BIOGRAPHICAL MEMOIRS
IAN FLEMING/PEMBROKE COLLEGE CAMBRIDGE
RICHARD HUME ADRIAN, SECOND
BARON ADRIAN OF CAMBRIDGE
16 October 1927 . 4 April 1995
PROCEEDINGS OF THE AMERICAN PHILOSOPHICAL SOCIETY
VOL. 145, NO. 1, MARCH 2001
richard hume adrian
R
ICHARD HUME ADRIAN , second Baron Adrian of Cambridge, was a man of many talents, and he applied these
with great effectiveness and integrity in a wide range of
endeavors during his lifetime. As a young man he enjoyed the excellent
educational opportunities that life in Cambridge and his strongly professional and intellectual family background provided him. In his professional life, he became a leading electrophysiologist, unraveling the
mysteries of the electrical properties of muscle cells in a scientific career
spanning three decades. Later in life, he devoted his considerable energy
and wisdom to more administrative and political (in the sense of policymaking) activities, both in Cambridge University and its colleges, and in
the British House of Lords. Those fortunate to have been among Richard
Adrian’s colleagues and friends respected and admired him for his absolute and unwavering honesty and fairness in every aspect of his distinguished career in science and education. He was my close friend and a
most valued colleague, and I shall forever cherish his friendship, his wisdom, his good humor, and his stellar example of what a life should be.
Early Life and Education
The Adrian line in England has been traced to a Huguenot refugee,
also named Richard, who fled from France or Flanders after the massacre of St. Bartholomew in 1572. Eleven generations later, Richard H.
Adrian was born on 16 October 1927 at his parents’ home in Grange
Road, Cambridge, near the “Backs” of the university. He was the only
son of the distinguished neurophysiologist E. D. Adrian (1889–1977),
Nobel laureate in 1932, and later first Baron Adrian (created 1955).
E. D. Adrian had been a student of physiologist Keith Lucas at Trinity
College in Cambridge, and had collaborated with many distinguished
physiologists, including Yngve Zotterman, Rachel Matthews, B.H.C.
Matthews, and Detlev Bronk. It was through his mother, Hester Agnes
Adrian (née Pinsent), that Richard Adrian gained his middle name: she
was related on her father’s side to the philosopher David Hume. Both
she and her mother were made D.B.E. for their work on mental health.
Richard had an older sister, Anne, and a twin sister, Jennet, both of
whom survive him.
Richard attended King’s College Choir School in Cambridge until
World War II broke out in Europe. Then he and his twin sister Jen were
sent to the United States, as were many children of professional English
families. The two twelve-year-old children came in June 1940 to the
suburban Philadelphia Sycamore Mills home of their father’s friend
and colleague Detlev Bronk and his wife, Helen. Richard and Jen lived
partly at the Bronks’ house, and partly in Swarthmore with Robert and
[84]
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Abbie Enders. Enders was professor of comparative anatomy at Swarthmore College, and his wife was an acknowledged expert in the field of
early education in urban areas. Exposure to a strongly intellectual
home atmosphere continued to be a constant in the early lives of these
two English children during their time in the United States.
Because the Enderses’ house was convenient to Swarthmore High
School, Richard tended to live there during the school week and at the
Bronks’ house on the weekends. Ramsey Bronk, who was a year younger
than Richard, recalls that conversations between Richard and the senior
Bronk at Sunday meals frequently were extremely lively, and even argumentative. We all recognize that argument is more often allowable
between a father and another man’s son (i.e., between a boy and another
boy’s father) than it is between a father and his own son. In later years,
Richard fondly recalled these stimulating weekends with the Bronks,
both at Sycamore Mills and at the Bronks’ summer house in Woods
Hole, Massachusetts, where many U.S. and foreign physiologists still
move their laboratories in the summer to take advantage of its intellectual atmosphere and its ample supply of marine organisms for experimental study.
In 1941, Richard spent the entire summer with the Bronks at
Woods Hole. There he met Herbert Gasser, who was head of the Rockefeller Institute for Medical Research in New York and a pioneer in the
field of electrophysiology. This meeting and other contacts with scientists made during that summer at Woods Hole almost certainly influenced Richard Adrian’s later decision to choose physiology for his
vocation. Richard said in the Henry LaBarre Jayne Lecture to the
American Philosophical Society on 22 April 1988 that it was an experience in hospital in 1941, when his appendix was removed by Dr. Jonathan
Rhoads, that turned him toward medicine and an academic career. Perhaps this explains his choice of medicine for his university education.
