The bilaminar embryonic disc, bilaminar blastoderm or embryonic disc is the distinct two-layered structure of cells formed in an

embryo An embryo ( ) is the initial stage of development for 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 sp ...

. In the development of the

human embryo Human embryonic development or human embryogenesis is the development and formation of the human embryo. It is characterised by the processes of cell division and cellular differentiation of the embryo that occurs during the early stages of deve ...

this takes place by day eight. It is formed when 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 known as the embryoblast, forms a bilaminar disc of two layers, an upper layer called the

epiblast In amniote embryonic development, the epiblast (also known as the primitive ectoderm) is one of two distinct cell layers arising from the inner cell mass in the mammalian blastocyst, or from the blastula in reptiles and birds. It drives the em ...

(primitive

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 ...

) and a lower layer called the

hypoblast In amniote embryology, the hypoblast is one of two distinct layers arising from the inner cell mass in the mammalian blastocyst, or from the blastodisc in reptiles and birds. The hypoblast gives rise to the yolk sac. The hypoblast is a layer of ...

(primitive

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 ...

), which will eventually form into

fetus A fetus or foetus (; : fetuses, foetuses, rarely feti or foeti) is the unborn offspring of a viviparous animal that develops from an embryo. Following the embryonic development, embryonic stage, the fetal stage of development takes place. Pren ...

. These two layers of cells are stretched between two fluid-filled cavities at either end: the primitive

yolk sac The yolk sac is a membranous wikt:sac, sac attached to an embryo, formed by cells of the hypoblast layer of the bilaminar embryonic disc. This is alternatively called the umbilical vesicle by the Terminologia Embryologica (TE), though ''yolk sac' ...

and the

amniotic sac The amniotic sac, also called the bag of waters or the membranes, is the sac in which the embryo and later fetus develops in amniotes. It is a thin but tough transparent pair of biological membrane, membranes that hold a developing embryo (and l ...

. The epiblast is adjacent to the

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 ...

and made of columnar

epithelial cells 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 hypoblast is closest to the

blastocoel 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 mammal ...

(blastocystic cavity) and made of cuboidal cells. As the two layers become evident, a

basement membrane The basement membrane, also known as base membrane, is a thin, pliable sheet-like type of extracellular matrix that provides cell and tissue support and acts as a platform for complex signalling. The basement membrane sits between epithelial tis ...

forms between the layers. This distinction of layers of the bilaminar disc defines the primitive dorso ventral axis and polarity in embryogenesis. The epiblast migrates away from the trophoblast downwards, forming the amniotic cavity in between, the lining of which is formed from amnioblasts developed from the epiblast. The hypoblast is pushed down and forms the yolk sac (exocoelomic cavity) lining. Some hypoblast cells migrate along the inner cytotrophoblast lining of the blastocoel, secreting an

extracellular matrix In biology, the extracellular matrix (ECM), also called intercellular matrix (ICM), is a network consisting of extracellular macromolecules and minerals, such as collagen, enzymes, glycoproteins and hydroxyapatite that provide structural and bio ...

along the way. These hypoblast cells and extracellular matrix are called Heuser's membrane (or the exocoelomic membrane), and they cover the blastocoel to form the yolk sac (or exocoelomic cavity). Cells of the hypoblast migrate along the outer edges of this reticulum and form the extraembryonic mesoderm; this disrupts the extraembryonic reticulum. Soon pockets form in the reticulum, which ultimately coalesce to form the chorionic cavity (extraembryonic coelom).

Initial formation

The one-celled

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 ...

, a eukaryotic cell formed by a

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 ...

event between two

gametes A gamete ( ) is a haploid cell that fuses with another haploid cell during fertilization in organisms that reproduce sexually. Gametes are an organism's reproductive cells, also referred to as sex cells. The name gamete was introduced by the Ge ...

at the start of

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 ...

, undergoes cleavage by

mitosis Mitosis () is a part of the cell cycle in eukaryote, eukaryotic cells in which replicated chromosomes are separated into two new Cell nucleus, nuclei. Cell division by mitosis is an equational division which gives rise to genetically identic ...

as it travels through the

fallopian tube The fallopian tubes, also known as uterine tubes, oviducts or salpinges (: salpinx), are paired tubular sex organs in the human female body that stretch from the Ovary, ovaries to the uterus. The fallopian tubes are part of the female reproduct ...

to the

uterus The uterus (from Latin ''uterus'', : uteri or uteruses) or womb () is the hollow organ, organ in the reproductive system of most female mammals, including humans, that accommodates the embryonic development, embryonic and prenatal development, f ...

