The Marine Biological Laboratory (MBL) in Woods Hole, Massachusetts, began in 1888 to offer opportunities for instruction and research in biological topics. For the first few years, this meant that individual investigators had a small lab space upstairs in the one wooden building on campus where students heard their lectures and did their research in a common area downstairs. The lectures for those first years offered an overview of general biology with a focus on zoology, and they were intended for teachers and graduate students interested in acquiring the background for teaching about and/or actually doing laboratory work. As the lab quickly grew, it added sets of lectures that made up courses in zoology, then botany, then physiology, and in 1893 what became the first Embryology Course.
The 1890s were a lively time in embryology, with new techniques and discoveries related to the roles of cells in development, and a growing excitement about the way that experimental embryological manipulations could reveal processes that otherwise remain inside the usually opaque embryo. The MBL recognized the importance of this work and began their course in embryology, which was offered as an elementary course in vertebrate embryology. This was designed for those who had already had the general course, and it had the appeal of offering something more advanced that would bring students back for an additional year. The goal that first year was to allow students to discover the details of development, and to learn the methods for doing the work. The six-week course was directed by Charles Otis Whitman, who was the MBL director, and his student and protégé Frank Rattray Lillie, who became Whitman’s Assistant Director and then his successor as Director both at the MBL and also at the Biology Department at the University of Chicago.
Students were expected to bring their own equipment, including a compound microscope, a dissecting microscope (and it was specified that the Paul Meyer pattern made by Zeiss was the best of the kind), a camera lucida , microtome, and other standard embryological equipment to make up a “complete outfit.” Each student was given a supply of fish eggs and expected to follow the stages of development starting with fertilization. The camera lucida was to facilitate drawing, which was an important part of embryological work until relatively recently. The students all learned the most up-to-date techniques for observing, preserving, embedding, fixing, staining, and then drawing, reconstructing, and modeling embryological processes. The course cost fifty dollars for a number of years and was limited to a dozen students.
For the second year, students were required to have not just a general biology course but also an anatomy course as a pre-requisite. The course continued until 1901, when the lab had grown enough that the course expanded its staff and added zoology as a pre-requisite. The goal of the course was not just to teach the basics of embryology but also to prepare students to take up independent investigations of their own. In addition, the course announcements emphasized the value of studying such a subject at the MBL. Here it was not necessary to rely on preserved developmental stages fixed to slides, but it was possible also to study the living material available during a summer at the seashore. The course continued, with new directors and instructors and with students going on to their own research and sometimes returning as instructors themselves.
Only in 1921 did the course begin to cost seventy-five dollars. Hubert Goodrich became director in 1922 and remained so until 1942. This was an important time for embryology, and the course clearly offered the basics in experimental techniques and introduction to modern theory. It probably served as a very valuable introduction for many young scientists, who encountered living material, interacted with established researchers, and learned how to use equipment and techniques not just to see what others had reported before but also how to ask new questions and prod the embryos to yield answers.
1942 brought a continuation of the basic approach but also a new director. Viktor Hamburger had served as an instructor (view PDF [10.1 MB] of original Hamburger lecture notes) since 1937, shortly after he had taken his experience in Hans Spemann’s lab in Germany to the University of Chicago. There, Lillie undoubtedly lured him to the MBL for the summer and put him to work. Hamburger directed the Embryology Course from 1942 through 1945, when long-time instructor, Donald Costello from the University of North Carolina, took over through 1950. Where Hamburger looked at neuroembryology in his own work and emphasized patterns of development and causes of differentiation, Costello was especially interested in comparative invertebrate embryology. No doubt their approaches worked together well, and the course retained its flavor of lab research drawing on the natural history of the marine offerings.