But one must think that it was Gasser and biophysicists such as Britton
Chance, whom he also met at the Bronks’ house, that led him to
choose for his research the rather biophysical branch of physiology
that deals with the electrical properties of cellular membranes, the field
in which Richard later gained his own international reputation.
Richard and Jen lived out the remainder of the European phase of
the war with the Bronks and Enderses in Pennsylvania. He graduated
from Swarthmore High School in 1943, at the relatively young age of
fifteen, and spent the following summer as an undergraduate at Swarthmore College. In November of that year, he returned to England to
continue his education for one year at Westminster School, which had
also been his father’s school. Richard went up to Trinity College, Cambridge, in January 1945. He graduated from Cambridge University in
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Top left, Richard Adrian as a boy
in Pennsylvania, with his father,
who was visiting in the spring of
1942; top right, sailing in Musa
at Norfolk after the war; middle,
in his Bentley on the Norfolk
coast at Easter of 1958; bottom
left, in his physiology laboratory
in Cambridge about 1960. Photographs courtesy of Lucy Adrian.
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1948, after reading the natural science tripos (medicine), Part I in anatomy, physiology, biochemistry, and pathology for two years, and Part
II in physiology. This preparation led to clinical study in London at
University College Hospital. He qualified M.B., B. Chir. in 1951.
After two years of national service in the Royal Army Medical
Corps at Porton Down, from 1952 to 1954, Richard returned to Cambridge University and to Trinity College, and started physiological
research under the supervision of Alan Hodgkin. This was an exciting
time at the Cambridge Physiological Laboratories. Hodgkin and Andrew
Huxley had just finished their experimental and mathematical analysis
of the action potential in the squid giant axon. The action potential is
an electrical disturbance that spreads along a nerve fiber and carries
signals to other nerve cells and to muscles, causing them to contract.
This was a great triumph in electrophysiology and the work for which
Hodgkin and Huxley later received the Nobel Prize. In a real sense, this
study of the squid axon marked the true emergence of the nascent field
of electrophysiology as a research area of the utmost importance and
potential in biology and medicine.
Scientific Research
Encouraged by Hodgkin and Huxley’s success with the squid nerve
axon, Richard Adrian began a study of the electrical properties of striated muscle cells, which display action potentials at their surface membranes very similar to those of squid axons. The properties of muscle
cells were known to exhibit several important differences from those of
squid axons, in addition to the obvious difference that muscle cells
contract following an action potential, whereas squid axons do not.
Muscle cell action potentials have considerably longer durations than
those of axons, and are conducted along the cell much more slowly: it
was suspected that this relative slowness might be related to the much
larger electrical capacity that muscle cells have compared with nerves.
Electron microscopy of muscle cells by Keith Porter and others suggested that this larger electrical capacity arises from an expansion of
the surface area of the muscle cell in the form of extensive invaginations of the surface membrane into the interior of the cell: more surface
membrane, greater capacity. These tubular invaginations, which are
not found in nerve cells, connect together inside the muscle cell to form
complex networks, known collectively as the transverse tubular system, or T-system. Much of Richard Adrian’s scientific work was related
in one way or another to these specialized structures and their associated electrical properties in muscle cells.
It was characteristic of Richard Adrian to be extremely cautious in
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his experimental work. Typically he would hold off beginning any
experimental study until he felt completely comfortable with his understanding of the theoretical basis of the experimental approach he planned
to use, and most assuredly with the adequacy and perfection of the
experimental setup itself. Early in his work on muscle, he made a significant technical advance in what probably started out for him as a
routine check on the accuracy of the microelectrodes used to record the
electrical potentials inside cells. These microelectrodes are produced by
heating a small, hollow tubing of glass, and suddenly pulling the softened glass out so that it forms a very small tube that eventually breaks,
leaving an open tip with a diameter of about 1 micrometer or even less.
