Placentation is the formation, type and structure, or modes of arrangement of the

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 ...

. The function of placentation is to transfer nutrients, respiratory gases, and water from maternal tissue to a growing

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 ...

, and in some instances to remove waste from the embryo. Placentation is best known in live-bearing

mammals A mammal () is a vertebrate animal of the 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 middle e ...

(

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 ...

), but also occurs in some fish, reptiles, amphibians, a diversity of invertebrates, and

flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Ancient Greek, Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed with ...

s. In vertebrates, placentas have evolved more than 100 times independently, with the majority of these instances occurring in

squamate Squamata (, Latin ''squamatus'', 'scaly, having scales') is the largest Order (biology), order of reptiles; most members of which are commonly known as Lizard, lizards, with the group also including Snake, snakes. With over 11,991 species, it i ...

reptiles. The placenta can be defined as an organ formed by the sustained apposition or fusion of fetal membranes and parental tissue for physiological exchange. This definition is modified from the original Mossman (1937) definition, which constrained placentation in animals to only those instances where it occurred in the uterus.

In mammals

In live bearing mammals, the placenta forms after the embryo implants into the wall of 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 ...

. 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 ...

is connected to the placenta via an

umbilical cord In Placentalia, placental mammals, the umbilical cord (also called the navel string, birth cord or ''funiculus umbilicalis'') is a conduit between the developing embryo or fetus and the placenta. During prenatal development, the umbilical cord i ...

. Mammalian placentas can be classified based on the number of tissues separating the maternal from the fetal blood. These include: ; endotheliochorial placentation : In this type of placentation, 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 ...

are in contact with the endothelium of maternal blood vessels. (e.g. in most

carnivore A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant Plants are the eukaryotes that form the Kingdom (biology), kingdom Plantae; they ar ...

s like

cat The cat (''Felis catus''), also referred to as the domestic cat or house cat, is a small domesticated carnivorous mammal. It is the only domesticated species of the family Felidae. Advances in archaeology and genetics have shown that the ...

s and

dog The dog (''Canis familiaris'' or ''Canis lupus familiaris'') is a domesticated descendant of the gray wolf. Also called the domestic dog, it was selectively bred from a population of wolves during the Late Pleistocene by hunter-gatherers. ...

s) ; epitheliochorial placentation : Chorionic villi, growing into the apertures of uterine glands ( epithelium). (e.g. in

ruminant Ruminants are herbivorous grazing or browsing artiodactyls belonging to the suborder Ruminantia that are able to acquire nutrients from plant-based food by fermenting it in a specialized stomach prior to digestion, principally through microb ...

horse The horse (''Equus ferus caballus'') is a domesticated, one-toed, hoofed mammal. It belongs to the taxonomic family Equidae and is one of two extant subspecies of ''Equus ferus''. The horse has evolved over the past 45 to 55 mi ...

whale Whales are a widely distributed and diverse group of fully Aquatic animal, aquatic placental mammal, placental marine mammals. As an informal and Colloquialism, colloquial grouping, they correspond to large members of the infraorder Cetacea ...

s, lower primates,

dugong The dugong (; ''Dugong dugon'') is a marine mammal. It is one of four living species of the order Sirenia, which also includes three species of manatees. It is the only living representative of the once-diverse family Dugongidae; its closest ...

s) ; hemochorial placentation : In hemochorial placentation maternal blood comes in direct contact with the fetal

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 ...

, which it does not in the other two types. It may avail for more efficient transfer of nutrients etc., but is also more challenging for the systems of

gestational immune tolerance Immune tolerance in pregnancy or maternal immune tolerance is the immune tolerance shown towards the fetus and placenta during pregnancy. This tolerance counters the immune response that would normally result in the rejection of something foreign i ...

to avoid rejection of the fetus. (e.g. in higher order primates, including

human Humans (''Homo sapiens'') or modern humans are the most common and widespread species of primate, and the last surviving species of the genus ''Homo''. They are Hominidae, great apes characterized by their Prehistory of nakedness and clothing ...

s, and also in

rabbit Rabbits are small mammals in the family Leporidae (which also includes the hares), which is in the order Lagomorpha (which also includes pikas). They are familiar throughout the world as a small herbivore, a prey animal, a domesticated ...

guinea pig The guinea pig or domestic guinea pig (''Cavia porcellus''), also known as the cavy or domestic cavy ( ), is a species of rodent belonging to the genus ''Cavia'', family Caviidae. Animal fancy, Breeders tend to use the name "cavy" for the ani ...

mice A mouse (: mice) is a small rodent. Characteristically, mice are known to have a pointed snout, small rounded ears, a body-length scaly tail, and a high breeding rate. The best known mouse species is the common house mouse (''Mus musculus' ...

, and

rats Rats are various medium-sized, long-tailed rodents. Species of rats are found throughout the order Rodentia, but stereotypical rats are found in the genus ''Rattus''. Other rat genera include ''Neotoma'' (pack rats), '' Bandicota'' (bandicoot ...

) 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 ...

