Cleavage and types
The process of cleavage reamains one of the earliest mechanical
activity in the conversion of a single celled egg into a multicellular embryo.
It is initiated by the sperm during fertilization. However in parthenogenetic
eggscleavage can commence without the influence of fertilization.
The process of cleavage or cellulation happens
through repeated mitotic divisions. These divisions result in cells called blastomeres.
In later stages of development the blastomeres occupy different regions and
differentiate into several types of body cells.
The first cleavage of frog’s egg was observed by Swammerdam
in 1738. The entire process of cleavage in frog’s egg was studied by Prevost
and Dumas in 1824. With the development of microscopes cleavages and
further stages were observed in the eggs of sea urchin, star fishes, amphioxus
and hen’s eggs.
From all these studies it has become clear that all
divisions in cleavage are mitotic. The mitotic process is very rapid. In
the eggs of sea urchin division of the blastomeres can be observed every 30
minutes.
As the cleavage
progresses the resultant daughter cells, namely the blastomeres get reduced in
size. During cleavage there is no growth in the blastomeres. The total size and
volume of the embryo remains the same.
The cleavages result in a compact mass of blastomeres
called morula. It gets transformed into blastula. While the wall
of the blastula is called the blastoderm, the central cavity is called
the blastocoel.
The planes of cleavage
An egg can be divided from different planes during
cleavage. Depending on the position of the cleavage furrow the planes of
cleavage are named.
1. Meridional plane: The plane of cleavage lies on
the animal vegetal axis. It bisects both the poles of the egg. Thus the egg is
divided into two equal halves.
2. Vertical plane: The cleavage furrows may lie on
either side of the
meridional plane. The furrows pass from animal to vegital
pole. The cleaved
cells may be unequal in size.
3. Equatorial plane: This cleavage plane bisects
the egg at right angles
to the main axis. It lies on the equatorial plane. It
divides the egg into two
halves.
4. Latitudinal plane: It is similar to the
equatorial plane, but it lies on either side of the equator. It is also called
as transverse or horizontal cleavage.
Influence of yolk on cleavage
Yolk is needed for embryonic development. However the
fertilized egg has to undergo all stages of development and result in a
suitable ‘young form’ initiating next generation. Somehow with all the
influences of yolk the developmental procedures are so adapted and modified
that a well formed embryo will result. The initial influence of yolk is felt
during the process of cleavage.The amount of the yolk and its distribution
affect the process of cleavage.
Accordingly several cleavage patterns have been
recognised.
1. Total or holoblastic cleavage - In this
type the cleavage furrow bisects
the entire egg. Such a cleavage may be either equal or
unequal.
(a) Equal holoblastic cleavage - In microlecithal
and isolecithal eggs, cleavage leads to the formation of blastomeres of equal
size. Eg: Amphioxus andplacental mammals.
(b) Unequal holoblastic cleavage - In mesolecithal
and telolocithal eggs,cleavage leads to the formation of blastomeres of unequal
size. Among the blastomeres there are many small sized micromeres and a few
large sized macromeres.
2. Meroblastic cleavage - In this type the
cleavage furrows are restricted to the active cytoplasm found either in the
animal pole (macrolecithal egg) or superficially surrounding the egg
(centrolecithal egg). Meroblastic cleavage may be of two types.
(a) Discoidal cleavage - Since the macrolecithal
eggs contain plenty of yolk, the cytoplasm is restricted to the narrow region
in the animal pole. Hencecleavage furrows can be formed only in the disc-like
animal pole region. Sucha cleavage is called discoidal meroblastic cleavage.
Eg: birds and reptiles.
(b) Superficial cleavage - In centrolecithal eggs,
the cleavage is restricted to
Laws of cleavage
Apparently there are several cleavage patterns. However,
all cleavages follow a common procedure. The cleavages are governed by certain
basic principles or laws.
1. Sach’s laws - These laws were proposed by Sach
in 1877.
i) Cells tend to divide into equal daughter cells
ii) Each new division plane tends to intersect the
preceding plane at right angles.
2. Balfour’s law (Balfour 1885) - “The speed or
rate of cleavage in any region of egg is inversely
proportional to the amount of yolk it contains”.
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