These extremely thin capillary tubes are filled with a strong electrolyte
solution to serve as a conductor of electricity, and then their fine tips
are carefully inserted into a cell through its surface membrane. Once
inside the cell, the electrode tips sample the intracellular electrical potential, which then can be measured by the experimenter at the larger end
of the electrode outside the cell.
One’s natural expectation for these electrodes would be that they
should record no potential when their tips are outside the cell, and then
accurately report the internal potential after they cross the membrane
and enter the interior of a cell. With characteristic thoroughness, Adrian
carefully checked this point, placing electrodes into the physiological
solution that would bathe the experimental cells, but without putting
the electrode tips into a cell. He then measured the electrical potential
reported by the electrodes. The expected result of this control experiment, zero potential, was frequently the result he obtained. But some
electrodes erroneously reported potentials significantly greater than
zero while still outside the cell. These electrodes could not be relied
upon to measure intracellular potentials accurately in a real experiment and they had to be discarded. Following publication of these
results, physiologists routinely checked and selected their electrodes
before using them. Adrian received much recognition for this technical
advance, in the form of bibliographic credit in experimental papers
written by other physiologists. He often expressed a modest feeling
that such recognition for such a small and obvious observation was
much overdone. To him it was, after all, only a matter of taking proper
care in carrying out one’s experimental work!
Adrian’s early experimental work on muscle, extending from 1956
to about 1966, was concerned primarily with the idea that the electrical potential inside a cell is due largely to the higher concentration of
potassium ions inside the cell compared with outside the cell. The simple idea is that potassium ions, which carry a positive electrical charge,
diffuse from inside to outside the cell through pores in the cell’s surface
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membrane, and that this outwardly-directed diffusion of positivelycharged potassium ions results in a deficit of positive electrical charge
inside the cell. This process continues until a sufficiently large negative
potential builds up inside the cell to provide a counter electrostatic
force to attract potassium ions back into the cell at the same rate at
which the concentration difference drives them out. Achievement of
this equality of inward and outward rates of diffusion results in a
steady-state condition in which there is no net movement of potassium
ions in either direction, and a steady negative electrical potential is
maintained inside the cell. This is known as the “resting potential” of
the cell. It can be measured by inserting the tip of a glass microelectrode into the cell, as mentioned above.
In a series of papers in the Journal of Physiology, Adrian first confirmed in muscle cells what had been seen earlier in nerve cells: altering
the external potassium concentration in the fluid surrounding muscle
cells alters the resting potential in the direction one would predict and
by the amount expected from the mechanism discussed above. Adrian
also performed a series of experiments in which he altered the internal
potassium ion concentration by exposing the muscle cells to solutions
with different osmotic pressures, which draws water into or out of the
cell. Again, the resulting resting potentials agreed with the idea that they
are due to potassium diffusion into and out of the cell under oppositely
directed concentration and electrical driving forces. An additional and
important result of these experiments was the observation of an “anomalous” difference in the ease with which potassium ions move through
the muscle cell’s membrane in the two directions, which can be much
greater in the inward direction than in the outward direction. This somewhat unexpected phenomenon has been called “inward rectification.”
Apparently it is specific for skeletal and certain cardiac muscle cells.
This initial work was important because a major unanswered question in muscle cell physiology at that time was how the signal that
causes a muscle cell to contract following an action potential on its surface is conveyed from the surface of the cell to its deep interior, where
the contractile apparatus is found. If muscle cells are like other kinds of
cells, including nerve cells, then the ionic currents responsible for its
action potentials should be confined to the membrane at the surface of
the fiber, a long distance from much of the contractile apparatus.
Adrian’s earlier results had defined some important differences between
the electrical properties of muscle cells and those of other cells. It
seemed apparent that these differences might provide clues about how
muscle cells are able to signal deep into the cell. Most of Adrian’s
experimental work for the remainder of his scientific career was
devoted to investigating this coupling problem. Much of his work
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was done in the Physiological Laboratories, initially in collaboration
with Alan Hodgkin and, many years later, Chris Huang. Through the
middle of his career, Adrian worked in Cambridge with a series of visiting investigators from the United States, including Walter Freygang
from NIH, Knox Chandler from Yale, the late Roy Costantin from
Washington University, Wolf Almers from the University of Washington, and the writer of this memoir. Adrian also made extended trips to
work in the laboratories of these collaborators.