In the 1950s S. Meryl Rose from the University of Illinois and then Mac V. Edds from Brown University directed the course. This is a period when such notables as John Tyler Bonner of Princeton University, with his enthusiasm for slime molds and problems of morphogenesis, John P. Trinkaus (known as Trink), Clifford Grobstein from the National Cancer Center, Philip Grant from Johns Hopkins University, John W. Saunders from Marquette University, and others brought new perspectives to the traditional course. Throughout this period and dating back to the 1940s, they listed a set of books that students should have, and the list remained surprisingly constant in ways that would be unlikely in this rapidly changing field today.
In 1962 James Ebert at the Carnegie Institution of Washington became director and things changed. The course description shifted for the next year, and the course fee rose to $300—$150 for the series of lectures plus $150 for those who wished to stay for an additional period of individual investigation. The course was becoming less introductory and more designed for would-be researchers such as graduate students and post-doctoral fellows who wanted to learn advanced techniques and to hear about theories and problems in development. The content remained, with additions of Ebert’s specialty work in organogenesis, but Ebert’s five year run as course director made the course at least look more professional in its focus on modern technical topics. This move to professionalism was reinforced by NIH training grants that supported the Embryology and sometimes other courses in the late 1960s and beyond.
In 1971 the course fees rose to $400 + $400, and the announcement made clear that the course was targeted to graduate students and post-doctoral fellows, and to advanced undergraduates seriously interested in pursuing research in this field. Also in 1971, Eric Davidson from Cal Tech served as an instructor. In 1972 he became course director and served in that role through 1974, then again from 1988 through 1996, making him the longest-running course director and giving him the opportunity for the greatest impact on the history of the course. He took great advantage of the opportunity to revise and update the course. In fact, Davidson brought the first major changes in the course which was now “centered around one particular conceptual area of developmental biology.” And the emphasis clearly shifted from instruction and introduction to research to a higher level of expected training and investment in research on the part of the participants.
The focus for 1972 was cytoplasmic localization phenomena, for 1973 the synthesis, storage, and utilization of developmental genetic information during oogenesis in vertebrates and invertebrates, and in 1974 “Sequence Organization in the animal genome and transcription-level gene regulation.” This was a significant shift, and arguably Ebert and Davidson in their different ways had done the most to move the traditional popular course into the professional and molecular era of the late twentieth century.
David Epel from the University of California at San Diego brought an emphasis on cells and cell-cell communication with “Cell Interactions, Cell Membranes, and Cell Surfaces in Development” (1975). Then Tom Humphreys from the University of Hawaii joined Epel in offering “Developmental Regulation of Gene Expression” (1976), “Extracellular Signals in Cell Growth and Differentiation” (1977), and “Localization, Pattern Formation, and Morphogenesis” (1978 and 1979). Rudolf Raff from Indiana University directed “The Control of Events in Early Embryology Development” (1980), “Gene Control and the Events of Early Embryonic Development” (1981), and “Cytoplasmic Localization, Determination, and Gene Control in Development” (1982). These years brought a much expanded list of instructors and lecturers, as the field became more complex and it was important to bring together speakers across the wide range of new ideas and techniques.
Then in 1984, under the direction of William Jeffery from the University of Texas at Austin and Bruce Brandhorst from McGill University, the course reverted to the simpler title of “Embryology: A Modern Course in Developmental Biology.” From 1989 through 1996 the title was “Embryology: Cell Differentiation and Gene Expression in Early Development” and from 1997 to present, it has been “Embryology: Concepts and Techniques in Modern Developmental Biology.”
Discussions by the Education Committee and course directors at various points show that the MBL considered whether to change the name to reflect more current thinking, namely with an emphasis on developmental biology and an emphasis on molecular genetics rather than the traditional embryology. They decided to stick with the traditional course that is now more than one hundred and fifteen years old and instead added other January short courses focused on developmental and molecular techniques. Of course the Embryology course, taught in traditional labs, includes a considerable dose of modern molecular work. And students in recent years admit that they no longer go out to muck about and collect specimens themselves, nor do they have any idea where to look in most cases. Yet there is a tie to tradition, to the observations and collections of the past century, and course instructors will send their students to the MBL Rare Books Room to discover something of the history and context of the work that is now so different from and yet so grounded in the traditions of its predecessors.
