Extra Embryonic Membranes Of Vertebrates

The embryos of reptiles, birds, and mammals produce 4 extraembryonic membranes. They are: a) b) c) d) amnion yolk sac chorion, and allantois.

In birds and most reptiles, the embryo with its extraembryonic membranes develops within a shelled egg. With these four membranes, the developing embryo is able to carry on essential metabolism while sealed within the egg. Surrounded by amniotic fluid, the embryo is kept as moist as a fish embryo in a pond. Although (most) mammals do not make a shelled egg, they do also enclose their embryo in an amnion. For this reason, the reptiles, birds, and mammals are collectively referred to as the amniota. In the embryonic development of vertebrates only a part of the egg or of cleavage mass of cells forms the actual embryo. The outer embryonic layers develop into extra-embryonic regions called embryonic or foetal membranes. Embryonic membranes are auxillary organs to protect the embryo and to provide its nutrition, respiration and excretion functions until the embryo develops fully and is capable of performing by its own. Four foetal or extra-embryonic membranes namely amnion, chorion, allantois and yolk-sac are formed around developing embryo of higher vertebrates. Their mode of formation and morphology may slightly alter in various animals.

Amnion and Chorion: Both these membranes develop simultaneously. The amnion protects the embryo in a sac filled with amniotic fluid. Development of amnion and chorion starts after gastrulation and neuralation in the form of folds of somatopleure which is somatic mesoderm and ectoderm. These folds are called amniotic folds and they arise from anterior or cephalic end as head folds and caudal end as tail folds which finally fuse at the mid dorsal part of the embryo. The inner somatopleuric layer becomes the amnion and the outer layer as chorion.

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The cavity between amnion and embryo is called amniotic cavity and that between amnion and chorion is called chorionic cavity or extra embryonic coelom. Amniotic cavity contains fluid called amniotic fluid. It cushions and acts as shock absorber to the developing embryo. It maintains constant temperature and prevents adhesion of foetal parts and amniotic sac. In viviparous animals it plays an important role in delivery of the young ones. Amnion is regarded as the membranous tent around the embryo. The chorion along with allantois serves as a respiratory organ in oviparous forms and in viviparous animals it involves in the formation of placenta. The chorion lines the inner surface of the shell (which is permeable to gases) and participates in the exchange of O2 and CO2 between the embryo and the outside air.

Mammals fall into three groups that differ in the way they use the amniotic egg. Monotremes These primitive mammals produce a shelled egg like their reptilian ancestors. Only four species exist today: three species of spiny anteater (echidna) and the duckbill platypus.

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Marsupials Marsupials do not produce a shelled egg. The egg, which is poorly supplied with yolk, is retained for a time within the reproductive tract of the mother. The embryo penetrates the wall of the uterus. The yolk sac provides a rudimentary connection to the mother's blood supply from which it receives food, oxygen, and other essentials. However, this interface between the tissues of the uterus and the extraembryonic membranes never becomes elaborately developed, and the young are born in a very immature state. The photo (courtesy of Dr. Carl G. Hartman) shows 18 newborn baby opossums fitting easily into a teaspoon. Despite their tiny size, they are able to crawl into a pouch on the mother's abdomen, attach themselves to nipples, and drink milk from her mammary glands. Marsupials are still abundant in Australia, but only the opossum is found in North America. Placental mammals In placental mammals, the extraembryonic membranes form a placenta and umbilical cord, which connect the embryo to the mother's uterus in a more elaborate and efficient way. The blood supply of the developing fetus is continuous with that of the placenta. The placenta extracts food and oxygen from the uterus. Carbon dioxide and other wastes (e.g., urea) are transferred to the mother for disposal by her excretory organs.

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Humans are placental mammals.

Allantois: The allantois stores metabolic wastes (chiefly uric acid) of the embryo and, as it grows larger, also participates in gas exchange. It arises from the hindgut as small sac and later spreads and grows as an extra-embryonic membrane over the embryo outside the amnion. The mesodermal lining of the alantois fuse with mesoderm of chorion to form allantochorion which give rise to blood vascular system in mammals. In oviparous forms the allantois spreads between amnion and yolk sac on one side and chorion on the other side. The mesoderm of allantois fuses with mesoderm of chorion to form chorio-allantois layer. This penetrates through the vitelline membrane and envelops the albumin sac and helps in the absorption of albumin and water. The distal part of the allantois is connected to the hind gut of the embryo by means of a allantoic stalk. When allantoic stalk is surrounded by yolk stalk it is called umbilical cord. It is a store house of nitrogenous wastes formed in the embryo. It also acts as embryonic lung. In viviparous forms allantoic circulation supplies the embryo with oxygen and nutrients absorbed from the maternal blood.

Yolk sac In mammals endoderm spreads over and encloses the cavity of yolk sac. It has respiratory, haemopoetic and storage function. The yolk sac contains yolk the sole source of food until hatching. Yolk is a mixture of proteins and lipoproteins. Along with allantois it forms yolk sac placenta.and umbilical stalk. In oviparous forms the extra embryonic splanchnopleure spreads and encloses the yolk to form yolk sac. The yolk is connected to the gut by a yolk stalk. Yolk sac like liver acts as haemopoetic organ and also performs biochemical functions. Albumin is enclosed at the distal end of the yolk sac and is supplied to the embryo.

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* The Monotremes Monotremes are a small but remarkable group of mammals that consists of a single species of duckbill platypus (Ornithorhynchus anatinus) found in Australia and three (perhaps four) species of spiny anteaters (echidnas) found in Australia and New Guinea. These animals retain several traits of their therapsid an cestors including

a cloaca the final segment of the digestive tract into which both the urinary and reproductive tracts empty (monotreme = single hole); lay shelled eggs that undergo merobastic cleavage like that of reptiles (and birds) rather than the holoblastic cleavage of all other mammals.

Despite these reptilian features, the monotremes meet all the criteria of true mammals: milk secreted from mammary glands (but no nipples); hair; teeth (only in the young; they are lost in adult montremes).

In the May 8 issue of Nature, a consortium of gene sequencers reported the results of sequencing the complete genome of the platypus. They identified 18,527 protein-coding genes distributed on 52 chromosomes. The mix of mammalian and reptilian phenotypic features turns out to be reflected in the genome as well. Examples: The platypus has genes for egg yolk proteins that are also found in birds but not in therians. The gene content of their X chromosomes resembles that of the Z chromosome in birds, not the X chromosome of other mammals like us.

Other features of their genome reflect their unique biology: The platypus produces a venom with genes which in other mammals encode for antimicrobial peptides called defensins. The platypus has some 1000 genes for receptors in its vomeronasal organ far more than found in other mammals. The platypus hunts for food underwater and probably uses these receptors to detect prey (as well as using its electroreceptors for this purpose).

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