Embryo Project Encyclopedia Articles

Conrad Hal Waddington's "Experiments on Embryonic Induction III," published in 1934 in the Journal of Experimental Biology, describes the discovery that the primitive streak induces the mammalian embryo. Waddington's hypothesis was that a transplanted primitive streak could induce neural tissue in the ectoderm of the rabbit embryo. The primitive streak defines the axis of an embryo and is capable of inducing the differentiation of various tissues in a developing embryo during gastrulation. In this experiment Waddington was, in fact, able to induce neural differentiation. Waddington noted that the tissue is "competent"; for a chick organizer, and by deduction a mammalian organizer must exist. Competence refers to a cell's ability to respond to an inducing signal, which is temporally limited to certain developmental stages. Waddington's initial work laid the foundation for many decades of research to follow, including further experiments by Waddington with the mammalian organizer.

Endoderm is one of the germ layers-- aggregates of cells that organize early during embryonic life and from which all organs and tissues develop. All animals, with the exception of sponges, form either two or three germ layers through a process known as gastrulation. During gastrulation, a ball of cells transforms into a two-layered embryo made of an inner layer of endoderm and an outer layer of ectoderm. In more complex organisms, like vertebrates, these two primary germ layers interact to give rise to a third germ layer, called mesoderm. Regardless of the presence of two or three layers, endoderm is always the inner-most layer. Endoderm forms the epithelium-- a type of tissue in which the cells are tightly linked together to form sheets-- that lines the primitive gut. From this epithelial lining of the primitive gut, organs like the digestive tract, liver, pancreas, and lungs develop.

The sex of a reptile embryo partly results from the production of sex hormones during development, and one process to produce those hormones depends on the temperature of the embryo's environment. The production of sex hormones can result solely from genetics or from genetics in combination with the influence of environmental factors. In genotypic sex determination, also called genetic or chromosomal sex determination, an organism's genes determine which hormones are produced. Non-genetic sex determination occurs when the sex of an organism can be altered during a sensitive period of development due to external factors such as temperature, humidity, or social interactions. Temperature-dependent sex determination (TSD), where the temperature of the embryo's environment influences its sex development, is a widespread non-genetic process of sex determination among vertebrates, including reptiles. All crocodilians, most turtles, many fish, and some lizards exhibit TSD.

The Cell-Theory was written by Thomas Henry Huxley in Britain and published in 1853 by The British and Foreign Medico-Chirurgical Review. The twenty-two page article reviews twelve works on cell theory, including those in Germany by Caspar Friedrich Wolff in the eighteenth century and by Karl Ernst von Baer in the nineteenth century. Huxley spends much of The Cell-Theory on a cell theory proposed in the late 1830s by Matthias Schleiden and Theodor Schwann in Germany. Schleiden and Schwann maintained that the cell was the most fundamental unit of life and that the nucleus was the most significant cellular component. Huxley, instead, promoted an epigenetic theory of the cell, for which properties of life emerge from the outer cytoplasm, cell membrane, and wall (the periplast), as opposed to the inner contents of the cell, including the nucleus (the endoplast). Huxley's arguments in The Cell-Theory influenced future scientists about the role of epigenetic processes in embryology and development.

Frederik Ruysch, working in the Netherlands, introduced the term epithelia in the third volume of his Thesaurus Anatomicus in 1703. Ruysch created the term from the Greek epi, which means on top of, and thele, which means nipple, to describe the type of tissue he found when dissecting the lip of a cadaver. In the mid nineteenth century, anatomist Albrecht von Haller adopted the word epithelium, designating Ruysch's original terminology as the plural version. In modern science, epithelium is a type of animal tissue in which cells are packed into neatly arranged sheets. The epithelial cells lie proximate to each other and attach to a thin, fibrous sheet called a basement membrane. Epithelia line the surfaces of cavities and structures throughout the body, and also form glands. Although they lack blood vessels, epithelia contain nerves and can function to receive sensation, absorb, protect, and secrete, depending on which part of the body the epithelia line. During development, epithelia act in conjunction with another tissue type, mesenchyme, to form nearly every organ in the body, from hair and teeth to the digestive tract. Epithelia are an essential part of embryonic development and the maintenance and function of the body throughout life.

