Students and faculty in the 2001 Embryology Course at the Marine Biological Laboratory in Woods Hole, MACreated: 2001
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
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.
In 1892 Lillie followed Whitman to the University of Chicago zoology department where Whitman had accepted a chair appointment. In 1894 Lillie graduated with a PhD in zoology. His dissertation was a descriptive study of cell lineage in freshwater mussels. From 1894 through 1899 Lillie worked as an instructor at the University of Michigan. In 1895 he married Frances Crane, sister of Chicago businessman Charles R. Crane. His brother-in-law would soon play a large role in introducing Lillie to the social elite in Chicago and helping to expand the MBL campus. During his marriage Lillie and his wife had five daughters, one son, and three adopted sons. Lillie briefly taught biology in upstate New York at Vassar College before returning to the University of Chicago as an assistant professor of embryology. In 1902 he was made an associate professor followed by a full professor in 1906. In 1908 Lillie published his classic book on chicken embryology, Development of the Chick: An Introduction to Embryology. Along with writing the text, Lillie prepared a large series of serial sections of the chick embryo at various stages to serve as illustrations. With revisions, the text and laboratory manual continue to be used to the present time, serving as one of the best accounts available on bird development.
In 1910 Lillie was made chair of the Department of Zoology. During this time he united the embryology program with the rest of the zoology department. With the combining of the departments, the budgets were also combined and Lillie was able to use his influence to obtain more money for embryological research. In 1931 Lillie was appointed dean of the Division of Biological Sciences at Chicago. After many years of distinguished teaching and research he was made the Andrew MacLeish Distinguished Service Professor of Embryology and the dean of biological sciences. In 1935 he was given emeritus professor status.
Although rarely written about, Lillie was a member of Chicago’s Eugenics Education Society, a committee member of the Second International Eugenics Congress, and served on the advisory council for the Eugenics Committee of the United States. In the early 1920s Lillie envisioned an Institute of Genetic Biology that would gather data to examine population problems, public health, and social control, but this never came to fruition.
Lillie is probably best known for his leadership at the Marine Biological Laboratory at Woods Hole. He organized the MBL’s first course in embryology in 1893 and became course director the following year. At that time the MBL consisted of one small building, a few skiffs, and a dock. In 1902 funding for the laboratory was so great that the corporation and board of the MBL considered transferring the laboratory to the Carnegie Institution of Washington to make the MBL Carnegie’s permanent marine research laboratory. Lillie and Whitman opposed the transfer and convinced the board to reverse its offer to Carnegie. To this day the MBL remains a laboratory relatively free from layers of outside control.
From 1900 to 1942 Lillie worked tirelessly to improve the laboratories and accommodations for the myriad of scientists who descended on Woods Hole during the summer months. Lillie called upon his brother-in-law to help finance the expansion of the MBL and Crane served as president of the corporation from 1904 through 1924. The Crane laboratory at MBL was named after the man who financed its building. Lillie was instrumental in making the marine laboratory into one of the leading research laboratories in the world. Not only did he serve as president of the corporation from 1925 to 1942 but he also served as Managing Editor of the MBL’s scientific publication The Biological Bulletin for twenty-five years.
Lillie was an outstanding administrator and teacher but the depth of his research in embryology and development is also remarkable. His early research primarily dealt with egg cleavage and early development in invertebrates. Although his early cell lineage work was mainly descriptive and comparative, it helped lay the foundation for experimental studies by Wilhelm Roux in 1888 and Hans Driesch in 1891. Lillie heavily influenced his former student and noted embryologist Ernest E. Just to continue working with Nereis to show the relationship of egg cleavage planes to the entry point of sperm.
In 1903 and 1904 Lillie published several papers on his studies of the chick embryo. Included in the papers was discussion about the formation of the amnion and his experiments with cauterizing parts of the embryo to see how further development of parts of the embryo were affected. Lillie had always shown interest in the chick embryo. He was convinced that chick embryos were the best choice for almost any type of experimental work of embryological problems.
From 1910 to 1921 Lillie’s research centered on fertilization in the annelid Nereis limbata and sea urchins Arbacia punctulata and Strongylocentrotus franciscanus and S. purpuratus. Lillie proposed that there were specific substances (fertilizin and antifertilizin) secreted by egg and sperm. Part of Lillie’s “fertilizin theory” likened the interaction between gametes to that of the lock-and-key fashion of antibodies and antigens. This was notable in that Lillie applied the then current immunological concepts and terminology to that of fertilization.