View a timeline of the directors and instructors here.
- Marine Biological Laboratory Annual Announcements: MBL-WHOI Library Special Collections.
- Marine Biological Laboratory Annual Reports: available in the MBL The Biological Bulletin Vols. 17 and 21–105 at http://www.archive.org/details/biologicalbullet01mari and beginning with 2004 at http://www.mbl.edu/governance/gov_annual_report.html.
The Marine Biological Laboratory (MBL) in Woods Hole, Massachusetts, began in 1888 to offer opportunities for instruction and research in biological topics. For the first few years, this meant that individual investigators had a small lab space upstairs in the one wooden building on campus where students heard their lectures and did their research in a common area downstairs.Created: 2007-10-24
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.
Just was born in Charleston, South Carolina, on 14 August 1883 to Charles Frazier Just Jr. and Mary Matthew Just. His father died in 1887 and soon after the Just family moved to James Island, off the coast of South Carolina. His early education consisted of attending the small school that his mother founded and directed. Just left James Island at the age of twelve to attend the Colored Normal Industrial Agricultural and Mechanics College at Orangeburg (now South Carolina State College). In 1899 Just graduated with a Licentiate of Instruction, meaning that he was certified to teach in any black school in South Carolina. He was only fifteen years old.
Teaching did not appeal to the teenager so he traveled north, picking up odd jobs along the way until he reached Kimball Union Academy in Meriden, New Hampshire. Just finished a classical course of study in three years, during which time his mother died. The young college graduate was advised by friends and teachers to attend Dartmouth College which he decided to do. At Dartmouth, Just studied biology, history, literature, and the classics. He graduated from Dartmouth in 1907, the only “magna cum laude” in his class, with an AB degree and joined the English faculty at Howard University, Washington, DC in the fall of that same year. He was asked to take over the biology department and teach physiology in 1910, in addition to his English teaching duties. Soon after, Just became the first head of the new Department of Zoology and gave up teaching English courses.
Through a Dartmouth contact, Just communicated with Lillie at the University of Chicago about a post-graduate degree in biology. Lillie directed Just to begin research in 1909 at Woods Hole and to take courses at the University of Chicago. Work at the marine station quickly turned into a research assistantship, working side-by-side with Lillie. All of this was done in addition to maintaining a full-time teaching position at Howard. After obtaining his PhD, Just returned annually to Woods Hole as an independent researcher.
Just focused his interests on marine invertebrate eggs, both in the laboratory and in their natural setting. Because of his tacit knowledge of how marine invertebrates reproduced in oceans and estuaries, Just was able to closely match his laboratory environment to that of the organism’s natural environment. In 1912, Just’s first paper, “The Relation of the First Cleavage Plane to the Entrance Point of the Sperm.” was published in the Biological Bulletin. Just showed that eggs of the marine worm (Nereis) cleave in different planes depending on the sperm’s point of entry. To Just, the egg’s surface was an important and robust factor in the fertilization process. By showing that sperm had an equal probability of entering the egg at any point on the egg’s surface, and that the direction of cleavage depended on the arbitrary point of entry of sperm, and not some predetermined cleavage plane, Just made a dent in preformationist theory. It was also during this time that he married Ethel Highwarden in 1912 and met Jacques Loeb while Loeb was at the Rockefeller Institute for Medical Research. Similar research interests and Loeb’s stand on social equality made the two embryologists fast, but not longstanding friends. In 1915 Just was the first to receive the Spingarn Medal, presented annually to the African-American who performs the greatest service to his or her race. This was followed by the University of Chicago awarding Just his PhD degree in experimental embryology in 1916.
During 1919 and 1920, Just published four papers in the Biological Bulletin, all focusing on his work with the sand dollar Echinarachnius parma. In one set of experiments Just measured the elevation of the egg membrane at sperm contact and the time that it took for the membrane to be penetrated by a sperm. He observed that the sperm was pulled into the egg rather than the commonly held view that it actively bored its way into the egg. Just also documented a “wave of instability” that moved from the sperm’s entry point to the opposite side of the egg. Since then, embryologists have proven that such an instability wave is a wave of cortical granule exocytosis that forms the fertilization envelope. Just also saw that the wave was associated with an immediate blocking of any further sperm penetrability of the egg.
