Embryo Project Encyclopedia

From a developing embryos three primary germ layers, ectoderm (green), mesoderm (pink) and endoderm (yellow), a variety of differentiated cell types and organ systems arise, far more than are shown here. The three primary germ layers are shown during the gastrula stage because they become distinct at the gastrula stage. The germ cells (blue) are pre- cursors to sperm and egg cells, and they are set aside early in development, and are thought to arise from the ectoderm.

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.

Tooth enamel contains relics of its formation process, in the form of microstructures, which indicate the incremental way in which it forms. These microstructures, called cross-striations and striae of Retzius, develop as enamel-forming cells called ameloblasts, whcih cyclically deposit enamel on developing teeth in accordance with two different biological clocks. Cross-striations result from a twenty-four hour cycle, called a Circadian rhythm, in the enamel deposition process, while striae of Retzius have a longer periodicity. Unlike other tissues, enamel does not remodel after it forms, leaving those microstructures intact after deposition. Cross-striations and striae of Retzius thus provide evidence of the timing and processes of tooth development, and they indicate how organisms in a lineage differently grow and develop across generations. Researchers have examined those microstructures to investigate human evolution.

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.

Frank R. Lillie was born in Toronto, Canada, on 27 June 1870. His mother was Emily Ann Rattray and his father was George Waddell Little, an accountant and co-owner of a wholesale drug company. While in high school Lillie took up interests in entomology and paleontology but went to the University of Toronto with the aim of studying ministry. He slowly became disillusioned with this career choice and decided to major in the natural sciences. It was during his senior year that he developed his lifelong interest in embryology. Graduating with a BA in 1891 Lillie then moved to the Marine Biological Laboratory (MBL) at Woods Hole, Massachusetts, to work and study with Charles Otis Whitman, the founding director of the MBL. Lillie collected and studied cell lineage side-by-side with some of the most prominent embryologists of the time: Edmund B. Wilson, Edwin G. Conklin, and Aaron L. Treadwell. Along with his cell lineage studies, Whitman guided Lillie to work on the question of how blastomeres contributed to the formation of organs in fresh water clams.

Ernest Everett Just was an early twentieth century American experimental embryologist involved in research at the Marine Biological Laboratory (MBL) at Woods Hole, Massachusetts, and the Stazione Zoologica in Naples, Italy. Just was known for simple but elegant experiments that supported the "fertilizing" theory of Frank R. Lillie and served as an antagonist to Jacques Loeb's work with artificial parthenogenesis. Just's many experiments with marine invertebrates showed that the egg surface, or ectoplasm, plays an important role in the fertilization and development of eggs.

In nineteenth century Great Britain, Thomas Henry Huxley proposed connections between the development of organisms and their evolutionary histories, critiqued previously held concepts of homology, and promoted Charles Darwin's theory of evolution. Many called him Darwin's Bulldog. Huxley helped professionalize and redefine British science. He wrote about philosophy, religion, and social issues, and researched and theorized in many biological fields. Huxley made several methodological contributions to both invertebrate and vertebrate embryology and development, and he helped shape the extra-scientific discourse for these fields.

Anatomical models have always been a mainstay of descriptive embryology. As the training of embryologists grew in the late 1800s, so too did the need for large-scale teaching models. Embryo wax models, such as those made by Adolf Ziegler and Gustav Born, were popular in the latter part of the nineteenth century and the early twentieth century as a way to visualize, in three dimensions, the fine detail of embryos without the aid of a microscope. While these models were found in many university laboratories, museums of science, and even expositions and world's fairs, they were anything but easy to make or obtain. Wax modeling required skill, patience, and specialized tools. Small laboratories with only one or two embryologists often found the prospect of wax modeling too laborious, too difficult, and too expensive to make the pursuit worthwhile. As an alternative, Susanna Phelps Gage, an embryologist at Cornell University, perfected a technique of using stacks of absorbent blotting paper rather than stacks of wax plates for constructing embryo models. She first demonstrated her blotting paper method to other embryologists at the annual meeting of the Association of American Anatomists in 1905 and later at the International Zoological Congress, held in Boston in August 1907.

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)."

Hans Spemann was an experimental embryologist best known for his transplantation studies and as the originator of the "organizer" concept. One of his earliest experiments involved constricting the blastomeres of a fertilized salamander egg with a noose of fine baby hair, resulting in a partially double embryo with two heads and one tail. Spemann continued changing variables such as the amount of time the embryo was constricted and the degree of constriction, all of which added more empirical evidence to Hans Driesch's studies showing that embryonic cells could self-regulate to varying degrees. Spemann's long list of "simple" experiments and significant findings were mainly carried out at his laboratory, the Spemann School at the University of Freiburg, Germany, where numerous graduate students collaborated with Spemann to investigate embryonic induction.