, placentation is the formation and growth of the placenta inside the uterus. It occurs after the implantation of the embryo into the uterine wall and involves the remodeling of

blood vessel Blood vessels are the tubular structures of a circulatory system that transport blood throughout many Animal, animals’ bodies. Blood vessels transport blood cells, nutrients, and oxygen to most of the Tissue (biology), tissues of a Body (bi ...

s in order to supply the needed amount of blood. In humans, placentation takes place 7–8 days after fertilization. In humans, the placenta develops in the following manner. Chorionic villi (from the embryo) on the embryonic pole grow, forming

chorion frondosum 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 chori ...

. Villi on the opposite side (abembryonic pole) degenerate and form the chorion laeve (or chorionic laevae), a smooth surface. The endometrium (from the mother) over the chorion frondosum (this part of the endometrium is called the decidua basalis) forms the decidual plate. The decidual plate is tightly attached to the chorion frondosum and goes on to form the actual placenta. Endometrium on the opposite side to the decidua basalis is the decidua parietalis. This fuses with the chorion laevae, thus filling up the uterine cavity. In the case of

twin 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 ...

s, dichorionic placentation refers to the presence of two placentas (in all

dizygotic 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 ...

and some

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 ...

twins). Monochorionic placentation occurs when monozygotic twins develop with only one placenta and bears a higher risk of complications during pregnancy. Abnormal placentation can lead to an early termination of pregnancy, for example in

pre-eclampsia Pre-eclampsia is a multi-system disorder specific to pregnancy, characterized by the new onset of hypertension, high blood pressure and often a significant amount of proteinuria, protein in the urine or by the new onset of high blood pressure a ...

Source-tissue types

Placenta can also be divided according to what kind of structure it develops from. There are two vessel-rich features in the

amniote Amniotes are tetrapod vertebrate animals belonging to the clade Amniota, a large group that comprises the vast majority of living terrestrial animal, terrestrial and semiaquatic vertebrates. Amniotes evolution, evolved from amphibious Stem tet ...

, the

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

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 ...

. When the chorion fuses with the former, the result is a

choriovitelline placenta A choriovitelline placenta is a placenta formed by the yolk sac and chorion. In a choriovitelline placenta, the yolk sac fuses with the chorion and, subsequently, wrinkles develop that hold the embryo to the uterine wall, thus forming the choriovit ...

. When it fuses with the latter, the result is a chorioallantoic placenta. Most mammals first form a temporaty choriovitelline placenta, then the chorioallantoic placenta takes over. (Primates do not form a definite choriovitelline placenta by fusion, but strong expression conservation suggest that the yolk sac remains useful.) Marsupials mostly have choriovitelline placental tissue. Rodents maintain both types throughout gestation.

In lizards and snakes

As placentation often results during the evolution of live birth, the more than 100 origins of live birth in lizards and snakes (

Squamata Squamata (, Latin ''squamatus'', 'scaly, having scales') is the largest Order (biology), order of reptiles; most members of which are commonly known as Lizard, lizards, with the group also including Snake, snakes. With over 11,991 species, it i ...

) have seen close to an equal number of independent origins of placentation. This means that the occurrence of placentation in Squamata is more frequent than in all other vertebrates combined, making them ideal for research on the evolution of placentation and viviparity itself. In most squamates, two separate placentae form, utilising separate embryonic tissue (the chorioallantoic and yolk-sac placentae). In species with more complex placentation, we see regional specialisation for gas, amino acid, and lipid transport. Placentae form following implantation into uterine tissue (as seen in mammals) and formation is likely facilitated by a plasma membrane transformation. Most reptiles exhibit strict epitheliochorial placentation (e.g. ''

Pseudemoia entrecasteauxii ''Pseudemoia entrecasteauxii'', also known commonly as Entrecasteaux's skink, the southern grass skink, the tussock cool-skink, and the tussock skink, is a species of lizard in the family Scincidae. The species is endemic to Australia. Geograp ...

)'' however at least two examples of endotheliochorial placentation have been identified ( ''Mabuya'''' sp.'' and '' Trachylepis ivensi''). Unlike eutherian mammals, epitheliochorial placentation is not maintained by maternal tissue as embryos do not readily invade tissues outside of the uterus.

Research

The placenta is an

organ Organ and organs may refer to: Biology * Organ (biology), a group of tissues organized to serve a common function * Organ system, a collection of organs that function together to carry out specific functions within the body. Musical instruments ...

that has evolved multiple times independently, evolved relatively recently in some lineages, and exists in intermediate forms in living species; for these reasons it is an outstanding model to study the evolution of complex organs in animals. Research into the genetic mechanisms that underpin the evolution of the placenta have been conducted in a diversity of animals including reptiles, seahorses, and mammals. The genetic processes that support the evolution of the placenta can be best understood by separating those that result in the evolution of new structures within the animal and those that result in the evolution of new functions within the placenta.

