

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Additionally, the cleavage planes are somewhat different from
other animals. First cleavage is meridional just like sea urchin
and frog. However, the second cleavage division sees one of the
blastomeres dividing meridionally and the other equatorially!
This type of cleavage is called ROTATIONAL HOLOBLASTIC
CLEAVAGE.

Another unique feature of mammalian cleavage is that the

blastomere cleavages are asynchronous. (compared with the
synchrony of sea urchin and frog up till the midblastula
transition). Cleavage of the mammalian embryo is regulated by
the zyotic nucleus from the very start.
Through the third cleavage the blastomeres form a ball of
loosely associated cells just like the other animals we’ve
studied. Before the fourth cleavage the cells of the blastula
dramatically change their behavior towards one another. They
rapidly try to maximize their contacts with the other blastomeres
and in doing cause the blastula to compact.

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MEROBLASTIC CLEAVAGE

In telolecithal and centrolecithal eggs the large dense yolk

prevents cleavage. Telolecithal eggs are characteristic of birds,
fishes, and reptiles while centrolecithal eggs are characteristic of
insects. Telolecithal eggs result in meroblastic discoidal
cleavage. Cleavage is restricted to the blastodisc at the animal
pole of the egg. At early cleavages, because cleavage cannot
proceed through the yolk, the blastomeres are continuous at their
vegetal margins.

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It's not until the equatorial cleavages that the cells of the
blastoderm separate from the yolk. Further equatorial cleavages
create a multilayered blastoderm three or four cells thick.

In birds a space forms between the blastoderm and the yolk

called the SUBGERMINAL cavity. By the 16 division (60,000
cells) cells of the blastoderm migrate into the subgerminal
cavity to form a second layer. The two layers are called the
outer EPIBLAST and inner HYPOBLAST.

Centrolecital eggs of arthropods undergo a SUPERFICIAL

CLEAVAGE. The large central mass of yolk confines the
cleavages to the cytoplasmic rim of the egg.

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