The chorion is the outermost

fetal membrane 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 memb ...

around the

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 ...

mammal A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...

bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...

s and

reptiles Reptiles, as commonly defined, are a group of tetrapods with an ectothermic metabolism and Amniotic egg, amniotic development. Living traditional reptiles comprise four Order (biology), orders: Testudines, Crocodilia, Squamata, and Rhynchocepha ...

(

amniotes Amniotes are tetrapod vertebrate animals belonging to the clade Amniota, a large group that comprises the vast majority of living terrestrial and semiaquatic vertebrates. Amniotes evolved from amphibious stem tetrapod ancestors during the ...

). It is also present around the embryo of other animals, like

insect Insects (from Latin ') are Hexapoda, hexapod invertebrates of the class (biology), class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (Insect morphology#Head, head, ...

s and

mollusc Mollusca is a phylum of protostome, protostomic invertebrate animals, whose members are known as molluscs or mollusks (). Around 76,000 extant taxon, extant species of molluscs are recognized, making it the second-largest animal phylum ...

Structure

In humans and other

theria Theria ( or ; ) is a scientific classification, subclass of mammals amongst the Theriiformes. Theria includes the eutherians (including the Placentalia, placental mammals) and the metatherians (including the marsupials) but excludes the egg-lay ...

n mammals, the chorion is one of the

s that exist during

pregnancy Pregnancy is the time during which one or more offspring gestation, gestates inside a woman's uterus. A multiple birth, multiple pregnancy involves more than one offspring, such as with twins. Conception (biology), Conception usually occurs ...

between the developing

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 ...

and mother. The chorion and 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 ...

together form 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 ...

. In humans it is formed by extraembryonic

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 the two layers of

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 ...

that surround the embryo and other membranes; the

chorionic villi Chorionic villi are Wiktionary:villus, villi that sprout from the chorion to provide maximal contact area with maternal blood. They are an essential element in pregnancy from a histology, histomorphologic perspective, and are, by definition, a pr ...

emerge from the chorion, invade 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 ...

, and allow the transfer of nutrients from maternal blood to fetal blood.

Layers

The chorion consists of two layers: an outer formed by the

, and an inner formed by the extra-embryonic

. The trophoblast is made up of an internal layer of cubical or prismatic cells, the

cytotrophoblast "Cytotrophoblast" is the name given to both the inner layer of the trophoblast (also called layer of Langhans) or the cells that live there. It is interior to the syncytiotrophoblast and external to the wall of the blastocyst in a developing embry ...

or layer of Langhans, and an external

multinucleate Multinucleate cells (also known as multinucleated cells or polynuclear cells) are eukaryotic cells that have more than one nucleus, i.e., multiple nuclei share one common cytoplasm. Mitosis in multinucleate cells can occur either in a coordinate ...

d layer, the

syncytiotrophoblast The syncytiotrophoblast (from the Greek 'syn'- "together"; 'cytio'- "of cells"; 'tropho'- "nutrition"; 'blast'- "bud") is the epithelial covering of the highly vascular embryonic placental villi, which invades the wall of the uterus to establish ...

Growth

The chorion undergoes rapid proliferation and forms numerous

process A process is a series or set of activities that interact to produce a result; it may occur once-only or be recurrent or periodic. Things called a process include: Business and management * Business process, activities that produce a specific s ...

es, the ''

'', which invade and destroy the uterine decidua, while simultaneously absorbing nutritive materials from it for the growth of the

. The chorionic villi are at first small and non-vascular, and consist of the trophoblast only, but they increase in size and ramify, whereas the mesoderm, carrying branches of the umbilical vessels, grows into them, and they are

vascularized Angiogenesis is the physiological process through which new blood vessels form from pre-existing vessels, formed in the earlier stage of vasculogenesis. Angiogenesis continues the growth of the vasculature mainly by processes of sprouting an ...

. Blood is carried to the villi by the paired

umbilical arteries The umbilical artery is a paired artery (with one for each half of the body) that is found in the abdominal and pelvic regions. In the fetus, it extends into the umbilical cord. Structure Development The umbilical arteries supply systemic arte ...