Details of these technically rather complex studies would not be
appropriate for inclusion in this memoir, the main goal of which is to
“capture the man,” but they can be summarized in moderately nontechnical terms, emphasizing the contributions made specifically by
Richard Adrian.
In 1966, Adrian made a second important technical contribution to
muscle physiology by showing how three microelectrodes could be
inserted into a single muscle cell, near one of its ends, and how these
electrodes could be used precisely to control the electrical potential difference across the surface membrane of the cell. In effect, this technique converts the experimenter from a mere observer of the potential
changes resulting from the cell’s own activity into someone who can
dictate the potential changes that the cell experiences. This greatly
expands one’s ability to discover just what is going on in the cell’s
membrane during an action potential and precisely how this relates to
contraction activation. Adrian used this “voltage-clamp” method with
Hodgkin and Chandler from 1965 to 1967 to explore in detail the relationship between membrane potential changes and the resulting contraction of the muscle cell. From 1967 to 1969, with Costantin and
Peachey, Adrian extended these studies by observing microscopically
the small movements of the contractile apparatus just at the onset of contraction, and exploring how this contraction spreads toward the center of
the cell from its surface as the surface membrane potential is changed by
small increments. These studies provided the first direct observation of
the process of the spread of activation of contraction from the surface
to the center of the muscle cell under conditions in which the membrane potential could be controlled and known with precision.
These results also provided the experimental background for a theoretical analysis of this inward spread of the activation signal in the
T-system, designed to see if such electrical effects in the tubules of
the T-system could provide a feasible mechanism for coupling the surface
action potential to contraction inside the muscle cell. And this is where
Adrian’s innate mathematical ability came to the fore. Initially his calculations were based on a simple model of “passive” spread of the electrical signal. A better fit to the experimental data was achieved later by
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assuming that an action potential mechanism similar to the one described
by Hodgkin and Huxley in nerve cells was present not only on the surface of the muscle fiber, but also in the T-system. These experimental
and theoretical studies, taken together, provided strong support for the
idea that an electrical activation signal spreads all the way to the center
of a muscle cell via the T-system, assuring that the entire cell can participate in its contraction.
Richard Adrian’s subsequent scientific research focused on understanding a small but essential step in the electrical events just described.
If a change in electrical potential difference across the T-system membrane is to trigger some further event (e.g. contraction) inside the cell,
there must be some molecule or group of molecules in the T-system
membrane that “senses” that the potential difference has changed.
This event would be detected experimentally as a movement of electrical charges within the membrane. Detection of these minuscule movements of charge is extremely difficult. Again Adrian, this time in
collaboration with Wolf Almers, first recorded such events in 1976
in experimental studies on frog muscle cells. Later studies by Adrian in
collaboration with A. Peres from Italy and with Chris Huang, and by
other laboratories, provided essential information on these charge
movements, which can be thought of as the last electrical step in the
coupling of action potentials to mechanical contraction. Beginning in
1956 and extending up to 1984, Adrian was a prime figure in the investigation of all the major electrical steps involved in coupling the cell’s
action potential to its contraction.
Through all of these studies, Richard Adrian was a constant source
of ideas and encouragement, a consummate experimenter, and a
superb interpreter of the observations. Words I wrote to Barry Till,
which he quoted at a memorial service for Richard Adrian in Cambridge on 24 June 1995, were my attempt at explaining why Richard
Adrian was so successful in his scientific research: “He always seemed
to be ahead of us in thinking about the experiments. Somewhere here is
one of the keys to his abilities as a scientist; he combined great practical ability with equipment and in the design of experimental protocols
with a keen theoretical understanding of the physics and mathematics
of the biological problem. That may sound simple and straightforward, but it is a rare and valuable talent. He knew when to use his
hands and when to use his brain, and all three were very keen.”
Academic Career
Richard Adrian remained at Cambridge for his entire academic and
research career, moving through the ranks to the highest level. His first
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college was Trinity, where he was both an undergraduate and a
research student. He then was a research fellow (1955–56) and director of medical studies and dean (1956–60) of Corpus Christi College.
