Charles Otis Whitman was an extremely curious and driven researcher who was not content to limit himself to one field of expertise. Among the fields of study to which he made significant contributions were: embryology; morphology, or the form of living organisms and the relationships between their structures; natural history; and behavior. Whitman served as director of several programs and institutions, including the Biology Department at the University of Chicago, where he helped establish a new style of biology and influenced the work of many researchers of his generation, as well as future ones. He also served as first director of the Marine Biological Laboratory (MBL) in Woods Hole, MA. Besides his considerable achievements with his own scientific research, Whitman was a tireless mentor who had many students who went on to achieve great success in the field of embryology.
Whitman was born in North Woodstock, Maine, to parents Marcia and Joseph Whitman on 14 December 1842. He grew up on a farm and developed an interest in natural history, particularly that of pigeons, at an early age. Whitman’s family was typical of the rural area where he grew up, and he was educated in the public school system, but despite his family’s lack of money he was highly motivated to receive a college education. Whitman earned money by teaching and tutoring in private schools, and in 1865 he began attending Bowdoin College in Brunswick, Maine. Whitman was enrolled in the accelerated program and finished his degree in 3 years, graduating in 1868 with a BA. After graduation from Bowdoin College, Whitman took a position as Principal of Westford Academy in Massachusetts, where he remained for four years. He then moved to Boston to accept a position as instructor in natural science at English High School. This move was one of great importance, as it was in Boston that he became aware of Harvard University Professor of Zoology Louis Agassiz and enrolled to become one of fifty participants in the first session of the summer marine biology program at the Anderson School of Natural History on Penikese Island in 1873. This experience had a profound impact on Whitman as well as on other of Agassiz’s students. In 1874 Whitman joined the Boston Society of Natural History and, after a second summer at Penikese, he decided to dedicate himself to the full-time study of zoology.
In 1875 Whitman went to study in Europe under Anton Dohrn at the Stazione Zoologica in Naples. After working with Dohrn in Naples, Whitman and his fellow Penikese Island student Charles Sedgwick Minot moved to Leipzig, Germany. There, under the direction of parasitologist Rudolf Leuckart, he learned the modern methods of embryology and microscopy. Whitman received his PhD from the University of Leipzig in 1878. His dissertation was “The Embryology of Clepsine (glossiphonia)”, with an emphasis on the direct role of cleavage in histogenesis, or the differentiation of cells into specialized tissue and organs during growth. This research was instrumental in laying the groundwork for future studies of cell lineage. Whitman found evidence that leech egg development was completely predetermined. This finding supported the regulative theory of embryo development, according to which the whole embryo regulates the development of each cell, in contrast to the mosaic theory in which each cell develops independently, like a mosaic tile. His discoveries while working with the leech were instrumental to future taxonomical and morphological studies.
In 1879 he was offered a postdoctoral fellowship at the Johns Hopkins University but turned it down when he was invited to become Professor of Zoology at the Imperial University of Tokyo. He only spent two years there, but his short tenure was extremely influential. Eight of Whitman’s students there went on to become prominent zoologists, including four who held major chairs, affording him the informal title “father of zoology” in Japan. From November 1881 until May 1882, Whitman returned to the Stazione Zoologica to study the embryology, life history, and classification of the dicyemids which led to the publication of a standard reference work on the parasite in 1883. From 1882 through 1886 Whitman worked as an assistant to Alexander Agassiz at the Museum of Comparative Zoology at Harvard University. During this time Whitman also served as the editor for the Department of Microscopy at the American Naturalist Magazine. After Harvard, Whitman took the job of tutoring amateur zoologist Edward Phelps Allis, Jr., in Milwaukee, Wisconsin. In addition to tutoring Allis, he took on the task of directing the very short lived Allis Lake Laboratory. While there, Whitman oversaw the work of many researchers, including William Morton Wheeler, who went on to become a prominent figure in the study of social insects.
