C. Ladd Prosser

C. Ladd Prosser
May 12, 1907 - February 3, 2002

Clifford Ladd Prosser, Professor Emeritus of Physiology and Neuroscience at the University of Illinois at Urbana–Champaign, passed away on Sunday, February 3, 2002, three months before his 95th birthday. Notwithstanding his eminence and scientific stature, he was fondly known as “Ladd” by his friends and colleagues all around the world. Ladd was the personification of a true scientist and an embodiment of a comparative physiologist. He devoted all of his adult professional life, from the early 1930s to the very end, to the pursuit of his love – comparative animal physiology.

Early Life. Ladd was born in the year 1907 in the village of Avon, in Western New York. His ancestors were farmers. His father, Clifford James Prosser, owned a General Store in Avon. As a young boy, Ladd’s Sunday afternoon outings with his father, along the woody paths of the Genesee River and its surrounding lakes, gave him an early start in the appreciation of Nature and the diversity of her animal and plant life.

Higher Education. At the age of eighteen, Ladd attended the University of Rochester, 20 miles North of Avon, where he received his degree in Zoology in 1929. His love of Nature spilled over into an enthusiasm for experimental studies in biology, so much so that he applied and was admitted to the Johns Hopkins University graduate program receiving a Ph.D. in Biology (1932). Part of his doctoral studies, carried out with the well-known protozoologist S.O. Mast, involved motor behavior of amoebae. Ladd also began, during his graduate studies, an independent study on the development of the nervous system and behavior in the earthworm. These early studies laid the foundation for his lifetime work in comparative animal physiology.

Postdoctoral Studies at Harvard, Cambridge & Oxford. A Parker Fellowship provided Ladd with the opportunity to carry out postdoctoral research at Harvard (1933) and England (1934). At Harvard Medical School, he worked with the well-known auditory neurophysiologist Hallowell Davis where he sharpened his skills in electrophysiology and discovered the caudal photoreceptor of crayfish. He also associated with such greats in American physiology as Walter Cannon and Alexander Forbes. Ladd was ambitious and went on to spend the next year at Cambridge and Oxford in England where he worked with such towering figures of Physiology as Edgar Adrian and John Eccles.

Research & Teaching: Clark University & Marine Biological Lab. Ladd’s first research and teaching position was at Clark University in Worcester, Massachusetts in 1934, the same year that he married the love of his life, Hazel Blanchard. While at Clark, he spent the summers working at the Marine Biological Laboratory (MBL) in Woods Hole, Massachusetts, doing research in invertebrate neurophysiology, in part focusing on the role of acetylcholine in marine organisms. During this period he met Kenneth “Kacy” Cole and Howard “Bim” Curtis, famous American biophysicists, with whom he developed a lifelong friendship; at MBL he also met and befriended such famous physiologists as Alan Hodgkin, Albert Szent-Gyorgi and Steve Kuffler. He remained a faithful visitor to MBL for many summers to come and became one of its Trustees (1950).

Moving to Urbana & the War Years in Chicago. In 1939 Ladd, in search of a larger and more research-oriented institution, was offered a faculty position at the University of Illinois at Urbana to teach Zoology to agriculture students. After the birth of Ellen, their first daughter, in 1939, the family took the train for Urbana where they settled for decades to come. The family grew larger with the birth of their second daughter Nancy in 1942 and their son Loring in 1945. After the attack on Pearl Harbor in 1941, Ladd was recruited by Kacy Cole to join a biomedical team in Chicago as part of the Manhattan Project to study the effects of high-level radiation on humans and other animals. These findings later formed part of the basis for post-war radiation safety standards.

Physiology Research, Training of Graduate & Postdoctoral Fellows at Illinois. After the war, Ladd returned to the University of Illinois, as a full professor and in 1949 helped form the Physiology Department, together with Bob Johnson, Fredrick “Teck” Steggerda, A. B. “Jack” Taylor and others. In his research lab in the Natural History Building, and later in Burrill Hall, Ladd began a 50-year career of research that included the training of about 45 doctoral students and numerous postdoctoral fellows from America, Europe, Asia and Australia. Many of his graduate trainees and associates, scientists such as Lloyd Barr, Geoff Burnstock, Andy Cossins, Brian Curtis, Asit Das, Mike Friedlander, Jeff Hazel, Madhu Kanungo, M. Kobayashi, Nick Kotchabhakdi, Jane Liu, Richard Meiss, Toshio Nagai, T. Tamaik, Bruce Sidell, Nick Sperelakis, Joe Szurszewski, Victor Wilson, and Jackie Wood, went on to become heads, chairs or directors of physiology, biochemistry, biophysics or neuroscience departments or programs in various states and countries.

Services to Physiology, Biophysics and Neuroscience at Illinois. From 1960 to 1969, Ladd was Head of the Physiology Department at Illinois. In 1962 he brought Biophysics into the Department, renaming it Physiology and Biophysics, an excellent pairing which lasted for more than three decades. At its height, this Department had nearly 40 faculty and 120 students, and was one of the top 10 in the nation, in terms of graduate teaching and productivity. During his headship Ladd recruited several new physiologists and biophysicists into the Department of whom Bill Sleator, Jim Heath, and Dennis Buetow, in turn, became Head of the Department. Ladd was instrumental in the building of Burrill Hall in the late 1950s. Upon its dedication, thanks to his leadership, the building served as the site of a joint meetings of the American Physiological Society and the Society of General Physiologists (Fall, 1960). Ladd’s services to the University of Illinois were not limited to Physiology. In the 1970s, together with Ed Banks and Bill Greenough, he helped establish the Neural and Behavioral Biology Graduate Program, later renamed the Neuroscience Program, an active interdisciplinary program currently comprising nearly 60 faculty and numerous graduate students.