Lillie’s investigation of the factors influencing the development of freemartins (sterile genetic female calves born as a twin to fertile male calves) helped Lillie answer the question of how sexually indifferent embryos at the beginning of development later turn into males or females. Beginning in 1914 Lillie worked with stockmen in the Swift and Company stockyard to obtain fifty-five pairs of in utero fetal twins from freshly slaughtered pregnant cows. In 1917 Lillie published his study in the Journal of Experimental Zoology with the finding that freemartin bovine twins are non-identical and that they share the same placenta, allowing for blood to be freely exchanged between the twin fetuses. A male’s testes form early in development and “masculinizing” substances (hormones) are released and circulate through the fused umbilical arteries. Lillie concluded that the freemartin was a genetic female calf that had certain male sexual characteristics due to the action of a fetal male sex hormone. This work led to the concept that once a gene directs a gonad to differentiate into a testes or an ovary, accessory reproductive structures in genetic males develop in the male direction due to the presence of male hormones. Genetic females develop rudimentary reproductive structures because they are not inhibited from developing due to a lack of male sex hormones. Lillie’s research with freemartins introduced the notion of the nature and action of sex hormones to embryologists when little was known about hormones. Soon, others at the University of Chicago attempted to produce freemartins in birds ( Benjamin Willier) and mammals ( Carl R. Moore). Castration experiments in fetal mammals in utero and research in the isolation and purification of sex hormones was undertaken in other laboratories at the University of Chicago. The subsequent work stemming from Lillie’s freemartin investigation helped form the field of reproductive biology.
Even after retiring from the University of Chicago in 1935, Lillie continued with his sex hormone studies by investigating the physiology and development of bird feathers. He used the Brown Leghorn fowl, a bird that displays a notable sexual dimorphism in feather color and patterns. He collaborated with Mary Juhn and His Wang to discover that embryonic feather papillae all start out with the same background color. Further feather coloration and patterns develop in an orderly fashion in response to both female sex hormones and thyroxin. Part of his research involved using castrated males into which injections of estrogens and thyroxin were given to induce “female” feather colorings in the birds’ regenerating feathers.
From 1935 to 1939 Lillie was president of the National Academy of Sciences and in 1935 to 1936 was chairman of the National Research Council. To date he is the only person ever to have held both leadership positions in the two organizations at the same time. Lillie was also appointed chairman of the National Academy of Sciences Oceanographic Committee to study the financing and construction of an Institute of Oceanography. In 1930 he helped secure a three million dollar grant from the Rockefeller Foundation to help locate and build the Oceanographic Institute next door to the MBL. Lillie served as president of the Woods Hole Oceanographic Institute from 1930 to 1939.
Lillie died of a stroke on 5 November 1947 in Billings Hospital at the University of Chicago, the campus at which his professional life had so intimately been connected. Thus ended the career of one of the world’s foremost embryologists and science administrators—a career that science historian Philip J. Pauly identified as having helped make biology the first science in which Americans became internationally recognized.
- Lillie, Frank R. The Problems of Fertilization. Chicago: University of Chicago Press, 1919.
- Manning, Kenneth J. Black Apollo of Science: The Life of Ernest Everett Just. New York, Oxford University Press, 1983.
- Mitman, Gregg. The State of Nature: Ecology, Community, and American Social Thought 1900–1950. Chicago: University of Chicago Press, 1992.
- Moore, Carl R. “Frank Rattray Lillie: 1870-1947.” Science 107 (1948): 33–35.
- Pauly, Philip J. Controlling Life: Jacques Loeb and the Engineering Ideal in Biology. New York: Oxford University Press, 1987.
- Watterson, Ray L. “The Striking Influence of the Leadership, Research, and Teaching of Frank R. Lillie (1870-1947).” American Zoologist 19 (1979): 1275–87.
- Willier, Benjamin. H., Ross G. Harrison, Henry. B. Bigelow, and Edwin. G. Conklin. “Addresses at the Lillie Memorial Meeting Woods Hole.” The Biological Bulletin 95 (1948): 151–62.
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.Created: 2009-07-22
Jacques Loeb experimented on embryos in Europe and the United States at the end of the nineteenth and beginning of the twentieth centuries. Among the first to study embryos through experimentation, Loeb helped found the new field of experimental embryology. Notably, Loeb showed scientists how to induce artificial parthenogenesis, thus refuting the idea that spermatozoa alone were necessary to develop eggs into embryos and confirming the idea that the chemical constitution of embryos’ environment affected their development. Furthermore, Loeb’s work showed that scientists could manipulate materials in a laboratory to create, as he called the process, the beginning stages of life.
Jacques Loeb was born in the Prussian town of Mayen to Barbara and Benedict Loeb in 1859. Named “Isaak,” he changed his name to “Jacques” just prior to entering the University of Strassburg in 1880. At Strassburg Loeb studied with the physiologist Friedrich Goltz and there he earned his MD in 1884.
Until 1891 Loeb taught and researched at various institutions, including the Naples Zoological Station in the winters of 1889 and 1890. In 1890 he met and married Anne Leonard, an American philologist. Moving to the United States, Loeb taught at Bryn Mawr College for a year prior to accepting an assistant professorship with the University of Chicago in 1892. While at Bryn Mawr Loeb met and initially disliked the young Thomas Hunt Morgan. At Chicago Loeb struggled to get along with Charles Otis Whitman, and to tolerate what Loeb termed the “romantic evolutionism” espoused most prominently by Chicago’s John Dewey.