. As the zygote undergoes

cell division Cell division is the process by which a parent cell (biology), cell divides into two daughter cells. Cell division usually occurs as part of a larger cell cycle in which the cell grows and replicates its chromosome(s) before dividing. In eukar ...

to form two, then four, then eight and then 16 cells (typically by day four after fertilization), it becomes a ball of cells called a

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 ...

. During these cellular divisions, the zygote remains the same size, but the number of cells increase. The morula enters the uterus after three or four days—during which a cavity, called the

, is formed to produce 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 ...

.Schoenwolf, Gary C., and William J. Larsen. Larsen's Human Embryology. 4th ed. Philadelphia: Churchill Livingstone/Elsevier, 2009. Print. Once the blastocyst is formed, it undergoes implantation into the

endometrium The endometrium is the inner epithelium, epithelial layer, along with its mucous membrane, of the mammalian uterus. It has a basal layer and a functional layer: the basal layer contains stem cells which regenerate the functional layer. The funct ...

. During implantation the blastocyst, which contains the

, undergoes

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 ...

into the two layers of the bilaminar embryonic disc. One of which is the epiblast, also known as the primitive ectoderm. The epiblast is the outer layer of the bilaminar embryonic disc and consists of columnar cells. The hypoblast, also known as the primitive endoderm, is the inner layer, closest to the endometrium, which consists of cuboidal cells. The epiblast will develop into the 'embryo proper', and the hypoblast into the outer layer of

fetal membranes The fetal membranes are the four extraembryonic membranes, associated with the developing embryo, and fetus in humans and other mammals. They are the amnion, chorion, allantois, and yolk sac. The amnion and the chorion are the chorioamniotic ...

(extraembryonic membranes). The blastocyst serves as a source of nutrients for the growing cells by

diffusion Diffusion is the net movement of anything (for example, atoms, ions, molecules, energy) generally from a region of higher concentration to a region of lower concentration. Diffusion is driven by a gradient in Gibbs free energy or chemical p ...

from the surrounding fluid."Bilaminar Embryonic Disc." Atlas of Human Embryology. Chronolab A.G. Switzerland, n.d. Web. 27 Nov. 2012. .

Amniotic sac formation

Beginning on day eight, the

is the first new cavity to form during the second week of development. Fluid collects between the epiblast and the hypoblast, which splits the epiblast into two portions. The layer at the embryonic pole grows around the amniotic sac, creating a barrier from the cytotrophoblast. This becomes known as the

amnion The amnion (: amnions or amnia) is a membrane that closely covers human and various other embryos when they first form. It fills with amniotic fluid, which causes the amnion to expand and become the amniotic sac that provides a protective envir ...

, which is one of the four fetal membranes and the cells it comprises are referred to as amnioblasts."10.1 Early Development and Implantation." The Embryoblast. N.p., n.d. Web. 29 Nov. 2012. Although the amniotic sac is initially smaller than the blastocyst it becomes larger by week eight until the entire embryo is encompassed by the amnion.

Yolk sac and gestational sac formation

The process of the formation of the

gestational sac The gestational sac is the large cavity of fluid surrounding the embryo. During early embryogenesis, it consists of the extraembryonic coelom, also called the chorionic cavity. The gestational sac is normally contained within the uterus. It is the ...

(chorionic cavity or extraembryonic coelom) and the

(umbilical vesicle) is still debated. The main theory states that formation of the membranes of the yolk sac begins with an increase in production of hypoblast cells, followed by different patterns of migration. On day eight, the first portion of hypoblast cells begin their migration and make what is known as the primary yolk sac, or Heuser's membrane (exocoelomic membrane). By day 12, the primary yolk sac has been disestablished by a new batch of migrating hypoblast cells that now contribute to the definitive yolk sac. While the primary yolk sac is forming, extraembryonic mesoderm migrate into the blastocyst cavity and fill it with loosely packed cells. When the extraembryonic mesoderm is separated into two portions, a new gap arises called the gestational sac. This new cavity is responsible for detaching the embryo and, its amnion and yolk sac, from the far wall of the blastocyst, which is now named the

chorion The chorion is the outermost fetal membrane around the embryo in mammals, birds and reptiles (amniotes). It is also present around the embryo of other animals, like insects and molluscs. Structure In humans and other therian mammals, the cho ...

. When the extraembryonic mesoderm splits into two layers, the amnion, yolk sac and chorion also become double-layered. The amnion and chorion are composed of extraembryonic ectoderm and mesoderm, whereas the yolk sac is composed of extraembryonic endoderm and mesoderm. By day 13, the connecting stalk, a dense portion of extraembryonic mesoderm, restrains the embryonic disc in the gestational sac.

