KEITH Roberts Porter died on May 2, 1997, just over
a month short of his 85th birthday. He had the
perspicacity, good fortune, and patience to take
advantage of the fast moving frontier of analytical biology
after the Second World War to provide many of the techniques and experimental approaches that established the
new field of biomedical research now known as cell biology. He was renowned for taking the first electron micrograph of an intact cell, but his contributions went far beyond that seminal instance. They ranged from technical
developments, such as the roller flask for cell culture and
the Porter-Blum ultramicrotome, to experimental and observational achievements, such as studies on the synthesis
and assembly of collagen, on the role of coated vesicles in
endocytosis, on lipid digestion in the intestine, and on the
universality of the 9 + 2 axoneme in cilia. The initial ultrastructure descriptions of the endoplasmic reticulum and
the sarcoplasmic reticulum, identification of the role of
T-tubules in excitation-contraction coupling in muscle and
the role of the cytoskeleton in cell transformation and
shape change, were his, as were many other contributions,
described in some detail elsewhere (Peachey and Brinkley,
1983
In addition to his specific scientific contributions, Keith
Porter also made more important philosophical contributions to the field that he helped to shape. These principles
include the understanding that the cell is not a "bag of enzymes"; that organelle structure is consistent from cell to
cell throughout a wide range of protists, animals, and plants;
that this means that cell structure and function have a
macromolecular basis; that self-assembly is a critical morphogenetic principle; and, most presciently, that the cell is
structurally integrated down to molecular resolutions by
an intricate network of cytoplasmic proteins, this integration having consequences for signal transduction and function. These contributions sometimes involved experienced
collaborators and, in later years, young disciples with whom
Porter developed particular empathy.
Keith Porter was a boy from Yarmouth, Nova Scotia,
who grew up far from the centers of science in the United
States and Europe. He went to college at Acadia University in Nova Scotia and then did graduate work at Harvard
University. He began his postdoctoral research career at
Princeton University and then moved to The Rockefeller
Institute for Medical Research (now The Rockefeller University) in the late 1930s, where he joined the laboratory of
James B. Murphy. In 1938, he married Elizabeth Lingley,
who nurtured him for over a half century. Their family life
was disrupted tragically by tuberculosis during the war years,
when their young son died. Porter became an American
citizen in 1947.
By 1946, Murphy's laboratory included Albert Claude,
G.C. Hogeboom, W.C. Schneider, George Palade, who
had arrived from Bucharest, and Keith Porter. It was, as
Palade described it later, the cradle of cell biology, where
cell fractionation and cell fine structure were born and
nurtured, and it remained so after Claude returned to Belgium in 1949. In an unusual move, when Murphy retired in 1950, Herbert Gasser, the Director of The Rockefeller Institute, placed Porter in charge of a newly created Laboratory of Cytology and recommended his promotion to Associate Member (equivalent to Associate Professor).
Palade also joined this new endeavor, and he became Associate Member in 1953. If Murphy's group nurtured the
newborn studies of the cell, the laboratory of Porter and Palade from 1953-1961 raised the field from infancy to
maturity.
During these years Keith Porter inspired, created, established, and led the institutions that we know today in the
field of cell biology, namely this journal (founded as the Journal of Biophysical and Biochemical Cytology) and the
American Society for Cell Biology. It is no coincidence
that Porter was both the chair of the committee that
founded the American Society for Cell Biology and the
first editor of the journal, that the Journal of Cell Biology
is published by The Rockefeller University Press, that the
American Society for Cell Biology has played an important role in the history of the journal, and that the present
editor of the journal is Keith Porter's scientific grandson.
Porter's account (with H. Stanley Bennett) of the founding
of the journal appeared in the December 1981 supplement
Discovery in Cell Biology (Journal of Cell Biology, Vol.
91, No. 3, Pt. 2).
