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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.
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