When Churchill College was established in 1960, he became a founding fellow and tutor for natural sciences. He maintained his fellowship
at Churchill until 1981, when he became master of Pembroke College.
Within the university, he was university demonstrator in physiology
from 1956 to 1961, lecturer from 1961 to 1968, reader in experimental biophysics from 1968 to 1978, and professor of cell physiology (ad
hominem) from 1978. While master of Pembroke College, he served as
vice chancellor of the university for a standard two-year term from
1985 to 1987. He retired from the college and from his professorship
in the university in 1992.
Adrian was honored by election as a fellow of the Royal Society in
1977 and of the Royal College of Physicians in 1987. He served on the
council of the Royal Society for 1984. He also was a trustee of the British Museum from 1979 to 1993 and of the British Museum (Natural
History) from 1984 until 1988. He served as a member of the British
Library Board from 1987 to 1993. He was on the council of the Baring
Foundation, a trustee of the Daiwa Anglo-Japanese Foundation, and a
governor of his old school, Westminster, and of the Imperial College of
Science and Technology. His contributions to science were recognized
with a degree as docteur honoris causa from Poitiers in 1975 and honorary fellowships at Churchill College and Darwin College at Cambridge.
In 1993, Adrian was appointed deputy lieutenant for Cambridgeshire,
a rather unusual distinction for an academic and an indication of his
success in bridging the town/gown divide.
House of Lords
Following his father’s death in 1977, Richard Adrian succeeded to his
father’s peerage and became the second Baron Adrian of Cambridge,
taking his seat in the House of Lords with a maiden speech on the care
and use of animals in medical research. Already in that speech there
were indications that he would become a significant force in that august
body, just as he had already become in the academic and scientific
world. Using carefully chosen words, the new Lord Adrian argued the
importance of the use of animals in the study of human diseases. He cleverly selected as his examples of medical advances that had depended critically upon animal research those disorders that he knew might well be
of special interest to his distinguished colleagues, who tended to be of a
certain age.
At this point in his life, Lord Adrian debated with himself the diffi-
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cult balance between his scientific research and teaching in Cambridge
and his participation in the House of Lords. He was well aware, and I
am sure he was reminded frequently by his academic colleagues, of the
need for a voice of reason backed up with strong academic credentials
in debates in the House of Lords on matters pertaining to higher education and scientific research. Beginning in about 1980, Lord Adrian
began to devote a greater part of his time and energy to participation
in the activities in the chamber of the House of Lords and as a member
of the Select Committee on Science and Technology. He became a vigorous defender of the independence of universities from unnecessary
and excessive government regulation and interference. He spoke with
considerable force about the arrogance of the British government and
its conviction that it alone knew how a university should be run. He
believed that it was dangerous and corrupting for institutions that
should be independent (such as universities) to rely too much on
money from the government. He led a cross-bench opposition to certain features of the Education Reform Bill of 1988, which he believed
to be a threat not only to Oxbridge but to all British universities. In the
end, this opposition secured an important amendment to the bill, guaranteeing a considerable insulation of the academic, teaching activities
in the universities from the source of funds through the government’s
University Grants Committee. This remains today as an important pillar supporting academic freedom in Britain.
Lord Adrian approached his work in the House with characteristic
thoroughness, always acquainting himself fully with the niceties of the
situation and arguments in advance of the debates. He was driven by a
fierce determination that, in every case, a just decision should be
reached. His effectiveness in the House clearly derived not only from
his intelligence and his skill as a speaker, and from his distinguished
academic career and extensive academic experience, but also from the
independence gained by his choice to sit on the cross-benches, where he
was less likely to be given a particular political label. One of his colleagues in the House has said that he didn’t realize how important Lord
Adrian’s thorough preparation was until he had to enter a debate without it. His academic colleagues, including many who were never privileged to know him, owe him a great debt for his efforts in London on
their behalf.
Personal Life
In 1967, Richard Adrian married Lucy Caroe, a historical geographer
with a special interest in the rural economy of nineteenth-century Britain, and a fellow of Newnham College Cambridge. Lucy’s family back-
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ground is no less intellectual and distinguished than Richard’s. Her
father and grandfather were the architects A.D.R. and W. D. Caroe,
while on her mother’s side she is the granddaughter of Sir William
Bragg, the English physicist and pioneer in x-ray diffraction. Lucy gave
unsparingly of herself to help Richard, and he valued her advice
immensely. When he became master of Pembroke College and vice
chancellor of the university, assuming positions that carry both heavy
responsibilities and considerable social and ceremonial obligations,
Lucy participated in these with charm and consummate skill.