Yolk sac during development

Like the amnion, the yolk sac is a fetal membrane that surrounds a cavity. Formation of the definitive yolk sac occurs after the extraembryonic mesoderm splits, and it becomes a double layered structure with hypoblast-derived endoderm on the inside and mesoderm surrounding the outside. The definitive yolk sac contributes greatly to the embryo during the fourth week of development, and executes critical functions for the embryo. One of which being the formation of blood, or

hematopoiesis Haematopoiesis (; ; also hematopoiesis in American English, sometimes h(a)emopoiesis) is the formation of blood cellular components. All cellular blood components are derived from haematopoietic stem cells. In a healthy adult human, roughly ten ...

. Also, primordial germ cells are first found in the wall of the yolk sac before primordial germ cell migration. After the fourth week of development, the growing embryonic disc becomes much larger than the yolk sac and eventually involutes before birth. Uncommonly, the yolk sac may persist as the

vitelline duct In the human embryo, the vitelline duct, also known as the vitellointestinal duct, the yolk stalk, the omphaloenteric duct, or the omphalomesenteric duct, is a long narrow tube that joins the yolk sac to the midgut lumen of the developing fetus. ...

and cause a congenital out pouching of the digestive tract called

Meckel's diverticulum A Meckel's diverticulum, a true congenital diverticulum, is a slight bulge in the small intestine present at birth and a vestigial remnant of the vitelline duct. It is the most common malformation of the Human gastrointestinal tract, gastrointesti ...

Epiblast cells during gastrulation

In the third week, gastrulation begins with the formation of the

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 ...

. Gastrulation occurs when pluripotent stem cells differentiate into the three germ cell layers:

mesoderm The mesoderm is the middle layer of the three germ layers that develops during gastrulation in the very early development of the embryo of most animals. The outer layer is the ectoderm, and the inner layer is the endoderm.Langman's Medical ...

and

. During gastrulation, cells of the epiblast migrate towards the primitive streak, enter it, and then move apart from it through a process called ingression.

Definitive endoderm development

On day 16, epiblast cells that are next to the primitive streak experience epithelial-to-mesenchymal transformation as they ingress through the primitive streak. The first wave of epiblast cells takes over the hypoblast, which slowly becomes replaced by new cells that eventually constitute the definitive endoderm. The definitive endoderm is what makes the lining of the gut and other associated gut structures.

Intraembryonic mesoderm development

Also beginning on day 16, some of the ingressing epiblast cells make their way into the area between the epiblast and the newly forming definitive endoderm. This layer of cells becomes known as intraembryonic mesoderm. After the cells have moved bilaterally from the primitive streak and matured, four divisions of intraembryonic mesoderm are made: cardiogenic mesoderm,

paraxial mesoderm Paraxial mesoderm, also known as presomitic or somitic mesoderm, is the area of mesoderm in the neurulating embryo that flanks and forms simultaneously with the neural tube. The cells of this region give rise to somites, blocks of tissue running ...

intermediate mesoderm Intermediate mesoderm or intermediate mesenchyme is a narrow section of the mesoderm (one of the three primary germ layers) located between the paraxial mesoderm and the lateral plate of the developing embryo. The intermediate mesoderm develops in ...

and

lateral plate mesoderm The lateral plate mesoderm is the mesoderm that is found at the periphery of the embryo. It is to the side of the paraxial mesoderm, and further to the axial mesoderm. The lateral plate mesoderm is separated from the paraxial mesoderm by a narrow r ...

Ectoderm development

After the definitive endoderm and intraembryonic mesoderm formations are complete, the remaining epiblast cells do not ingress through the primitive streak; rather they remain on the outside and form the ectoderm. It is not long until the ectoderm becomes the

neural plate In embryology, the neural plate is a key Development of the human body, developmental structure that serves as the basis for the nervous system. Cranial to the primitive node of the embryonic primitive streak, Ectoderm, ectodermal tissue thickens ...

and

surface ectoderm The surface ectoderm, AKA external ectoderm, is one of the two early embryonic divisions of the ectoderm. The other early division of the ectoderm is the neuroectoderm. The surface ectoderm develops into the following structures: *Skin (only th ...

. Due to the fact that an embryo develops cranial to caudal, the formation of ectoderm does not happen at the same rate during development. The more inferior portion of the primitive streak will still have epiblast cells ingressing to make intraembryonic mesoderm, while the more superior portion has already stopped ingressing. However, eventually gastrulation finishes and the three germ layers are complete.

References

External links

Diagram at manchester.ac.uk
{{Authority control Embryology