Evolution of placental structures

In all placental animals, placentas have evolved through the utilisation of existing tissues. In viviparous mammals and reptiles placentas form from the intimate interaction of the uterus and a series of embryonic membranes including the chorioallantoic and yolk sac membranes. In guppies placental tissues form between the ovarian tissue and the egg membrane. In pipefish placentas form following the interaction with the egg and the skin. Despite the placenta forming from pre-existing tissues, in many instances new structures can evolve within these pre-existing tissues. For example, in male seahorses the underbelly skin has become highly modified to form a pouch in which embryos can develop. In mammals and some reptiles, including the viviparous southern grass skink, the uterus becomes regionally specialised to support placental functions, within each of these regions being a new specialised uterine structure. In the southern grass skink three distinct regions of the placenta form which likely perform different functions; the placentome supports nutrient transfer via membrane bound transport proteins, the paraplacentome supports the exchange of respiratory gasses, and the yolk sac placenta supports lipid transport via apocrine secretion.

Evolution of placental functions

Placental functions include nutrient transport, gas exchange, maternal-fetal communication, and waste removal from the embryo. These functions have evolved by a series of general processes such as re-purposing processes found in the ancestral tissues from which a placenta is derived, recruiting the expression of genes expressed elsewhere in the organism to perform new functions in placental tissues, and the evolution of new molecular processes following the formation of new placenta specific genes. In mammals, maternal-fetal communication occurs via the production of a range of signalling molecules and their receptors in the chorioallantoic membrane of the embryo and the endometrium of the mother. Examination of these tissues in egg-laying and other independently evolved live bearing vertebrates has shown us that many of these signalling molecules are expressed widely in vertebrate species and were probably expressed in the ancestral amniote vertebrate. This suggests that maternal fetal communication has evolved by utilising the existing signalling molecules and their receptors, from which placental tissues are derived.

In plants

s, placentation is the attachment of

ovule In seed plants, the ovule is the structure that gives rise to and contains the female reproductive cells. It consists of three parts: the ''integument'', forming its outer layer, the ''nucellus'' (or remnant of the sporangium, megasporangium), ...

s inside 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 ...

."Flowers" At: ''Botany Online'' At: University of Hamburg Department of Biology. (see ''External links'' below). The ovules inside a

flower Flowers, also known as blooms and blossoms, are the reproductive structures of flowering plants ( angiosperms). Typically, they are structured in four circular levels, called whorls, around the end of a stalk. These whorls include: calyx, m ...

's ovary (which later become the

seed In botany, a seed is a plant structure containing an embryo and stored nutrients in a protective coat called a ''testa''. More generally, the term "seed" means anything that can be Sowing, sown, which may include seed and husk or tuber. Seeds ...

s inside a

fruit In botany, a fruit is the seed-bearing structure in flowering plants (angiosperms) that is formed from the ovary after flowering. Fruits are the means by which angiosperms disseminate their seeds. Edible fruits in particular have long propaga ...

) are attached via ''funiculi'', the plant part equivalent to an umbilical cord. The part of the ovary where the funiculus attaches is referred to as the ''placenta''. In botany, the term placentation most commonly refers to the arrangement of ovules inside an ovary. Placentation types include: *Basal: The placenta is found in mono to multi carpellary, syncarpous ovary. Usually a single ovule is attached at the base (bottom). It is unilocular(Single chamber). E.g.: ''Helianthus'', ''Tridex'', ''Tagetus''. *Parietal: It is found in bicarpellary to multicarpellary syncarpous ovary. Unilocular ovary becomes bilocular due to formation of false septum.E.g.: Cucumber *Axile : it is found in bicarpellary to multicarpellary syncarpous ovary. The carpels fuse to form septa forming a central axis and ovules are arranged on the axis. E.g.: Hibiscus, lemon, tomato, lilly. *Apical: The placenta is at the apex (top) of the ovary. Simple or compound ovary. *Superficial: Similar to axile, but placentae are on inner surfaces of multilocular ovary (e.g. Nymphaea) *Free central : It is found in bicarpellary to multicarpellary syncarpous ovary. Due to degradation of false septum unilocular condition is formed and ovules are arranged on the central axis. E.g.: ''

Dianthus ''Dianthus'' ( ) is a genus of about 340 species of flowering plants in the family Caryophyllaceae, native mainly to Europe and Asia, with a few species in north Africa and in southern Africa, and one species (''D. repens'') in arctic North Am ...

'', ''

Primula ''Primula'' () is a genus of herbaceous plant, herbaceous flowering plants in the family (biology), family Primulaceae. They include the primrose (''Primula vulgaris, P. vulgaris''), a familiar wildflower of banks and verges. Other common specie ...

'' (primroses) *Marginal : It is found in monocarpellary unilocular ovary, placenta forms a rigid along ventral side and ovules are arranged in two vertical rows. E.g.: ''

Pisum sativum Pea (''pisum'' in Latin) is a pulse or fodder crop, but the word often refers to the seed or sometimes the pod of this flowering plant species. Peas are eaten as a vegetable. Carl Linnaeus gave the species the scientific name ''Pisum sativum ...

'' (pea) File:Gynoecium morphology placentation basal longitudinal section.png, Basal File:Gynoecium morphology placentation parietal-1 cross section.png, Parietal File:Gynoecium morphology placentation axile-2 cross section.png, Axile File:Gynoecium morphology placentation free-central cross section.png, Free central File:Gynoecium morphology placentation marginal longitudinal section.png, Marginal