In 1961, Porter moved to Harvard University to become
Professor of Biology and subsequently Chairman of the
department. In 1968, he moved to the University of Colorado at Boulder as Chairman of the new Department of
Molecular, Cellular, and Developmental Biology. There
he developed facets of scanning microscopy and he established a High Voltage Electron Microscope Facility that continues to be a national resource. From each instrument he
employed, he drew a set of compelling images that he disseminated in talks, publications, and atlases. He retired
from the University of Colorado in 1983, at which time
the building that housed his laboratory was named in his
honor. He did not rest on his laurels but moved again to
become Wilson Elkins Distinguished Professor in the Department of Biological Sciences at the University of Maryland, Baltimore County. At age 75, he finally moved once
again to become Distinguished Research Professor of Biology at the University of Pennsylvania, returning to collaborate with Lee Peachey, who was his first graduate student.
His interest in students was whetted at The Rockefeller University, where he and Palade established a premier
course in comprehensive cell biology. At Rockefeller, Harvard, Colorado, and Maryland, he trained a coterie of
postdoctoral associates and graduate students who spread
the gospel of cell biology and fine structure throughout the
world. While he had a pungent wit, a sharp sense of humor, and a critical eye, for which his lectures were known,
many a cell biologist throughout the country owes his/her
position to Keith Porter's kind recommendations and sympathetic encouragement. Even now, he continues to support the field through the Keith Porter Endowment to
which his estate is the major contributor.
Porter was celebrated and honored in many ways, with
festschrifts, with a dedicated volume of the Journal of Cell
Biology (Palade, 1977 In 1956, as preface to the Proceedings of a Conference
on Tissue Fine Structure that appeared as a supplement to
Volume 2 of this journal, Porter wrote: "It must be evident
by now to even the most confirmed skeptic that electron
microscopy is destined to have a profound influence on
the future development of biology and related sciences.
Investigations carried on with the microscope . . . are currently revealing unexpectedly numerous and complex details of structure in the cells of plants and animals. Such studies usher in a period of microscopic discovery that will
surely match if not surpass in importance the 40 or 50 years of activity that followed the introduction of improved
optical microscopes, and techniques of fixation, sectioning,
and staining about a century ago.
For those of us who are fortunate to be part of this new
development, these are days of great interest and opportunity."
The 40 years have passed and now the father of the field
has died. We would do well to leave our successors as rich
a legacy of opportunity in cell biology as he has left us.
Peter Satir
Department of Anatomy and Structural Biology, Albert
Einstein College of Medicine, Bronx, New York 10461-1602
; Moberg, 1996
). Absent from this list are his early pioneering work establishing the androgenetic haploid in
frogs, an exercise in nuclear transplantation with consequences for the recent cloning of mammals, and his later adventures with pigment migration in fish chromatophores.
Photo courtesy of Dr. Lee D. Peachey.
[View Larger Version of this Image (142K GIF file)]
), and with a number of important
prizes. Among his other honors, he was a member of the
National Academy of Sciences, a corecipient with George
Palade and Albert Claude of the Louisa Gross Horwitz
Prize, and a corecipient with Palade and Daniel Mazia of
the first E.B. Wilson Award of the American Society for Cell Biology. The Porter Lecture, named for him, is the
premier lecture at the annual meeting of the society. The
Nobel prize, presented in 1974 for work in which he pioneered, eluded him. In 1977, Porter received the National
Medal of Science from President Carter.
.
1. | Moberg, C.L.. 1996. Keith Porter and the founding of the tissue culture association: a fiftieth anniversary tribute, 1946-1996. In Vitro Cell. Dev. Biol. Anim. 32: 663-669 . |
2. | Palade, G.E.. 1977. Keith Roberts Porter and the development of contemporary cell biology. J. Cell Biol. 75: D1-D19 . |
3. | Peachey, L.D., and B.R. Brinkley. 1983. Scientific achievements and contributions of Keith R. Porter to modern cell biology. Mod. Cell Biol. 2: 1-12 . |