, which branch into

chorionic arteries Chorionic (plate) vessels, also fetal surface vessels are blood vessels, including both arteries and veins, that carry blood through the chorion in the fetoplacental circulation. Chorionic arteries branch off the umbilical artery, and supply the ...

and enter the

cotyledon arteries The placenta of humans, and certain other mammals contains structures known as cotyledons, which transmit fetal blood and allow exchange of oxygen and nutrients with the maternal blood. Ruminants The Artiodactyla have a cotyledonary placenta. I ...

. After circulating through the capillaries of the villi, the blood is returned to the embryo by the

umbilical vein The umbilical vein is a vein present during fetal development that carries oxygenated blood from the placenta into the growing fetus. The umbilical vein provides convenient access to the central circulation of a neonate for restoration of blood vo ...

. Until about the end of the second month of

, the villi cover the entire chorion, and are almost uniform in size; but, after this, they develop unequally.

Parts

The part of the chorion that is in contact with the decidua capsularis undergoes atrophy, so that by the fourth month scarcely a trace of the villi is left. This part of the chorion becomes smooth, and is named the chorion laeve (from the Latin word ''levis'', meaning smooth). As it takes no share in the formation of the placenta, this is also named the non-placental part of the chorion. As the chorion grows, the chorion laeve comes in contact with the decidua parietalis and these layers fuse. The villi at the embryonic pole, which is in contact with the

decidua basalis The decidua is the modified mucosal lining of the uterus (that is, modified endometrium) that forms every month, in preparation for pregnancy. It is shed off each month when there is no fertilized egg to support. The decidua is under the influe ...

, increase greatly in size and complexity, and hence this part is named the chorion frondosum. Thus the placenta develops from the chorion frondosum and the decidua basalis.

Monochorionic twins

''Monochorionic twins'' are

twins Twins are two offspring produced by the same pregnancy.MedicineNet > Definition of Twin Last Editorial Review: 19 June 2000 Twins can be either ''monozygotic'' ('identical'), meaning that they develop from one zygote, which splits and forms two e ...

that share the same

placenta The placenta (: placentas or placentae) is a temporary embryonic and later fetal organ that begins developing from the blastocyst shortly after implantation. It plays critical roles in facilitating nutrient, gas, and waste exchange between ...

. This occurs in 0.3% of all pregnancies, and in 75% of

monozygotic Twins are two offspring produced by the same pregnancy.MedicineNet > Definition of Twin Last Editorial Review: 19 June 2000 Twins can be either ''monozygotic'' ('identical'), meaning that they develop from one zygote, which splits and forms two e ...

(identical) twins, when the split takes place on or after the third day 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 ...

. The remaining 25% of monozygous twins become ''dichorionic diamniotic''. The condition may affect any type of

multiple birth A multiple birth is the culmination of a multiple pregnancy, wherein the mother gives birth to two or more babies. A term most applicable to vertebrate species, multiple births occur in most kinds of mammals, with varying frequencies. Such births ...

, resulting in ''monochorionic multiples''.

Infections

Recent studies indicate that the chorion may be susceptible to pathogenic infections. Recent findings indicate that '' Ureaplasma parvum''

bacteria Bacteria (; : bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of Prokaryote, prokaryotic microorganisms. Typically a few micr ...

can infect the chorion tissue, thereby impacting pregnancy outcome. In addition, footprints of

JC polyomavirus Human polyomavirus 2, commonly referred to as the JC virus or John Cunningham virus, is a type of human polyomavirus. It was identified by electron microscopy in 1965 by ZuRhein and Chou, and by Silverman and Rubinstein. It was later isolated in ...

and

Merkel cell polyomavirus Merkel cell polyomavirus (MCV or MCPyV) was first described in January 2008 in Pittsburgh, Pennsylvania. It was the first example of a human viral pathogen discovered using unbiased metagenomic next-generation sequencing with a technique called ...

have been detected in chorionic villi from females affected by spontaneous abortion as well as pregnant women. Another

virus A virus is a submicroscopic infectious agent that replicates only inside the living Cell (biology), cells of an organism. Viruses infect all life forms, from animals and plants to microorganisms, including bacteria and archaea. Viruses are ...

