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 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.

Elizabeth Dexter Hay studied the cellular processes that affect development of embryos in the US during the mid-twentieth and early twenty-first centuries. In 1974, Hay showed that the extracellular matrix, a collection of structural molecules that surround cells, influences cell behavior. Cell growth, cell migration, and gene expression are influenced by the interaction between cells and their extracellular matrix. Hay also discovered a phenomenon later called epithelial-mesenchymal transition, a process that occurs during normal embryo and adult development in which epithelial cells, cells that line external and internal surfaces of the body, transform into mesenchymal stem cells, connective tissue cells that are capable of turning into other cell types. Hay's work helped researchers explain normal developmental processes and enabled research into abnormal processes that can cause developmental defects and diseases.

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.

Edmund Beecher Wilson contributed to cell biology, the study of cells, in the US during the end of the nineteenth and the beginning of the twentieth centuries. His three editions of The Cell in Development and Inheritance (or Heredity) in 1896, 1900, and 1925 introduced generations of students to cell biology. In The Cell, Wilson described the evidence and theories of his time about cells and identified topics for future study. He helped show how each part of the cell works during cell division and in every step of early development of an organism. Developmental biologists trained in the mid-twentieth century reported WilsonÕs text as their foundation for understanding biology, including about how cells, development, heredity, and evolution interact. Wilson considered cells as the center of all biological phenomena.

The Cell in Development and Inheritance, by Edmund Beecher Wilson, provided a textbook introduction to cell biology for generations of biologists in the twentieth century. In his book, Wilson integrated information about development, inheritance, chromosomes, organelles, and the structure and functions of cells. First published in 1896, the book started with 371 pages, grew to 483 pages in the second edition that appeared in 1900, and expanded to 1,231 pages by the third and final edition in 1925. Wilson dedicated the book to the cell biologist Theodor Boveri, whose work established the roles of chromosomes in cell division. With its explanations and many illustrations and diagrams, The Cell in Development and Inheritance enabled embryologists to better understand development in terms of cell structure and function.