Embryo Project Encyclopedia

Mechanism of Notch Signaling: The image depicts a type of cell signaling, in which two animal cells interact and transmit a molecular signal from one to the other. The process results in the production of proteins, which influence the cells as they differentiate, move, and contribute to embryological development. In the membrane of the signaling cell, there is a ligand (represented by a green oval). The ligand functions to activate a change in a receptor molecule. In the receiving cell, there are receptors; in this case, Notch proteins (represented by orange forks). The Notch proteins are embedded in the receiving cell membrane, and they have at least two parts: an intracellular domain (inside the cell) and the receptor (outside the cell). Once the ligand and receptor bind to each other, a protease (represented by the dark red triangle) can sever the intracellular domain from the rest of the Notch receptor. Inside the nucleus of the receiving cell (represented by the gray area) are the cellês DNA (represented by the multi-colored helices) and its transcription factors (blue rectangles). Transcription factors are proteins that bind to DNA to regulate transcription, the first step in gene expression, which eventually yields proteins or other products. Initially, repressor proteins (represented by a red irregular hexagon) prevent transcription factors from allowing transcription. When the severed Notch receptor intracellular domain reaches the nucleus, it displaces the repressor. The transcription factor can then signal for transcription to occur. 1) There is a Notch receptor protein in the membrane of a receiving cell, and a ligand for this receptor (for example, Delta) in the membrane of the signaling cell. When the ligand binds to the receptor, the intracellular domain of the receptor changes shape. 2) Inside the receiving cell, there are proteases. Once the intracellular domain of the receptor changes shape, the protease can bind to it and shear the intracellular domain away from the rest of the receptor molecule. 3) The severed intracellular domain is shuttled to the receiving cell nucleus. Here, the intracellular domain displaces a repressor protein. This allows a transcription factor to initiate DNA transcription. During transcription, DNA is used as a template to create RNA. Following transcription, the process of translation occurs, which uses RNA as a template to create proteins. These proteins influence the behavior, fate, and differentiation of cells, which contribute to normal embryonic development

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.

'On the Permanent Life of Tissues outside of the Organism' reports Alexis Carrel's 1912 experiments on the maintenance of tissue in culture media. At the time, Carrel was a French surgeon and biologist working at the Rockefeller Institute in New York City. In his paper, Carrel reported that he had successfully maintained tissue cultures, which derived from connective tissues of developing chicks and other tissue sources, by serially culturing them. Among all the tissue cultures Carrel reported, one was maintained for more than two months, whereas previous efforts had only been able to keep tissues in vitro for three to fifteen days. Carrel’s experiments contributed to the development of long-term tissue culture techniques, which were useful in the study of embryology and eventually became instrumental in stem cell research. Despite later evidence to the contrary, Carrel believed that as long as the tissue culture method was accurately applied, tissues kept outside of the organisms should be able to divide indefinitely and have permanent life.

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.

Christiane Nusslein-Volhard studied how genes control embryonic development in flies and in fish in Europe during the twentieth and twenty-first centuries. In the 1970s, Nusslein-Volhard focused her career on studying the genetic control of development in the fruit fly Drosophila melanogaster. In 1988, Nusslein-Volhard identified the first described morphogen, a protein coded by the gene bicoid in flies. In 1995, along with Eric F. Wieschaus and Edward B. Lewis, she received the Nobel Prize in Physiology or Medicine for the discovery of genes that establish the body plan and segmentation in Drosophila. Nusslein-Volhard also investigated the genetic control of embryonic development to zebrafish, further generalizing her findings and helping establishing zebrafish as a model organism for studies of vertebrate development.

In 1969, Roy J. Britten and Eric H. Davidson published Gene Regulation for Higher Cells: A Theory, in Science. A Theory proposes a minimal model of gene regulation, in which various types of genes interact to control the differentiation of cells through differential gene expression. Britten worked at the Carnegie Institute of Washington in Washington, D.C., while Davidson worked at the California Institute of Technology in Pasadena, California. Their paper was an early theoretical and mechanistic description of gene regulation in higher organisms.

Of Sir D'Arcy Thompson's nearly 300 publications, the theoretical treatise On Growth and Form, first published in 1917, remains the principal work for which he is remembered. This substantial book is still in print today, and merited an editorial review and introductory essays by two important twentieth century biologists, John Tyler Bonner and Stephen Jay Gould. Growth and Form was immediately well-received for both its literary style and its scientific significance, as discussed by the biologist Sir Peter Medawar. Despite being almost continuously in print since its first publication, the exact influence of Growth and Form on the biological sciences, although widely acknowledged, is yet difficult to characterize. In this work Thompson aimed to unite physics and biology through an analysis of the physical limitations to the growth and structure of organisms. For developmental biologists in particular, Thompson's theory on the transformation of biological forms, presented in the final chapter of Growth and Form, was thought provoking.

Edward Stuart Russell was born 23 March 1887 to Helen Cockburn Young and the Reverend John N. Russell in Port Glasgow, Scotland. Friends and co-workers alike knew Russell as a quiet and focused, though always kind and helpful person. Trained in classics and biology, Russell's interests drew him to the study of historical and philosophical issues in the biological sciences, particularly morphology and animal behavior. According to Nils Roll-Hansen, Russell was one of the most influential philosophers of biology in the second third of the twentieth century. It was through history and philosophy, rather than his equally important work as a fisheries biologist, Russell argued that developmental and embryological studies deserve a central role in the biological sciences.

In 1916, at the age of twenty-nine, Edward Stuart Russell published his first major work, Form and Function: a Contribution to the History of Animal Morphology. This book has maintained wide readership among scientists and historians since its initial publication, and today is generally recognized as the first modern, sustained study of the history of morphology. In particular, Form and Function incorporates an extensive theoretical analysis of the relationship between embryological studies and comparative morphology in the nineteenth century. Russell employs a history-of-ideas approach in this book, describing the most significant morphologists and their theories. The first chapters of Form and Function discuss early investigators into morphology, such as Hippocrates and Aristotle. The book concludes with a discussion of the opening decade of the twentieth century and the works of Russell’s contemporaries, such as Ernst Mehnert, Hans Driesch, Oscar Hertwig, and Albert Oppel. The broad structure of these chapters, and thus Russell’s overall history, is organized into three main “currents”: a functionalist approach, which includes evolutionary morphologists; a transcendental or idealistic morphology; and finally a focus on experimental embryology or “causal morphology,” to use Russell’s terminology. Consequently the overall framework of Form and Function explains the emerging importance of embryology for an understanding of biological form.

Known by many for his wide-reaching interests and keen thinking, D'Arcy Wentworth Thompson was one of Britain's leading scientific academics in the first few decades of the twentieth century. A prodigious author, Thompson published some 300 papers, books, and articles in the biological sciences, classics, oceanography, and mathematics. He was a famous lecturer and conversationalist-a true "scholar-naturalist," as his daughter wrote in her biography of her father. Of his numerous publications, the acclaimed On Growth and Form (1917, 1945) is generally considered to be his most influential. Many highly respected biologists-like John Tyler Bonner, Joseph Woodger, Sir Peter Medawar, and Stephen Jay Gould-have argued for the importance of On Growth and Form for the history of twentieth century biology. In this work Thompson integrates a causal understanding of biological growth and structure with the mathematics of physical laws. Many developmental biologists have drawn inspiration from reading Thompson's magnum opus, by focusing on this approach to understanding the physical limitations and mathematical processes of developmental growth and morphological form.