In embryology, cleavage is the division of cells in the early development of the embryo, following fertilization. [1] The zygotes of many species undergo rapid cell cycles with no significant overall growth, producing a cluster of cells the same size as the original zygote. The different cells derived from cleavage are called blastomeres and form a compact mass called the morula. Cleavage ends with the formation of the blastula, or of the blastocyst in mammals.
Depending mostly on the concentration 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.
Cleavage differs from other forms of cell division in that it increases the number of cells and nuclear mass without increasing the cytoplasmic mass. This means that with each successive subdivision, there is roughly half the cytoplasm in each daughter cell than before that division, and thus the ratio of nuclear to cytoplasmic material increases. [2]
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 (CDKs). The complex cyclin B/CDK1 also known as 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.
The end of cleavage coincides with the beginning of zygotic transcription. This point in non-mammals is referred to as the midblastula transition and appears to be controlled by the nuclear-cytoplasmic ratio (about 1:6).
Determinate cleavage (also called mosaic cleavage) is in most protostomes. It results in the developmental fate of the cells being set early in the embryo development. Each blastomere produced by early embryonic cleavage does not have the capacity to develop into a complete embryo.
A cell can only be indeterminate (also called regulative) if it has a complete set of undisturbed animal/vegetal cytoarchitectural features. It is characteristic of deuterostomes—when the original cell in a deuterostome embryo divides, the two resulting cells can be separated, and each one can individually develop into a whole organism.
In holoblastic cleavage, the zygote and blastomeres are completely divided during the cleavage, so the number of blastomeres doubles with each 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 or microlecithal cells (moderate concentration of yolk in a gradient)—bilateral holoblastic, radial holoblastic, rotational holoblastic, and spiral holoblastic, cleavage. [3] 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 cleavages. 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 concentration 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.[ citation needed ]
I. Holoblastic (complete) cleavage | II. Meroblastic (incomplete) cleavage |
---|---|
A. Isolecithal (sparse, evenly distributed yolk)
B. Mesolecithal (moderate vegetal yolk disposition)
| A. Telolecithal (dense yolk throughout most of cell)
B. Centrolecithal (yolk in center of egg)
|
Mammals have a slow rate of division that is between 12 and 24 hours. These cellular divisions are asynchronous. Zygotic transcription starts at the two-, four-, or eight-cell stage. Cleavage is holoblastic and rotational.
In human embryonic development at the eight-cell stage, having undergone three cleavages the embryo goes through some changes as it develops into a blastocyst. At the eight-cell stage the blastomeres are round, and only loosely adhered. With further division in the process of compaction the cells become flattened, and develop an inside-out polarity that optimises the cell to cell contact between them. They begin to tightly adhere as gap junctions are formed, and tight junctions are developed with the other blastomeres. [13] [14] At the 16–32 cell stage the compacted embryo is called a morula. [14] [15] Once the embryo has divided into 16 cells, it begins to resemble a mulberry, hence the name morula (Latin, morus: mulberry). [16] With further compaction the individual outer blastomeres, the trophoblasts, become indistinguishable as they become organised into a thin sheet of tightly adhered epithelial cells. They are still enclosed within the zona pellucida. This compaction serves to make the structure watertight, to contain the fluid that the cells will later secrete.
In the human the morula enters the uterus after three or four days, and begins to take in fluid, as sodium-potassium pumps on the trophoblasts pump sodium into the morula, drawing in water from the maternal environment to become blastocoelic fluid. Hydrostatic pressure of the fluid creates a large cavity in the morula called a blastocoel. Embryoblast cells also known as the inner cell mass form a compact mass of cells at the embryonic pole on one side of the cavity that will go on to produce the embryo proper. The embryo is now termed a blastocyst. [14] [17] The trophoblasts will eventually give rise to the embryonic contribution to the placenta called the chorion.
A single cell can be removed from a pre-compaction eight-cell embryo and used for genetic screening, and the embryo will recover. [18] [19]
Differences exist between cleavage in placental mammals and other mammals.
A zygote is a eukaryotic cell formed by a fertilization event between two gametes. The zygote's genome is a combination of the DNA in each gamete, and contains all of the genetic information of a new individual organism.
An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male sperm cell. The resulting fusion of these two cells produces a single-celled zygote that undergoes many cell divisions that produce cells known as blastomeres. The blastomeres are arranged as a solid ball that when reaching a certain size, called a morula, takes in fluid to create a cavity called a blastocoel. The structure is then termed a blastula, or a blastocyst in mammals.
Hemichordata is a phylum which consists of triploblastic, enterocoelomate, and bilaterally symmetrical marine deuterostome animals, generally considered the sister group of the echinoderms. They appear in the Lower or Middle Cambrian and include two main classes: Enteropneusta, and Pterobranchia. A third class, Planctosphaeroidea, is known only from the larva of a single species, Planctosphaera pelagica. The class Graptolithina, formerly considered extinct, is now placed within the pterobranchs, represented by a single living genus Rhabdopleura.
Blastulation is the stage in early animal embryonic development that produces the blastula. In mammalian development the blastula develops into the blastocyst with a differentiated inner cell mass and an outer trophectoderm. The blastula is a hollow sphere of cells known as blastomeres surrounding an inner fluid-filled cavity called the blastocoel. Embryonic development begins with a sperm fertilizing an egg cell to become a zygote, which undergoes many cleavages to develop into a ball of cells called a morula. Only when the blastocoel is formed does the early embryo become a blastula. The blastula precedes the formation of the gastrula in which the germ layers of the embryo form.