National Scientific Services. In addition to his services to the University, Ladd was, in the words of an eminent national colleague, “A statesman of science”, ever ready to serve and representing the needs of science and the scientists. Thus he served as the President of the Society of General Physiologists (1958), the American Society of Zoologists (1961), and the American Physiological Society (1969). He also served on the editorial boards and as editor of many journals including Comparative Physiology and Biochemistry, American Journal of Physiology and Physiological Zoology. He also served on numerous governmental and national committees, including NASA, the National Academy of Sciences, and the study sections of the National Science Foundation, National Institutes of Health, as well as a post-war Committee on the Biological effects of Radiation in the Bikini atoll and a National Academy committee on the ecology of Panama Canal.

Prosser’s Books and Retirement Years. Ladd Prosser’s magnum opus was his massive textbook of “Comparative Animal Physiology” (Wiley, New York), which underwent four editions from 1950 to 1991 and was translated into three languages, including Russian. This work made his name a familiar one in the physiological circles all around the world. Through its pages, generations of students around the world were introduced to comparative physiology. Although he retired in 1975, he continued active research until the mid-1990s. We fondly remember Ladd carrying his bucket full of small fish in the Burrill Hall hallways, in pursuit of his many experiments on fish adaptation. His last Ph.D. trainee, William “Bill” Seddon, graduated in 1994. The result of Bill’s thesis was the subject of Ladd’s last collaborative research paper, published in “Comparative Physiology & Biochemistry” (1999). Interestingly, his first research paper had also been published, nearly seven decades earlier, in a similar comparative publication, “Journal of Cellular & Comparative Physiology” (1932).

Continuing Work in Spite of Failing Health. In 1997 Ladd suffered a serious hip fracture; this together with the debilitating stroke of his beloved wife Hazel in the same year, put an end to his life in the lab. Nevertheless he continued working at home on his two book projects, “Scientific Autobiography and Personal Memoir” (Stipes Publishing, Champaign, 2001) and a more massive undertaking, “A History of Nerve, Muscle and Synapse Physiology” (Essie Meisami, Ed., Stipes Publishing, due summer 2002). He also diligently read issues of “Science” and “Nature” as well as books on biography and American history.

Prosser’s Scientific Legacy. During his 70-year long scientific career, Ladd Prosser produced 7 books, over 50 reviews and monographs and nearly 150 research papers, exclusive of abstracts and reports. His most innovative book was “Adaptational Biology: From Molecules to Organisms” (Wiley, New York, 1986) in which he tried to develop a unified theory of evolutionary adaptation by studying various levels of organismic complexity. Ladd’s early research concerned the properties of the nervous system of simple animals and invertebrates. One of his early and seminal discoveries was the demonstration of spontaneous activity in the isolated nervous system of invertebrates, a thesis contrary to the prevailing views of the behaviorists who at the time thought that all behavior is stimulus-dependent, and which led ultimately to modern-day views of central pattern generators in the nervous system.

In the 1950s Ladd’s research emphasis changed from nerve to muscle, particularly smooth muscle, its functional diversity and adaptation. In his words, “I decided that animal speed was due more to muscle than nervous system”. This focus remained central through the rest of his career. Among his theoretical insights was the concept of a relationship between muscle size (diameter) and its speed of contraction. He also helped develop some of the concepts of slow waves in intestinal smooth muscles and the role of the interstitial cells of Cajal. Another legacy of Ladd was the many in vivo and in vitro studies on biochemical and physiological adaptation of fish and other marine invertebrates to changes in environmental temperature.

Awards & Honors. For his lifetime of research and dedication to comparative physiology, Dr. Prosser received numerous local, national and international awards and honors, including a Guggenheim Fellowship, a Fellow of the American Academy of Arts and Sciences, 50th Anniversary Award of the American Society of General Physiologists. He considered, however, election to the National Academy of Sciences (1974), the highest honor bestowed upon him. Ladd Prosser was among the few giants of comparative physiology in the second half of the twentieth century. He will be dearly missed but to borrow a quote from General MacArthur, “Old scientists never die, they just fade away gradually.”

The Prosser Family. Ladd Prosser leaves two daughters, a son, six grandchildren and one great-grandchild. His older daughter, “Jane” Ellen Prosser Armstrong, lives in Ladd’s favorite summer place, Woods Hole, Massachusetts. Ladd’s second daughter, Nancy Ladd Prosser Meinertzhagen, makes her home in Halifax, Nova Scotia, Canada. Ladd’s son, Loring Blanchard Prosser lives in Indianapolis, Indiana. Ladd’s wife, Hazel Blanchard Prosser, died in 1999, just before her 92nd birthday, from complications after an earlier stroke.

Prepared by Essie Meisami, with input from Phil Best, Dennis Buetow, Howard Ducoff and Bill Greenough, Department of Molecular & Integrative Physiology, University of Illinois, Urbana-Champaign and Ian Meinertzhagen, Neuroscience Institute, Dalhousie University, Halifax.