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. They typically perform roles in nutrition, gas exchange and waste removal. [1]
There are four standard extraembryonic membranes in amniotes, i.e. reptiles (including birds) and mammals:
In humans and other mammals they are more usually called fetal membranes.
The extraembryonic membranes in insects include a serous membrane (serosa) originating from blastoderm cells, an amnion or amniotic cavity whose expression is controlled by the Zerknüllt gene, and a yolk sac. [3]
A body cavity is any space or compartment, or potential space, in an animal body. Cavities accommodate organs and other structures; cavities as potential spaces contain fluid.
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 membranes that hold a developing embryo until shortly before birth. The inner of these membranes, the amnion, encloses the amniotic cavity, containing the amniotic fluid and the embryo. The outer membrane, the chorion, contains the amnion and is part of the placenta. On the outer side, the amniotic sac is connected to the yolk sac, the allantois, and via the umbilical cord, the placenta.
The amnion 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 environment for the developing embryo. The amnion, along with the chorion, the yolk sac and the allantois protect the embryo. In birds, reptiles and monotremes, the protective sac is enclosed in a shell. In marsupials and placental mammals, it is enclosed in a uterus.
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 Carboniferous period and further diverged into two groups, namely the sauropsids and synapsids, an event that marks the appearance of Amniota, according to the definition established under the PhyloCode. This basal divergence within Amniota has been dated by molecular studies at 310–329 Ma or 312–330 Ma, but the presence of Hylonomus at Joggins implies a minimal age of about 317 Ma. A fossilized birth-death process study of early amniotes suggested an age of 322–340 Ma. Amniotes are distinguished from the other living tetrapod clade — the non-amniote lissamphibians — by the development of three extraembryonic membranes, thicker and keratinized skin, and costal respiration. Additional unique features are the presence of adrenocortical and chromaffin tissues as a discrete pair of glands near their kidneys, which are more complex, the presence of an astragalus for better extremity range of motion, the diminished role of skin breathing, and the complete loss of metamorphosis, gills, and lateral lines.
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.
An egg is an organic vessel grown by an animal to carry a possibly fertilized egg cell 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 animal hatches.
The allantois is a hollow sac-like structure filled with clear fluid that forms part of a developing amniote's conceptus. It helps the embryo exchange gases and handle liquid waste.
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 only available structure that can be used to determine if an intrauterine pregnancy exists until the embryo can be identified.
The yolk sac is a membranous 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 is far more widely used. In humans, the yolk sac is important in early embryonic blood supply, and much of it is incorporated into the primordial gut during the fourth week of embryonic development.
In amniote embryonic development, the epiblast 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, the other layer is the hypoblast. It drives the embryo proper through its differentiation into the three primary germ layers, ectoderm, mesoderm and endoderm, during gastrulation. The amniotic ectoderm and extraembryonic mesoderm also originate from the epiblast.
The bilaminar embryonic disc, bilaminar blastoderm or embryonic disc is the distinct two-layered structure of cells formed in an embryo. In the development of the human embryo this takes place by day eight. It is formed when the inner cell mass, also known as the embryoblast, forms a bilaminar disc of two layers, an upper layer called the epiblast and a lower layer called the hypoblast, which will eventually form into fetus. These two layers of cells are stretched between two fluid-filled cavities at either end: the primitive yolk sac and the amniotic sac.
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 development. In biological terms, the development of the human body entails growth from a one-celled zygote to an adult human being. Fertilization occurs when the sperm cell successfully enters and fuses with an egg cell (ovum). The genetic material of the sperm and egg then combine to form the single cell zygote and the germinal stage of development commences. Human embryonic development covers the first eight weeks of development, which have 23 stages, called Carnegie stages. At the beginning of the ninth week, the embryo is termed a fetus. In comparison to the embryo, the fetus has more recognizable external features and a more complete set of developing organs.
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, which in turn gives rise to the chorion.
The anamniotes are an informal group of craniates comprising all fishes and amphibians, which lay their eggs in aquatic environments. They are distinguished from the amniotes, which can reproduce on dry land either by laying shelled eggs or by carrying fertilized eggs within the female. Older sources, particularly before the 20th century, may refer to anamniotes as "lower vertebrates" and amniotes as "higher vertebrates", based on the antiquated idea of the evolutionary great chain of being.
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 membranes that make up the amniotic sac which surrounds and protects the embryo. The fetal membranes are four of six accessory organs developed by the conceptus that are not part of the embryo itself, the other two are the placenta, and the umbilical cord.
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 membranes – the chorion and the allantois. It is the avian homologue of the mammalian placenta. It is the outermost extra-embryonic membrane which lines the non-vascular egg shell membrane.
The connecting stalk, or body stalk, is an embryonic structure that is formed by the third week of development and connects the embryo to its shell of trophoblasts. The connecting stalk is derived from the extraembryonic mesoderm. Initially it lies caudally to the trilaminar germ disc, but, with subsequent embryonic folding, the body stalk assume a more ventral position. Progressive expansion of the amnion from the umbilical ring creates a tube with a covering of amniotic membrane with allantois and umbilical vessels as its content and mesoderm of the connecting stalk as the ground substance. This extraembryonic mesodermal ground substance forms the future Wharton's jelly. The amniotic membrane and its contents form the umbilical cord that connects the embryo and the placenta.
The development of the digestive system in the human embryo concerns the epithelium of the digestive system and the parenchyma of its derivatives, which originate from the endoderm. Connective tissue, muscular components, and peritoneal components originate in the mesoderm. Different regions of the gut tube such as the esophagus, stomach, duodenum, etc. are specified by a retinoic acid gradient that causes transcription factors unique to each region to be expressed. Differentiation of the gut and its derivatives depends upon reciprocal interactions between the gut endoderm and its surrounding mesoderm. Hox genes in the mesoderm are induced by a Hedgehog signaling pathway secreted by gut endoderm and regulate the craniocaudal organization of the gut and its derivatives. The gut system extends from the oropharyngeal membrane to the cloacal membrane and is divided into the foregut, midgut, and hindgut.
Retinol-binding proteins (RBP) are a family of proteins with diverse functions. They are carrier proteins that bind retinol. Assessment of retinol-binding protein is used to determine visceral protein mass in health-related nutritional studies.
This glossary of developmental biology is a list of definitions of terms and concepts commonly used in the study of developmental biology and related disciplines in biology, including embryology and reproductive biology, primarily as they pertain to vertebrate animals and particularly to humans and other mammals. The developmental biology of invertebrates, plants, fungi, and other organisms is treated in other articles; e.g terms relating to the reproduction and development of insects are listed in Glossary of entomology, and those relating to plants are listed in Glossary of botany.