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 (fr ...

during very early

embryonic development 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, sperm cell (spermat ...

. At this stage in mammals the blastula is called 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 ...

, which consists of an outer

epithelium Epithelium or epithelial tissue is a thin, continuous, protective layer of cells with little extracellular matrix. An example is the epidermis, the outermost layer of the skin. Epithelial ( mesothelial) tissues line the outer surfaces of man ...

, the

trophectoderm The trophoblast (from Greek : to feed; and : germinator) is the outer layer of cells of the blastocyst. Trophoblasts are present four days after fertilization in humans. They provide nutrients to the embryo and develop into a large part of the pl ...

, enveloping 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 ...

and the blastocoel . It develops following

cleavage Cleavage may refer to: Science * Cleavage (crystal), the way in which a crystal or mineral tends to split * Cleavage (embryo), the division of cells in an early embryo * Cleavage (geology), foliation of rock perpendicular to stress, a result of ...

of the

zygote A zygote (; , ) is a eukaryote, eukaryotic cell (biology), 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 individ ...

after

fertilization Fertilisation or fertilization (see American and British English spelling differences#-ise, -ize (-isation, -ization), spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give ...

. It is the first fluid-filled cavity or lumen formed as the embryo enlarges, and is the essential precursor for the differentiated

gastrula Gastrulation is the stage in the early embryonic development of most animals, during which the blastula (a single-layered hollow sphere of Cell (biology), cells), or in mammals, the blastocyst, is reorganized into a two-layered or three-layered e ...

. 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 with ...

'' a very small cavity has been described in the two-cell stage of development.

In mammals

After fertilization, the

undergoes several rounds of

divisions forming 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. At the 8- or 16-cell stage, the embryo undergoes compaction and forms the

morula 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 siz ...

. Eventually, the

is a solid ball of cells that has a small group of internal cells surrounded by a larger group of external cells. Then blastomeres undergo

cellular differentiation Cellular differentiation is the process in which a stem cell changes 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 multicellula ...

with internal cells adopting the

fate and the external layer becoming

. The inner cell mass will go on to become the actual embryo. The external, surrounding cells develop into

trophoblast The trophoblast (from Greek language, Greek : to feed; and : germinator) is the outer layer of cells of the blastocyst. Trophoblasts are present four days after Human fertilization, fertilization in humans. They provide nutrients to the embryo an ...

cells, which only contribute to extra-embryonic tissues. At this stage there is no lumen within the embryo. In a process called

cavitation Cavitation in fluid mechanics and engineering normally is the phenomenon in which the static pressure of a liquid reduces to below the liquid's vapor pressure, leading to the formation of small vapor-filled cavities in the liquid. When sub ...

cells transport fluid into the embryo to create a blastocoel, the fluid-filled lumen. The membranes of the

cells contain sodium (Na⁺) pumps, Na⁺/K⁺- ATPase and Na⁺/H⁺ exchangers, that pump sodium into the embryo. The oviduct cells stimulate these trophoblast sodium pumps as the fertilized egg travels down the fallopian tube towards the uterus. The accumulation of sodium pulls in water through

osmosis Osmosis (, ) is the spontaneous net movement or diffusion of solvent molecules through a selectively permeable membrane, selectively-permeable membrane from a region of high water potential (region of lower solute concentration) to a region of ...

. The accumulation of water breaks open cell-cell contacts via

hydraulic fracturing Fracking (also known as hydraulic fracturing, fracing, hydrofracturing, or hydrofracking) is a well stimulation technique involving the fracturing of Formation (geology), formations in bedrock by a pressurized liquid. The process involves the ...

. To form a single lumen, the fluid from multiple water pockets collects into a single entity in process akin to

Ostwald ripening Ostwald ripening is a phenomenon observed in solid solutions and liquid sols that involves the change of an inhomogeneous structure over time, in that small crystals or sol particles first dissolve and then redeposit onto larger crystals or s ...

. The blastocoel further expands and the inner cell mass becomes positioned on one side of the trophoblast cells forming a mammalian blastula, called a

. The axis formed by the

and the blastocoel is the first axis of symmetry of mammalian embryo and determines its attachment point to the uterus.

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 In developmental biology, an embryo is divided into two hemispheres: the animal pole and the vegetal pole within a blastula. The animal pole consists of small cells that divide rapidly, in contrast with the vegetal pole below it. In some cases, t ...

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 divisio ...

. 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 gastr ...

from coming in contact with those cells in the

ectoderm The ectoderm is one of the three primary germ layers formed in early embryonic development. It is the outermost layer, and is superficial to the mesoderm (the middle layer) and endoderm (the innermost layer). It emerges and originates from the o ...

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

oligonucleotide Oligonucleotides are short DNA or RNA molecules, oligomers, that have a wide range of applications in genetic testing, Recombinant DNA, research, and Forensic DNA, forensics. Commonly made in the laboratory by Oligonucleotide synthesis, solid-phase ...

s. 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 Gastrulation is the stage in the early embryonic development of most animals, during which the blastula (a single-layered hollow sphere of cells), or in mammals, the blastocyst, is reorganized into a two-layered or three-layered embryo known as ...

, the blastocoel is displaced by the formation of the

archenteron The archenteron, also called the gastrocoel, is the internal cavity formed in the gastrulation stage in early embryonic development that becomes the cavity of the primitive gut. Formation in sea urchins As primary mesenchyme cells detach fro ...

, 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 Cell junction, junctional complexes between epithelial cells, sealing and preventing leakage of solutes and water. Th ...

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. 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 Filopodia (: filopodium) are slender cytoplasmic projections that extend beyond the leading edge of lamellipodia in migrating cells. Within the lamellipodium, actin ribs are known as ''microspikes'', and when they extend beyond the lamellipod ...

. 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 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 ...

, 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 siz ...

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 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 ...

. As these cells converge inward, a depression forms called the primitive groove 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.

In zebrafish

Unlike amphibian, echinoderm, mammalian, and avian embryos,

zebrafish The zebrafish (''Danio rerio'') is a species of freshwater ray-finned fish belonging to the family Danionidae of the order Cypriniformes. Native to South Asia, it is a popular aquarium fish, frequently sold under the trade name zebra danio (an ...

do not have a defined blastocoel. Rather, they have small, irregular extracellular spaces that are formed between the cells of the

sitting atop the yolk.