In 1920, while continuing to teach at Howard, Just obtained a ten-year research fellowship from Julius Rosenwald through the National Research Council. He wasted no time in returning to his work at Woods Hole where he continued to study the process of fertilization with results that strengthened Lillie’s work and questioned Loeb’s idea of “superficial cytolysis.” According to Loeb’s studies, egg development could be initiated by exposing eggs to butyric acid. Development was then immediately followed by the release of lysine, Loeb’s cytolytic agent, to break down the egg cortex. Just showed that putting eggs in butyric acid for a short period of time actually slowed cytolysis rather than sped it up. Just was able to prove that the cytolytic effect of the butyric acid was due to overexposure of the eggs to the acid and nothing more. He went even further in dismissing Loeb’s experimental findings on artificial parthenogenesis, attacking Loeb’s method of record-keeping and his apparent failure to maintain experimental conditions to mimic his experimental organisms’ natural environments.
The friendship that Just and Loeb had forged together at Woods Hole quickly vanished. Their disagreements played out for many years, with Loeb providing negative evaluations of Just to the Rockefeller Institute and the Carnegie Foundation. Every time Just tried to obtain grant money, Loeb’s evaluations seemed to rise up and quell any grant award.
In 1929 Just made his first trip to Europe and worked at Anton Dohrn’s Stazione Zoologica in Naples. For six months he experimented with sea urchins (Paracentrotus lividus and Echinus microtuberculatus) to see how these organisms develop and to continue testing Lillie’s “fertilizin” theory of fertilization. Around 1906 Lillie had hypothesized that eggs release a substance that he coined fertilizin. Upon contact with spermatozoa, said Lillie, fertilizin causes sperm to attach to it. Lillie believed that fertilizin molecules served as receptors on the egg’s surface. With receptors for egg and sperm surfaces, the molecule helped to “agglutinate” egg and sperm together.
In 1930 Just was invited to the Kaiser Wilhelm Institute in Berlin where he continued his studies of the ectoplasm with other species, including Amoeba. Just strove to show the importance of ectoplasm in initiation of development. While in Europe he published three articles on the role of ectoplasm: “ Die Rolle des kortikalen Cytoplasmas bei vitalen Erscheinungen” (“The Role of Cortical Cytoplasmin Vital Phenomena”) in Naturwissenschaften (1931) and “On Origin of Mutations” (1932) and “Cortical Cytoplasm and Evolution” (1933), both published in the American Naturalist. All three articles pointed to Just’s view that ectoplasm is necessary for fertilization to occur.
With little hope of ever being able to teach anywhere but a black college, and his continued failure to secure research funds, Just went to Europe in 1938 with the intent of leaving Howard and finishing out his research career on a new continent. In 1939 he published two books: Basic Methods for Experiments on Eggs of Marine Animals and The Biology of the Cell Surface. Both books reflected Just’s holistic view of eggs and embryos: that is, eggs are to be taken seriously in their own right rather than seen simply as tools to manipulate in order to prove a theory. While Just’s experiments may have been simple, he was an intense perfectionist when it came to laboratory procedure. His ability to keep laboratory environments similar to actual marine environments aided in the integrity of his research. It also led to his life-long criticism of experimental embryologists who failed to appreciate his tacit knowledge about inducing marine invertebrate reproduction. To Just, too many embryologists were busy taking eggs out of natural environments and subjecting them to unnatural manipulations while ignoring the importance of the eggs’ environment as an important factor in development. In The Biology of the Cell Surface, Just also continued his attack on the role of genes in development. He remained adamant that cytoplasm was the key to development and not the nucleus. This countered the growing enthusiasm by geneticists who held the idea that the nucleus controlled fertilization and development.