After a decade at Chicago, Loeb moved to the University of California at Berkeley for eight years. In California he experimented at Stanford University’s Hopkins Marine Station in Pacific Grove. He then returned east to New York’s Rockefeller Institute for Medical Research in 1910, working there until he died in 1924. During his career, he conducted many of his experiments at Bryn Mawr, Chicago, Rockefeller, and at the Marine Biological Laboratory in Woods Hole, Massachusetts.
Loeb’s early work at Strassburg was on brain physiology. Having judged the field’s dominant theory misguided, he decided to instead study traditional issues in biology such as embryology. He used experiments to answer questions about development and embryology rather than observation alone, which was the common method of biological inquiry. Loeb looked to the scientific practices of physiologists Goltz, Eduard Pflüger, and the eminent plant physiologist Julius Sachs for methodological inspiration.
Loeb worked with Sachs extensively while the former worked at the University of Würzburg for two years starting in 1886. Loeb then developed a correspondence and friendship with the Austrian physicist and philosopher Ernst Mach, whose writings provided the theoretical foundation for Loeb’s work. For Loeb, any explanation of phenomena could come only from manipulating the physical structures of things and their chemical makeup. That conception of scientific knowledge, based on physicochemical manipulations, paralleled Wilhelm Roux’s mechanistic conception of science encapsulated in Entwicklungsmechanik.
Like Mach, Loeb thought science was not a mere description of nature but was instead a tool for humans to interact with nature. To him, a biologist was like an engineer and organisms were biologists’ material. Loeb earned his reputation as a biologist-engineer in 1899 when he published experimental results showing artificial parthenogenesis in sea urchins, in “ On the Nature of the Process of Fertilization and the Artificial Production of Normal Larvae (Plutei) from the Unfertilized Eggs of the Sea Urchin”.
Artificial parthenogenesis is the human manipulation of egg cells causing embryonic development without spermatozoa. Loeb manipulated unfertilized sea urchin eggs with inorganic solutions of salt water. The result was eggs developing into larvae, or early stage embryos, which he later reproduced using frog eggs. Loeb elaborated on his results in 1913’s Artificial Parthenogenesis and Fertilization.
Loeb’s 1899 results stirred the popular presses, and many viewed him as a creator of life, perhaps with the ability to engineer new types of organisms. Loeb was never able to create new forms of life, as he intended, but he inspired scientists such as John Howard Northrop, John Broadus Watson, Hermann Joseph Müller, Burrhus Frederic (B. F.) Skinner, and Gregory Pincus. Loeb also contributed to the study of animal tropisms (environment-caused orientation).
Loeb helped transform biology into a largely experimental science. His 1912 The Mechanistic Conception of Life established his reputation as a researcher who treated organisms as machines. In that work, he stated that biologists explain organic phenomena only when they could control those phenomena. Loeb later believed that biologists explain phenomena by detailing the mechanisms, the step-by-step processes, by which a component of an organism achieves its function in physical and chemical terms. In The Organism as a Whole (1916), Loeb discussed how a mechanist could investigate organisms considered as wholes. His commitment to physicochemical explanations led him to study protein chemistry for the last few years of his life. Loeb died in 1924.
- Loeb, Jacques. Artificial Parthenogenesis and Fertilization. Chicago: University of Chicago Press, 1913.
- Loeb, Jacques. “Mechanistic Science and Metaphysical Romance.” Yale Review 4 (1915): 766–85.
- Loeb, Jacques. “On the Nature of the Process of Fertilization and the Artificial Production of Normal Larvae (Plutei) from the Unfertilized Eggs of Sea Urchins.” The American Journal of Physiology 3 (1899): 135–38.
- Loeb, Jacques. The Organism as a Whole: From a Physicochemical Viewpoint. New York: G.P. Putnam’s Sons, 1916.
- Loeb, Jacques. The Mechanistic Conception of Life. Ed. Donald Fleming. Cambridge, Mass.: Harvard University Press, 1964.
- Pauly, Philip. Controlling Life: Jacques Loeb and the Engineering Ideal in Biology. New York: Oxford University Press, 1987.
Jacques Loeb experimented on embryos in Europe and the United States at the end of the nineteenth and beginning of the twentieth centuries. Among the first to study embryos through experimentation, Loeb helped found the new field of experimental embryology. Notably, Loeb showed scientists how to create artificial parthenogenesis, thus refuting the idea that spermatozoa alone were necessary to develop eggs into embryos and confirming the idea that the chemical constitution of embryos environment affected their development.Created: 2009-06-10
Students and faculty in the 1993 Embryology Course at the Marine Biological Laboratory in Woods Hole, MACreated: 1993