Placenta

Placenta (Lat. for a cake), in anatomy, the organ by which the embryo is nourished within the womb of its mother. When the young one is born the placenta and membranes come away as the "afterbirth." In human anatomy the organ is a circular disk about seven or eight inches in diameter and one and a quarter inches in thickness at its centre, while at its margin it is very thin and is continuous with the foetal membranes. It weighs about a pound.

In order to explain the formation of the placenta it is necessary to encroach to some extent on the domain of physiology. Before each menstrual period, during the child-bearing age of a woman, the mucous membrane of the uterus hypertrophies, and, at the period, is cast off and renewed, but if a fertilized ovum reaches the uterus the casting off is postponed until the birth of the child. From the fact that the thickened mucous membrane lining the interior of the uterus is cast off sooner or later, it is spoken of as the "decidua." The fertilized ovum, on reaching the uterus, sinks into and embeds. itself in the already prepared decidua, and, as it enlarges, there is, one part of the decidua lying between it and the uterine wall ("decidua serrotina" or "basalis"), one part stretched over the surface of the enlarging ovum ("decidua reflexa" or "capsularis") and one part lining the rest of the uterus ("decidua vera") (see fig I.).

Decidua basalis.

It is the decidua basalis which is specially interesting in considering the formation of the placenta. That part which is nearest the ovum is called the "stratum compactum," but farther away the uterine glands dilate and give a spongy appearance to the mucous membrane which earns this particular layer the name of "stratum spongiosum." Processes grow out from the surface of the ovum which penetrate thestratum compactum of the decidua basalis and capsularis and push their way into the enlarged maternal blood sinuses; these are the "chorionic villi." Later, the "allantoic" or "abdominal stalk" grows from the mesoderm of the hind end of the embryo into the chorionic villi which enter the decidua basalis, and in this bloodvessels pass which push their way into the maternal blood sinuses. Eventually the original walls of these sinuses, together with the false amnion, disappear, and nothing now separates the maternal from the foetal blood except the delicate walls of the foetal vessels covered by some nucleated noncellular tissue, known as syncytium, derived from the chorionic epithelium, so that the embryo is able to take its supply of oxygen and materials for growth from the blood of its mother and to give up carbonic acid and excretory matters. It is the gradual enlargement of the chorionic villi in the decidua basalis together with the intervillous maternal blood sinuses that forms the placenta; the decidua capsularis and vera eventually become pressed From A. H. Young and A. Robinson, in Cunningham's Text-Book of Anatomy.  FIG. I. - Diagram representing a very young human ovum almost immediately after its entrance into the decidua, and whilst the place of its entrance is still covered with a plug of fibrin. The ectoderm has already proliferated and embraced spaces which contain maternal blood and are continuous with the maternal blood-vessels.

Uterine mucosa..

Gland.

Dilated p Unchanged pa Unchanged layer.

Stratum spongiosum. Stratum compactum.

Ectodermal villus enclosing space containing maternal blood.

Cavity which becomes coelom.

Fibrin plug. Decidua vera.

Inner mass (Entoderm)Decidua vera.

Cavity of Uterus.

From A. H. Young and A. Robinson, in Cunningham's Text-Book of Anatomy.  FIG 2. - Diagram. Later stage in the development of the placenta, showing the relations of the foetal villi to the placental sinuses, the fusion of the amnion with the inner surface of the chorion, and the thinning of the fused deciduae (capsularis and vera).

together as the embryo enlarges, and then, as pressure continues, atrophy. The allantoic stalk elongates enormously, and in its later stages contains two arteries (umbilical) and only one vein (owing to the obliteration of the right one) embedded in some loose connective tissue known as "Wharton's jelly." At first the stalk of the yolk-sac is quite distinct from this, but later the two structures become bound up together (see fig. 2), after which they are known as the "umbilical cord." A distinction must be made between the allantoic stalkand the allantois; the latter is an entodermal outgrowth from the hind end of the mesodaeum or primitive alimentary canal, which in the human subject only reaches a little way toward the placenta. The allantoic stalk is the mass of mesoderm containing blood-vessels which is pushed in front of the allantois and, as has been shown, reaches and blends with the decidua basalis to form the placenta. For further details see Quain's Anatomy, vol. i. (London, 1908); and, for literature, O. Hertwig's Handbuch der Entwickelungslehre  (Jena).

## Comparative Anatomy

If the placenta is to be regarded as a close union between the vascular system of the parent and embryo, the condition may be found casually scattered throughout the phylum of the Chordata. In such a very lowly member of the Placenta.

Unchanged layer.

Placental villus.

phylum as Saipa, a placenta is formed, and the embryo is nourished within the body of its parent. In some of the viviparous sharks, e.g. the blue shark (Carcharias), the yolk-sac has ridges which fit into grooves in the wall of the oviduct and allow an interchange of materials between the maternal and foetal blood. This is an example of an "umbilical placenta." In the viviparous blennies (Zoarces viviparus), among the teleostean fishes, two or three hundred young are nourished in the hollow ovary, which develops villi secreting nutritive material. Among the Amphibia the alpine salamander (Salamandra atra) nourishes its young in its oviducts until the gilled stage of development is past, while in the Reptilia the young of a viviparous lizard (Sees chalcides) establish a communication between the yolk-sac anteriorly and the allantois posteriorly, on the one hand, and the walls of the oviduct on the other. In this way both an umbilical and an allantoic placenta are formed.

The mammals are divided into Placentalia and Aplacentalia; in the latter group, to which the monotremes and most marsupials belong, the ova have a great deal of yolk, and the young, born in a very immature condition, finish their development in their mother's pouch; but although these mammals have no allantoic placenta there is an intimate connexion between the walls of the yolk-sac and the uterine mucous membrane, and so an umbilical or omphalic placenta exists. The name Aplacentalia therefore only means that they have no allantoic placenta. Among the Placentalia the umbilical and allantoic placentae sometimes coexist for some time, as in the case of the hedgehog, the bandicoot and the mouse. In most of the lower placental mammals the allantois is much more developed than in man, and the most primitive type of placenta is that in which villi are formed over the whole surface of the chorion projecting into the decidua of the tubular cornu of the uterus. This is known as a "diffuse placenta," and is met with in the pangolin, pig, hippopotamus, camel, chevrotain, horse, rhinoceros, tapir and whale. When the villi are collected into a number of round tufts or cotyledons, as in most ruminants, the type is spoken of as a "cotyledonous placenta," and an intermediate stage between this and the last is found in the giraffe.

In the Carnivora, elephant, procavia (Hyrax) and aard vark (Orycteropus), there is a "zonary-placenta" which forms a girdle round the embryo. In sloths and lemurs the placenta is domeshaped, while in rodents, insectivores and bats, it is a ventral disk or closely applied pair of disks, thus differing from the dorsal disk of the ant-eater, armadillo and higher Primates, which is known as a "metadiscoidal placenta." It will thus be seen that the form of the placenta is not an altogether trustworthy indication of the systemic position of its owner. In the diffuse and cotyledonous placentae the villi do not penetrate very deeply into the decidua, and at birth are simply withdrawn, the decidua being left behind in the uterus, so that these placentae are spoken of as non-deciduate while other kinds are deciduate.

For further details see S. W. W. Turner, Lectures on the Comparative Anatomy of the Placenta (Edinburgh, 1876); A.Robinson, "Mammalian Ova and the Formation of the Placenta," Journ. Anat. and Phys. (1904) xxxviii., 186, 325. For literature up to 1906, R. Wiedersheim's Comparative Anatomy of Vertebrates, translated and adapted by W. N. Parker (London, 1907).

(F. G. P.)