The Nazi invasion of France in 1940 forced Just to return to the US and Howard University, one of the few institutions at the time that would hire a black scientist. His attempt to recareer again in the United States was short-lived however. Just died of pancreatic cancer on 27 October 1941.
- Byrnes, Malcolm W. “Ernest Everett Just.” New Dictionary of Scientific Biography 4: 66– 70.
- Byrnes, Malcolm W., and William R. Eckberg. “Ernest Everett Just (1883-1941): An Early Ecological Developmental Biologist,” Developmental Biology 296 (2006): 1– 11.
- Gilbert, Scott F. “Cellular Politics: Ernest Everett Just, Richard B. Goldschmidt, and the Attempt to Reconcile Embryology and Genetics.” In The American Development of Biology, eds. Ronald Rainger, Keith R. Benson, and Jane Maienschein, 311–42. Philadelphia, PA: University of Pennsylvania Press, 1988.
- Gould, Stephen Jay. “Just in the Middle: A Solution to the Mechanist-Vitalist Controversy.” In The Flamingo’s Smile: Reflections in Natural History, 337-391. New York: W. W. Norton, 1985.
- Just, Ernest E., “The Relation of the First Cleavage Plane at the Entrance Point of the Sperm,” Biological Bulletin 22 (1912): 239–52.
- Lillie, Frank R. “Obituary of Ernest Everett,” Science 95 (1942): 10–11.
- Manning, Kenneth J. Black Apollo of Science: The Life of Ernest Everett Just. New York: Oxford University Press, 1983.
- Pauly, Philip J. Controlling Life: Jacques Loeb and the Engineering Ideal in Biology. New York: Oxford University Press, 1987.
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.Created: 2010-06-16
James David Ebert studied the developmental processes of chicks and of viruses in the US during the twentieth century. He also helped build and grow many research institutions, such as the Department of Embryology in the Carnegie Institution of Washington in Baltimore, Maryland and the Marine Biological Laboratory (MBL) in Woods Hole, Massachusetts. When few biologists studied the biochemistry of embryos, Ebert built programs and courses around the foci of biochemistry and genetics, especially with regards to embryology. He eventually directed the MBL's Embryology Course, and later, the MBL itself.
Ebert was born on 11 December 1921 in the town of Bentleyville, Pennsylvania. He attended public schools while growing up and then graduated from Washington and Jefferson College in Washington, Pennsylvania in 1942. Not long after graduation he joined the United States Navy and eventually became a lieutenant. Ebert was stationed on a destroyer in the Pacific Ocean that was attacked by a kamikaze pilot. The destroyer sank and Ebert spent twenty-four hours in the ocean until being rescued. Afterwards, as a biologist, Ebert befriended and trained several Japanese developmental biologists.
In 1946 Ebert began working towards his PhD in developmental biology under the instruction of Benjamin Willier at the Johns Hopkins University in Baltimore, Maryland. In the same year he married Alma Goodwin, who was a Women Accepted for Volunteer Emergency during the war. Ebert received his PhD in 1950 and immediately became a member of the faculty at the Massachusetts Institute of Technology (MIT) in Cambridge, Massachusetts. After one year at MIT, Ebert moved to Indiana University in Bloomington, Indiana. Ebert became an associate professor of zoology by 1955, and he had started a program of experimental embryology. He studied chick embryos and the processes by which the protein make-up of the embryos changed throughout development.
Six years after receiving his PhD, Ebert became the director of the Carnegie Institution of Washington's Department of Embryology, in Baltimore, Maryland. Prior to Ebert's term as director, the department had had three other directors. The Institution's president, Caryl Haskins, had contemplated closing the department and starting something new. However, with some persuasion from Willier, Haskins spoke with Ebert and decided to give him the opportunity to run the department. Ebert and Haskins agreed that the department needed to focus on the study of genes and their regulation as well as the ways cells influence one another. Haskins said that Ebert's youth and enthusiastic personality made Haskins believe that Ebert would provide a fresh perspective to the department.