Gastrulation is the stage in the early embryonic development of most animals, during which the blastula, or in mammals the blastocyst is reorganized into a two-layered or three-layered embryo known as the gastrula. Before gastrulation, the embryo is a continuous epithelial sheet of cells; by the end of gastrulation, the embryo has begun differentiation to establish distinct cell lineages, set up the basic axes of the body, and internalized one or more cell types including the prospective gut.
The blastocyst is a structure formed in the early embryonic development of mammals. It possesses an inner cell mass (ICM) also known as the embryoblast which subsequently forms the embryo, and an outer layer of trophoblast cells called the trophectoderm. This layer surrounds the inner cell mass and a fluid-filled cavity known as the blastocoel. In the late blastocyst the trophectoderm is known as the trophoblast. The trophoblast gives rise to the chorion and amnion, the two fetal membranes that surround the embryo. The placenta derives from the embryonic chorion and the underlying uterine tissue of the mother.
In biology, a blastomere is a type of cell produced by cell division (cleavage) of the zygote after fertilization; blastomeres are an essential part of blastula formation, and blastocyst formation in mammals.
The blastocoel, also spelled blastocoele and blastocele, and also called cleavage cavity, or segmentation cavity is a fluid-filled or yolk-filled cavity that forms in the blastula during very early embryonic development. At this stage in mammals the blastula develops into the blastocyst containing an inner cell mass, and outer trophectoderm.
In developmental biology, animal embryonic development, also known as animal embryogenesis, is the developmental stage of an animal embryo. Embryonic development starts with the fertilization of an egg cell (ovum) by a sperm cell, (spermatozoon). Once fertilized, the ovum becomes a single diploid cell known as a zygote. The zygote undergoes mitotic divisions with no significant growth and cellular differentiation, leading to development of a multicellular embryo after passing through an organizational checkpoint during mid-embryogenesis. In mammals, the term refers chiefly to the early stages of prenatal development, whereas the terms fetus and fetal development describe later stages.
Isolecithal refers to the even distribution of yolk in the cytoplasm of ova of mammals and other vertebrates, notably fishes of the families Petromyzontidae, Amiidae, and Lepisosteidae. Isolecithal cells have two equal hemispheres of yolk. However, during cellular development, normally under the influence of gravity, some of the yolk settles to the bottom of the egg, producing an uneven distribution of yolky hemispheres. Such uneven cells are known as telolecithal and are common where there is sufficient yolk mass.
In embryology, Carnegie stages are a standardized system of 23 stages used to provide a unified developmental chronology of the vertebrate embryo.
The inner cell mass (ICM) or embryoblast is a structure in the early development of an embryo. It is the mass of cells inside the blastocyst that will eventually give rise to the definitive structures of the fetus. The inner cell mass forms in the earliest stages of embryonic development, before implantation into the endometrium of the uterus. The ICM is entirely surrounded by the single layer of trophoblast cells of the trophectoderm.
An asymmetric cell division produces two daughter cells with different cellular fates. This is in contrast to symmetric cell divisions which give rise to daughter cells of equivalent fates. Notably, stem cells divide asymmetrically to give rise to two distinct daughter cells: one copy of the original stem cell as well as a second daughter programmed to differentiate into a non-stem cell fate.
In the field of developmental biology, regional differentiation is the process by which different areas are identified in the development of the early embryo. The process by which the cells become specified differs between organisms.
The development of fishes is unique in some specific aspects compared to the development of other animals.
The blastodisc, also called the germinal disc, is the embryo-forming part on the yolk of the egg of an animal that undergoes discoidal meroblastic cleavage. Discoidal cleavage occurs in those animals with a large proportion of yolk in their eggs, and include insects, fish, reptiles and birds. The blastodisc is a small disc of cytoplasm that sits on top of the yolk. In birds it is a small, circular, white spot on the surface of the yellow yolk of an egg, at the animal pole.
Cavitation is a process in early embryonic development that follows cleavage. Cavitation is the formation of the blastocoel, a fluid-filled cavity that defines the blastula, or in mammals the blastocyst. After fertilization, cell division of the zygote occurs which results in the formation of a solid ball of cells (blastomeres) called the morula. Further division of cells increases their number in the morula, and the morula differentiates them into two groups. The internal cells become the inner cell mass, and the outer cells become the trophoblast. Before cell differentiation takes place there are two transcription factors, Oct-4 and nanog that are uniformly expressed on all of the cells, but both of these transcription factors are turned off in the trophoblast once it has formed.
Leech embryogenesis is the process by which the embryo of the leech forms and develops. The embryonic development of the larva occurs as a series of stages. During stage 1, the first cleavage occurs, which gives rise to an AB and a CD blastomere, and is in the interphase of this cell division when a yolk-free cytoplasm called teloplasm is formed. The teloplasm is known to be a determinant for the specification of the D cell fate. In stage 3, during the second cleavage, an unequal division occurs in the CD blastomere. As a consequence, it creates a large D cell on the left and a smaller C cell to the right. This unequal division process is dependent on actomyosin, and by the end of stage 3 the AB cell divides. On stage 4 of development, the micromeres and teloblast stem cells are formed and subsequently, the D quadrant divides to form the DM and the DNOPQ teloblast precursor cells. By the end stage 6, the zygote contains a set of 25 micromeres, 3 macromeres and 10 teloblasts derived from the D quadrant.
This glossary of developmental biology is a list of definitions of terms and concepts commonly used in the study of developmental biology and related disciplines in biology, including embryology and reproductive biology, primarily as they pertain to vertebrate animals and particularly to humans and other mammals. The developmental biology of invertebrates, plants, fungi, and other organisms is treated in other articles; e.g terms relating to the reproduction and development of insects are listed in Glossary of entomology, and those relating to plants are listed in Glossary of botany.