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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 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 (fro ...
during very early embryonic development. At this stage in mammals the blastula develops into the
blastocyst 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 t ...
containing an
inner cell mass The inner cell mass (ICM) or embryoblast (known as the pluriblast in marsupials) 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 t ...
, and outer trophectoderm. It develops following cleavage of the zygote after
fertilization Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a new individual organism or offspring and initiate its development. Proce ...
. It is the first cell cavity formed as the embryo enlarges, and is the essential precursor for the differentiated gastrula. In the ''
Xenopus ''Xenopus'' () (Gk., ξενος, ''xenos''=strange, πους, ''pous''=foot, commonly known as the clawed frog) is a genus of highly aquatic frogs native to sub-Saharan Africa. Twenty species are currently described within it. The two best-known ...
'' a very small cavity has been described in the two-cell stage of development.


In mammals

After fertilization, the zygote undergoes mitotic divisions during rotational cleavage resulting in daughter cells known as
blastomere 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. Human blastomere characteristics In ...
s until they are at the 16-cell stage called the morula. The morula is a solid ball of cells that has a small group of internal cells surrounded by a larger group of external cells. Following further cleavages and compaction the blastomeres undergo
cellular differentiation Cellular differentiation is the process in which a stem cell alters from one type to a differentiated one. Usually, the cell changes to a more specialized type. Differentiation happens multiple times during the development of a multicellular ...
that gives rise to an internal group of cells called the
inner cell mass The inner cell mass (ICM) or embryoblast (known as the pluriblast in marsupials) 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 t ...
, (also ''embryoblast'') and an external layer of surrounding cells known as trophoblasts. The inner cell mass will go on to become the actual embryo. The external, surrounding cells develop into the trophoblast cells. At this stage there is no cavity within the morula; the embryo is still a ball of dividing cells. In a process called
cavitation Cavitation is a phenomenon in which the static pressure of a liquid reduces to below the liquid's vapour pressure, leading to the formation of small vapor-filled cavities in the liquid. When subjected to higher pressure, these cavities, cal ...
, the trophoblast cells secrete fluid into the morula to create a blastocoel, the fluid-filled cavity. The membranes of the trophoblast cells contain sodium (Na+) pumps, Na+/K+- ATPase and Na+/H+ exchangers, that pump sodium into the centrally forming cavity. The accumulation of sodium pulls in water osmotically, creating and enlarging the blastocoel. The oviduct cells stimulate these trophoblast sodium pumps as the fertilized egg travels down the fallopian tube towards the uterus. As the embryo further divides, the blastocoel expands and the inner cell mass is positioned on one side of the trophoblast cells forming a mammalian blastula, called a
blastocyst 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 t ...
.


In amphibians

An amphibian embryo in the 128- cell stage is considered a blastula as the blastocoel in the embryo becomes apparent during this stage. The fluid-filled cavity forms in the animal hemisphere of the frog. However, the early formation of the blastocoel has been traced back to the very first
cleavage furrow In cell biology, the cleavage furrow is the indentation of the cell's surface that begins the progression of cleavage, by which animal and some algal cells undergo cytokinesis, the final splitting of the membrane, in the process of cell division. ...
. It was demonstrated in the frog embryo that the first cleavage furrow widens in the animal hemisphere creating a small intercellular cavity that is sealed off via tight junctions. As cleavage continues, the cavity expands to become the developed blastocoel. The blastocoel is a crucial component of amphibian embryo development. It permits cell migration during gastrulation and prevents the cells beneath the blastocoel from interacting prematurely with the cells above the blastocoel. For instance, the blastocoel prevents the vegetal cells destined to become
endoderm Endoderm is the innermost of the three primary germ layers in the very early embryo. The other two layers are the ectoderm (outside layer) and mesoderm (middle layer). Cells migrating inward along the archenteron form the inner layer of the gastru ...
from coming in contact with those cells in the ectoderm fated to give rise to the skin and nerves.


Damage to blastocoel

The blastocoel can be damaged and abolished if the adhesion between blastomeres, provided by cell adhesion molecules like EP-cadherin, is destroyed as mRNA by oligonucleotides. If the mRNA is destroyed, then there’s no EP-cadherin, little to no blastomere adhesion and the blastocoel is non-existent. During the next stage of embryonic development, amphibian gastrulation, the blastocoel is displaced by the formation of the
archenteron The primary gut that forms during gastrulation in a developing embryo is known as the archenteron, the gastrocoel or the primitive digestive tube. It develops into the endoderm and mesoderm of an animal. Formation in sea urchins As primary mesenc ...
, during mid-gastrulation. At the end of gastrulation, the blastocoel has been obliterated.