Ebert argued that it was his job to recognize and to recruit new talent and then support them in their work. He stressed the use of biochemistry and genetics, which in the 1960s blended together to form molecular biology. During this time, Ebert started to study the relationship between muscle cell differentiation and the propensity to infection in the Rous sarcoma virus.
While still director of the Carnegie embryology department, in 1970 Ebert also became the president and nonresident director of the Marine Biological Laboratory (MBL) at Woods Hole. At the MBL he researched, with Keiko Ozato, the response of murine lymphocytes to mitogens.
In 1977 Ebert ended his term at the Carnegie Department of Embryology, but he remained the director of the MBL. From 1978 until 1987, Ebert lived in Washington, D.C., and he was the president of the whole Carnegie Institution of Washington. As the institution's president, he made the decision to help build a large optical telescope in Chile at Las Campanas Observatory, and he worked towards the creation of a common campus for both Carnegie departments in Washington.
Ebert remained involved with scientific institutions for the rest of his life. When leaving one institution, he found another one to join. He retired from the Carnegie Institution in 1987 and became the president of the Chesapeake Bay Institute at the Johns Hopkins University, where he was a professor of biology for six years. Ebert was elected to many societies including the National Academy of Sciences, the American Philosophical Society, the American Academy of Arts and Sciences, and the Institute of Medicine. He was the vice president of the National Academy of Sciences from 1981 through 1993 and he also chaired its Government-University-Industry Research Roundtable from 1987 through 1993. His colleagues elected him as president of the Society for the Study of Development and Growth, the American Institute of Biological Sciences, and the American Society of Zoologists.
In retirement, Ebert and his wife Alma spent half of each year in Woods Hole and at the MBL. Ebert and Alma died on 22 May 2001 in an automobile accident while en route to Woods Hole.
- DeHaan, Robert L., and James D. Ebert. "Morphogenesis." Annual Review of Physiology 26 (1964): 15–46.
- Ebert, James D. "An analysis of the effects of anti-organ sera on the development, in vitro, of the early chick blastoderm." Journal of Experimental Zoology 115 (1950): 351–77.
- Ebert, James D. "An analysis of the synthesis and distribution of the contractile protein, myosin, in the development of the heart." Proceedings of the National Academy of Sciences 39 (1953): 333–44.
- Ebert, James D. "The effects of chorioallantoic transplants of adult chicken tissues on homologous tissues of the host chick embryo." Proceedings of the National Academy of Sciences 40 (1954): 337–47.
- Ebert, James D. "The formation of muscle and muscle-like elements in the chorioallantoic membrane following inoculation of a mixture of cardiac microsomes and Rous sarcoma virus." Journal of Experimental Zoology 142 (1959): 587–621.
- Ebert, James D., and Ian M. Sussex. Interacting Systems in Development. New York: Holt, Rinehart and Winston, 1970.
- Ebert, James D., and Fred H. Wilt. "Animal Viruses and Embryos." The Quarterly Review of Biology 35 (1960): 261–312.
- Obituaries. "Jim and Alma Ebert." Marine Biological Laboratory. http://www.mbl.edu/news/obit/obit_ebert.html (Accessed December 8, 2007).
- Ozato, Keiko, William H. Adler, and James D. Ebert. "Synergism of bacterial lipopolysaccharides and concanavalin A in the activation of thymic lymphocytes." Cellular Immunology 17 (1975): 532–41.
- Singer, Maxine. "James David Ebert." Proceedings of the American Philosophical Society 148 (2004): 124–27.
James David Ebert studied the developmental processes of chicks and of viruses in the US during the twentieth century. He also helped build and grow many research institutions, such as the Department of Embryology in the Carnegie Institution of Washington in Baltimore, Maryland and the Marine Biological Laboratory (MBL) in Woods Hole, Massachusetts. When few biologists studied the biochemistry of embryos, Ebert built programs and courses around the foci of biochemistry and genetics, especially with regards to embryology.Created: 2008-09-12