During the summer of 1888 Whitman was invited to direct the newly established Marine Biological Laboratory in Woods Hole, a position he held until 1908. In 1889 Whitman left the Allis Lake facility to take the position of Chair of Zoology at Clark University in Worcester, Massachusetts. In 1892 Whitman moved again to become head of the biology department at the newly founded University of Chicago. There Whitman had several students who went on to make names for themselves in embryology. One of the most prominent was Frank Rattray Lillie, who took over as director at the MBL after Whitman and succeeded Whitman at the University of Chicago, as well. Whitman had many embryologist colleagues at Chicago, including Frank Rattray Lillie, Jacques Loeb, Franklin Paine Mall, Albert Davis Mead, Shosaburo Watase, and William Morton Wheeler. Whitman remained at the University of Chicago until his death on 6 December 1910.
Whitman’s study of sexual dimorphism, the morphological differences between male and female organisms of the same species, was an influence on Oscar Riddle and his endocrinological research. Whitman’s 1898 paper “Animal Behavior” contains many examples of innate, non-learned, behavior. In his later work, he analyzed the relation between innate and learned behavior and the ability of animals to adjust their behavior to new experiences. Whitman saw a similarity of variation in related species, and the trends of evolutionary change in all species from the simplest of organisms to the most advanced. In 1900, when researchers were torn between the theories of mutation and selection, Whitman was a strong proponent of selection.
Whitman published papers and journal articles on every aspect of his work, but is probably best known for his posthumously published three-volume work The Orthogenic Evolution in Pigeons, considered to be the first extensive study in comparative ethology. Whitman was instrumental in the founding of several journals and academic institutions, including the Journal of Morphology, the Biological Bulletin, and the American Morphological Society which, through a merger with the Western Branch of the American Society of Naturalists (known as the Society of American Zoologists in 1901 and 1902), became the American Society of Zoologists in 1902.
Whitman’s work significantly impacted the field of embryology. It greatly influenced the researchers of his generation as well as future generations. Whitman made significant contributions in the fields of embryology, morphology, taxonomy, and ethology. He published numerous books and papers in all of these subjects. Whitman was a mentor to biology students in several institutions around the world. Many of the institutions and publications he founded continue to be at the top of the field of embryology today.
- Carr, Harvey A., and Oscar Riddle, eds. Posthumous Works of Charles Otis Whitman. Vol. I–III. Washington DC: Carnegie Institution of Washington, 1919.
- Davenport, Charles B. “The Personality, Heredity and Work of Charles Otis Whitman.” American Naturalist 51 (1917): 5–30.
- Gilbert, Scott. Developmental Biology, 7th ed. Sunderland, MA: Sinauer Associates Inc., 2003.
- Mayr, Ernst. “Whitman, Charles Otis.” Dictionary of Scientific Biography 13: 313–15.
- Newman, Horatio Hackett. “History of the Department of Zoology in the University of Chicago.” Bios 19 (1948): 215–39.
- Pauly, Philip J. “From Adventism to Biology—the Development of Whitman, Charles Otis.” Perspectives in Biology and Medicine 37 (Spring 1994): 395–408.
Charles Otis Whitman was an extremely curious and driven researcher who was not content to limit himself to one field of expertise. Among the fields of study to which he made significant contributions were: embryology; morphology, or the form of living organisms and the relationships between their structures; natural history; and behavior.Created: 2009-01-21
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
Audio recording from the powerpoint presentation "Jean and Katsuma Dan: A Personal Tribute to Their Legacy in Science and Teaching", by Dr. Shinya Inoue. "Listening to Life Through the Microscope" symposium, held at the Marine Biological Laboratory, Woods Hole, MA, July 14, 2011.
Katsuma Dan reflects on his first meeting with Dr. Victor Heilbrunn at the University of Pennsylvania in December 1930. Recorded at the University of Washington, Friday Harbor group in 1978.Created: 1978
1 black and white video; sound (musical accompaniment only); reformatted digital
By the 1930s, the MBL had become "the" place to go during the summer for biological research and training. Luminaries such as Frank Lillie, Edmund Beecher Wilson, Edwin Grant Conklin, and Thomas Hunt Morgan took their students, packed up their families and research labs, and headed to the MBL. They worked in labs, ate together in the Mess, and they often lived in the limited on-campus housing. Life at the MBL was a life where fun, family, and science intertwined. This film, taken in 1935 by B. R. Coonfield of Brooklyn College, captures snippets of life at the MBL.Created: 1935