BK polyomavirus The BK virus, also known as Human polyomavirus 1, is a member of the polyomavirus family. Past infection with the BK virus is widespread, but significant consequences of infection are uncommon, with the exception of the immunocompromised and the ...

has been detected in the same tissues, but with lesser extent.

Other animals

reptile Reptiles, as commonly defined, are a group of tetrapods with an ectothermic metabolism and Amniotic egg, amniotic development. Living traditional reptiles comprise four Order (biology), orders: Testudines, Crocodilia, Squamata, and Rhynchocepha ...

s, and

monotreme Monotremes () are mammals of the order Monotremata. They are the only group of living mammals that lay eggs, rather than bearing live young. The extant monotreme species are the platypus and the four species of echidnas. Monotremes are typified ...

s, the chorion is one of the four

extraembryonic membrane The extraembryonic membranes are four membranes which assist in the development of an animal's embryo. Such membranes occur in a range of animals from humans to insects. They originate from the zygote, but are not considered part of the embryo. The ...

s that make up the amniotic egg that provide for the nutrients and protection needed for the embryo's survival. It is located inside the albumen, which is the white of the egg. It encloses the embryo and the rest of the embryonic system. The chorion is also present in

s. During growth and development of the embryo, there is an increased need for oxygen. To compensate for this, the chorion and the

allantois The allantois ( ; : allantoides or allantoises) is one of the extraembryonic membranes arising from the yolk sac. It is a hollow sac-like structure filled with clear fluid that forms part of the developing conceptus in an amniote that helps the ...

fuse together to form the

chorioallantoic membrane The chorioallantoic membrane (CAM), also known as the chorioallantois, is a highly vascularized membrane found in the eggs of certain amniotes like birds and reptiles. It is formed by the fusion of the mesodermal layers of two extra-embryonic memb ...

. Together these form a double membrane, which functions to remove carbon dioxide and to replenish oxygen through the porous shell. At the time of hatching, the fetus becomes detached from the chorion as it emerges from the shell. In

s, it develops by the follicle cells while the

egg An egg is an organic vessel grown by an animal to carry a possibly fertilized egg cell (a zygote) and to incubate from it an embryo within the egg until the embryo has become an animal fetus that can survive on its own, at which point the ...

is in the

ovary The ovary () is a gonad in the female reproductive system that produces ova; when released, an ovum travels through the fallopian tube/ oviduct into the uterus. There is an ovary on the left and the right side of the body. The ovaries are end ...

.Chapman, R.F. (1998) "The insects: structure and function", Section ''The egg and embryology''. Previewed in Google Book

on 26 Sep 2009. Some

mollusks Mollusca is a phylum of protostomic invertebrate animals, whose members are known as molluscs or mollusks (). Around 76,000 extant species of molluscs are recognized, making it the second-largest animal phylum after Arthropoda. The num ...

also have chorions as part of their eggs. For example, fragile

octopus An octopus (: octopuses or octopodes) is a soft-bodied, eight-limbed mollusc of the order Octopoda (, ). The order consists of some 300 species and is grouped within the class Cephalopoda with squids, cuttlefish, and nautiloids. Like oth ...

eggs have only a chorion as their envelope. “The Octopoda are characterized by eggs that have only a chorion as an envelope”https://www.sciencedirect.com/topics/agricultural-and-biological-sciences/octopoda

Additional images

Image:Gray21.png, Section through the embryo. Image:Gray24.png, Diagram of the human embryo. Image:Gray25.png, Diagram illustrating early formation of

and differentiation of body-stalk. Image:Gray26.png, Diagram showing later stage of allantoic development with commencing constriction of the yolk-sac. Image:Gray39.png, Scheme of placental circulation.

References

External links

* — "Female Reproductive System: placenta, chorionic plate"
McGill
{{Authority control Amniotic sac Embryology