In sea urchins

At the 120- cell stage, the sea urchin embryo is considered a blastula because of its developed blastocoel, which every embryonic cell surrounds and touches. Every cell is in contact with the proteinaceous fluid of the blastocoel on the inside and touches the hyaline layer on the outside. The loosely connected blastomeres are now tightly connected because of
tight junction Tight junctions, also known as occluding junctions or ''zonulae occludentes'' (singular, ''zonula occludens''), are multiprotein junctional complexes whose canonical function is to prevent leakage of solutes and water and seals between the epith ...
s that create a seamless epithelium that completely encircles the blastocoel. Even as the blastomeres continue to divide, the blastula remains one-cell thick and thins out as the embryo expands outward. This is accomplished in part due to the influx of water that expands the blastocoel and pushes the cells surrounding it outwards. At this point, the cells have become specified and are ciliated on the opposite side of the blastocoel. The vegetal plate and animal hemisphere develop and secrete a hatching enzyme that digests the fertilization envelope and allows the embryo to now become a free-swimming hatched blastula.


Development of primary mesenchyme

Important to the sea urchin blastula is the ingression of the
primary mesenchyme Mesenchymal stem cells (MSCs) also known as mesenchymal stromal cells or medicinal signaling cells are multipotent stromal cells that can differentiate into a variety of cell types, including osteoblasts (bone cells), chondrocytes (cartilage ce ...
. After the blastula hatches from the fertilization envelope, the vegetal side of the blastula begins to flatten and thicken as a small cluster of these cells develop long, thin processes called filopodia. These cells then dissociate and ingress into the blastocoel and are called the primary mesenchyme. The cells move randomly along the inside of the blastocoel, until they become localized in the ventrolateral region of the blastocoel.


In birds

Similar to mammals, fertilization of the avian ovum occurs in the oviduct. From there the blastodisc, a small cluster of cells in the animal pole of the egg, then undergoes
discoidal In embryology, cleavage is the division of cells in the early development of the embryo, following fertilization. The zygotes of many species undergo rapid cell cycles with no significant overall growth, producing a cluster of cells the same size ...
meroblastic In embryology, cleavage is the division of cells in the early development of the embryo, following fertilization. The zygotes of many species undergo rapid cell cycles with no significant overall growth, producing a cluster of cells the same size ...
cleavage. The blastoderm develops into the epiblast and hypoblast and it is between these layers that the blastocoel will form. The shape and formation of the avian blastodisc differs from amphibian, fish, and echinoderm blastulas, but the overall spatial relationship of the blastocoel remains the same.


Formation of primitive streak

The avian blastocoel is important during the development of the primitive streak. The ingression of the endodermal precursor cells form the epiblast into the blastocoel and the migration of lateral cells of the posterior epiblast towards the center form the early primitive streak. As these cells converge inward, a depression forms called the
primitive groove The primitive streak is a structure that forms in the early embryo in amniotes. In amphibians the equivalent structure is the blastopore. During early embryonic development, the embryonic disc becomes oval shaped, and then pear-shaped with the b ...
and functions as an opening through which cells travel into the blastocoel. As cells migrate into the blastocoel, they undergo an epithelial-to-mesenchymal transformation.


Zebrafish: a special case

Unlike amphibian, echinoderm, mammalian, and avian embryos,
zebrafish The zebrafish (''Danio rerio'') is a freshwater fish belonging to the minnow family (Cyprinidae) of the order Cypriniformes. Native to South Asia, it is a popular aquarium fish, frequently sold under the trade name zebra danio (and thus often ...
do not have a defined blastocoel. Rather, they have small, irregular extracellular spaces that are formed between the cells of the
blastodisc 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, ...
sitting atop the yolk.


References


Further reading

*Dorlands Staff (2004). "blastocoel istionary entry. ''Dorland's Illustrated Medical Dictionary'' (online). Amsterdam, NDE: Elsevier-Saunders. Retrieved 30 January 2016. "blastocoel... laso- + -coelethe fluid-filled cavity of the mass of cells (blastula) produced by cleavage of fertilized ovum. Sometimes spelled... lled...'Also' blastocoelic ...pertaining to the blastocoele."
Dorlands.com
* * * {{Extraembryonic and fetal membranes Animal developmental biology