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

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.

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.

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.

Historically the exact age of human embryo specimens has long perplexed embryologists. With the menstrual history of the mother often unknown or not exact, and the premenstrual and postmenstrual phases varying considerably among women, age sometimes came down to a best guess based on the weight and size of the embryo. Wilhelm His was one of the first to write comparative descriptions of human embryos in the late 1800s. Soon afterward, Franklin P. Mall, the first director of the Carnegie Institution of Washington's (CIW) Department of Embryology, expanded upon His' work. Mall's first efforts were to place embryos into stages based on menstrual ages and body length. This method ran into problems however when it became apparent that obtaining menstrual ages was often impossible or simply too inaccurate even if the information could be obtained from the women who carried the embryos. Mall decided instead to look for patterns among embryos to come up with some type of staging system whereby embryo age could be more accurately determined.

Jacques Loeb is best known for his embryological work investigating parthenogenesis in invertebrates. Artificial Parthenogenesis and Fertilization is a revised and English-translated work from his earlier book, Die chemische Entwicklungserregung des tierischen Eies (1900). Artificial Parthenogenesis describes Loeb's many and varied methodical experiments to initiate egg development without fertilization by sperm. As is true with much of science, some of Loeb's experiments were successful and many were not. Artificial Parthenogenesis presents a sense of what early twentieth century embryology looked like: experimenters' overarching desire for manipulation and control, coupled with their use of chemicals and macromolecules as agents of change. The book also illuminates the historical role of the sea urchin in the study of embryological development.

Felix Anton Dohrn is best remembered as the founder of the Stazione Zoologica di Napoli, the world' s first permanent laboratory devoted to the study of marine organisms. Dohrn was born on 29 December 1840 in Stettin, Pomerania (now Poland), to a wealthy merchant family. Dohrn's paternal grandfather, Heinrich, trained as a surgeon and then established a sugar refinery, while Dohrn's father, Carl August Dohrn, who inherited the family business, became interested in natural history through Alexander von Humboldt, a family friend. Once settled in his career, Anton Dohrn's own research never strayed far from the origin of vertebrates. He promoted the theory that vertebrates closely resemble and are derived from annelid-like ancestors and he spent years studying the early embryogenesis of lower vertebrates in attempts to prove this.

'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 1931 embryologist and historian Joseph Needham published a well-received three-volume treatise titled Chemical Embryology. The first four chapters from this work were delivered as lectures on Speculation, Observation, and Experiment, as Illustrated by the History of Embryology at the University of London. The same lectures were later released as a book published in 1934 titled A History of Embryology. This monograph represents one of the first general accounts of the history of embryology and presents embryology as a history of intertwined ideas, a style of historical writing advanced by noted biology historian Jane Oppenheimer. A revised 1959 edition of the text published by Abelard and Schuman, New York, examines the history of embryology from antiquities to the mid-nineteenth century. Arthur Hughes, lecturer in anatomy at Cambridge University, is credited by Needham as providing technical assistance with the new version.