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.

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.

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)."

During the 1870s and early 1880s, the British morphologist Francis Maitland Balfour contributed in important ways to the budding field of evolutionary embryology, especially through his comparative embryological approach to uncovering ancestral relationships between groups. As developmental biologist and historian Brian Hall has observed, the field of evolutionary embryology in the nineteenth century was the historical ancestor of modern-day evolutionary developmental biology. Balfour's work was notably inspired by Charles Darwin's theory of evolution and Ernst Haeckel's account of the relationships between embryology and evolution. Only a decade after Balfour's program of research began, an alpine climbing accident robbed Britain of its most promising embryologist.

A germ layer is a group of cells in an embryo that interact with each other as the embryo develops and contribute to the formation of all organs and tissues. All animals, except perhaps sponges, form two or three germ layers. The germ layers develop early in embryonic life, through the process of gastrulation. During gastrulation, a hollow cluster of cells called a blastula reorganizes into two primary germ layers: an inner layer, called endoderm, and an outer layer, called ectoderm. Diploblastic organisms have only the two primary germ layers; these organisms characteristically have multiple symmetrical body axes (radial symmetry), as is true of jellyfish, sea anemones, and the rest of the phylum Cnidaria. All other animals are triploblastic, as endoderm and ectoderm interact to produce a third germ layer, called mesoderm. Together, the three germ layers will give rise to every organ in the body, from skin and hair to the digestive tract.

In 1962 the journal Acta Biotheoretica published the final work of the biologist Edward Stuart Russell, a full eight years after his death. Entitled The Diversity of Animals: an Evolutionary Study, this short, unfinished manuscript on evolution received little recognition in the scientific presses despite both its technical discussion of adaptations in decapods (crabs, shrimp, etc.) and its different approach to evolutionary theory. The precise reason for this neglect is unclear. This book is a continuation of Russell's philosophical perspective, organicism, an interpretation that focuses on the organism as the primary unit of analysis for the biological sciences. Russell first argued for this position in several of his earlier works, such as The Interpretation of Development and Heredity (1930) and The Directiveness of Organic Activities (1946). What was new in The Diversity of Animals lies in Russell's orthogenetic theory of evolution. By "orthogenetic" he means evolutionary change in definite directions. The overall thesis of this work is that transformations in evolution that occur in early ontogenesis, or development, are the best explanation for most diversity in nature. The consequence of Russell's argument is that an understanding of development is fundamental to an explanation of the major transformations in the evolutionary history of life.

Between February 1969 and August 1970 Edward Kollar and Grace Baird, from the University of Chicago in Chicago, Illinois, published three papers that established the role of the mesenchyme in tooth induction. Drawing upon a history of using tissue interactions to understand differentiation, Kollar and Baird designed their experiments to understand how differentiated structures become specified. Their work overturned a widely accepted model that epithelium controls the identity of the structure, a phenomenon called structural specificity. Interactions between epithelium and mesenchyme control the development and differentiation of many parts during embryonic development, including structures like the gastrointestinal tract and hair. Thus, the realization that mesenchyme drives induction and differentiation during epithelio-mesenchymal interactions had far-reaching effects.