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In embryology, cleavage is the division of cells in the early embryo. The zygotes of many species undergo rapid cell cycles with no significant growth, producing a cluster of cells the same size as the original zygote. Depending mostly on the amount of yolk in the egg, the cleavage can be holoblastic (total or entire cleavage) or meroblastic (partial cleavage). The pole of the egg with the highest concentration of yolk is referred to as the vegetal pole while the opposite is referred to as the animal pole.
The end of cleavage coincides with the beginning of zygotic transcription. This point is referred to as the mid-blastula transition and appears to be controlled by the nuclear/cytoplasmic ratio (about 1/6). The different cells derived from cleavage (up to the blastula stage) are called blastomeres. A blastula is formed in many animals with a fluid-filled central cavity called a blastocoel.
Additional recommended knowledge
The rapid cell cycles are facilitated by maintaining high levels of proteins that control cell cycle progression such as the cyclins and their associated cyclin dependent kinases (cdk). The complex CyclinB/cdc2 aka MPF (maturation promoting factor) promotes entry into mitosis.
The processes of karyokinesis (mitosis) and cytokinesis work together to result in cleavage. The mitotic apparatus is made up of a central spindle and polar asters made up of polymers of tubulin protein called microtubules. The asters are nucleated by centrosomes and the centrosomes are organized by centrioles brought into the egg by the sperm as basal bodies. Cytokinesis is mediated by the contractile ring made up of polymers of actin protein called microfilaments. Karyokinesis and cytokinesis are independent but spatially and temporally coordinated processes. While mitosis can occur in the absence of cytokinesis, cytokinesis requires the mitotic apparatus.
Types of cleavage
In the absence of a large concentration of yolk, four major cleavage types can be observed in isolecithal cells (cells with a small even distribution of yolk) or in mesolecithal cells (moderate amount of yolk in a gradient) - bilateral holoblastic, radial holoblastic, rotational holoblastic, and spiral holoblastic, cleavage. These holoblastic cleavage planes pass all the way through isolecithal zygotes during the process of cytokinesis. Coeloblastula is the next stage of development for eggs that undergo these radial cleavaging. In holoblastic eggs the first cleavage always occurs along the vegetal-animal axis of the egg, the second cleavage is perpendicular to the first. From here the spatial arrangement of blastomeres can follow various patterns, due to different planes of cleavage, in various organisms.
In the presence of a large amount of yolk in the fertilized egg cell, the cell can undergo partial, or meroblastic, cleavage. Two major types of meroblastic cleavage are discoidal and superficial.
There are several differences between the cleavage in mammals and the cleavage in other animals. Mammals have a slow rate of division that is between 12 and 24 hours. These cellular division are asynchronous. Zygotic transcription starts at the two, four, or eight-cell stage. Cleavage is holoblastic and rotational.
At the eight-cell stage, the embryo goes through a process called compaction. Most of the blastomeres in this stage become polarized and develop tight junctions with the other blastomeres. This process leads to the development of two different populations of cells: polar cells on the outside and apolar cells on the inside. The outer cells, called the trophoblast cells, secrete fluid on their basal (inner) surface to form a blastocoel cavity through the process of cavitation. These trophoblast cells will eventually give rise to the embryonic contribution to the placenta called the chorion. The inner cells adhere to one side of the cavity to form the inner cell mass (ICM) and will give rise to the embryo and some extraembryonic membranes. At this stage, the embryo is called a blastocyst.
|This article is licensed under the GNU Free Documentation License. It uses material from the Wikipedia article "Cleavage_(embryo)". A list of authors is available in Wikipedia.|