Hilde Proscholdt Mangold was a doctoral student at the Zoological Institute at the University of Freiburg in Freiburg, Germany, from 1920-1923. Mangold conducted research for her dissertation 'On the Induction of Embryonic Primordia by Implantation of Organizers from Different Species' ('Ueber Induktion von Embryonanlagen durch Implantation artfremder Organisatoren'), under the guidance of Hans Spemann, a professor of zoology at the University of Freiburg. The dissertation was the culmination of five experiments on three species of newt embryos, of the genus Triton (presently, Triturus), performed during the summers of 1921 and 1922, which resulted in a confirmation of Spemann's organizer concept. Spemann and Mangold published the dissertation in a 1924 edition of Roux's Archives for Microscopic Anatomy and Developmental Mechanics (Roux's Archiv fur Mikroskopische Anatomie und Entwicklungsmechanik)."

In 1962 researcher John Bertrand Gurdon at the University of Oxford in Oxford, England, conducted a series of experiments on the developmental capacity of nuclei taken from intestinal epithelium cells of feeding tadpoles. In the experiments, Gurdon conducted nuclear transplantation, or cloning, of differentiated cells, or cells that have already specialized to become one cell type or another, in tadpoles. Gurdon's experiment showed that differentiated adult cells could be induced to an undifferentiated state, where they could once again become multiple cell types. Gurdon's experiment disproved the theory that differentiated cells could not be undifferentiated or dedifferentiated into a new type of differentiated cell. Gurdon's experiment demonstrated nuclear transplantation, also called cloning, using differentiated cells.

Eugen Steinach researched sex hormones and their effects on mammals in the late nineteenth and early twentieth centuries in Europe. He experimented on rats by removing their testicles and implanting them elsewhere in their bodies, and he found that the testes interstitial cells produce male sex hormones. He developed the Steinach Rejuvenation Procedure, which he claimed could rejuvenate men by increasing their production of sex hormones. Steinach’s work on female sex hormones and on ovarian extracts led to the development of the first standardized injectable estrogen. Steinach’s research on reproductive hormones helped researchers explain the roles of sex hormones and develop hormone drugs.

In 1996, the US Congress mandated that the US Environmental Protection Agency (EPA) create and regulate the Endocrine Disruptor Screening Program. The program tests industrial and agricultural chemicals for hormonal impacts in humans and in wildlife that may disrupt organisms' endocrine systems. The endocrine system regulates the release of small amounts of chemical substances called hormones to keep the body functioning normally. Some chemicals can impede the endocrine system's function by mimicking or blocking hormone reception, which can disrupt processes of development and reproduction and harm organisms. As of 2017, the Endocrine Disruptor Screening Program is the largest US program to identify and regulate chemicals that affect the normal production of sex hormones like estrogen and androgen, which can have long-term effects on development and reproduction.

In the early 1920s, researchers Edgar Allen and Edward Adelbert Doisy conducted an experiment that demonstrated that ovarian follicles, which produce eggs in mammals, also contain and produce what they called the primary ovarian hormone, later renamed estrogen. In their experiment, Doisy and Allen extracted estrogen from the ovarian follicles of hogs and proved that they had isolated estrogen by using a measurement later renamed the Allen-Doisy test. Allen and Doisy’s 1923 experiment to isolate estrogen showed it was made within the ovaries and also established a method for isolating the sex hormone. That method provided a basis for future research on hormones. Later researchers showed that estrogen functions in the menstrual cycles of primates by signaling for the tissue lining the uterus (endometrium) to thicken in preparation for possible implantation of a fertilized egg.