On becoming master of Pembroke College in 1981, Richard spoke
of his personal delights with his usual honesty and humility, saying,
“Among my pleasures are (or were) climbing, sailing, skiing and driving fast cars fast; in none of these was I any better than average.” As a
young man he rode a motor bike, wearing what his sister Jen has
described as “a fabulous leather kit.” His massive Bentley, which
appears in one of the family pictures reproduced here, was from his time
at Porton Down. A red Austin-Healy, and then a Mini Cooper-S followed. I remember flying along the narrow and curved roads of Norfolk
at a seemingly impossible speed in that little but powerful Mini, with
Richard coolly in control at the wheel. Later in his life, the Cooper-S
yielded to a Rover, then to a minivan of some sort, and, last, to a VW
Passat: his practical side took over from the young bachelor, though I
suspect not without some struggle!
In addition to being a gifted experimentalist in the laboratory,
Richard was a competent and enthusiastic woodworker. He planned,
on retirement, to extend the woodworking skills derived from building
model airplanes as a boy, and the lessons learned in “shop” at Swarthmore High School. He had already started assembling a shop in his
garage in Norfolk when he retired. He had a keen eye for quality in
antique furniture and in craftsmanship of all kinds. This made his association with the Goldsmiths’ Company, one of the Great Twelve of the
City of London Livery Companies, a special delight for him. There he
served as prime warden from 1990 to 1991, presiding at feasts and ceremonies in their newly renovated hall. A commemorative medal was
designed by the artist Philip Nathan and struck at the Royal Mint to
mark Richard’s prime wardenship. Following tradition, the obverse of
this medal shows the honoree’s head. Following Richard’s wishes, the
reverse features a scientific drawing of part of a muscle cell, together
with the insignia of Pembroke College, the Goldsmiths’ Company, and
the Adrian family. This balance among family and academic background on the one hand, and science on the other, demonstrates his
continuing commitment to these multiple aspects of his life and career.
Richard Adrian retired from his university position as professor of
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cell physiology and from the mastership of Pembroke College in 1992.
He and Lucy had put considerable time and effort into renovating
Frostlake Cottage in Malting Lane in Cambridge as a house to retire
to, and planned to live there and at their home, Umgeni, at Cley-nextthe-Sea on the north Norfolk coast. However, after only six months of
retirement, Richard was stricken with abdominal pain while attending
a reception given by the American Philosophical Society at the University of Pennsylvania Museum during its April 1993 meeting. By chance,
at the time he took ill, he was in conversation with Dr. Jonathan
Rhoads, emeritus professor of surgery at the hospital of the university,
and the surgeon who had removed Richard’s appendix at that hospital
in 1941. Richard was taken immediately across Thirty-fourth Street to
the university hospital. When he was released two days later, he and
Lucy canceled trips to Tokyo for a meeting of the Daiwa Foundation
and to Melbourne, where he was to speak on university reform in the
United Kingdom. They extended their stay in Philadelphia, staying
with Helen and me at our house in Narberth for another week, until he
felt able to return to Cambridge. Later that spring came a diagnosis of
cancer. When he died on 4 April 1995, the world lost a great scientist
and statesman. His family and his colleagues, including myself, lost a
great friend. He was a model for us, a model of honesty and sincerity
in science and in life, and we miss him greatly.
I cannot close without recollecting our long and close relationship.
We worked together over two decades, in England and in the U.S. We
taught muscle physiology courses in Colorado and Mexico. We attended
scientific conferences together, often adding an excursion. We visited
Indian cave temples in Aurangabad and Ellora after meetings in New
Delhi and Bombay. We backpacked in the Colorado Rockies, and we
and our wives combined camping in the deserts of Colorado and New
Mexico with a Strauss Salome in Santa Fe. We also sailed on the Norfolk coast in a boat Richard had built himself. Lucy Adrian has said
that her husband greatly enjoyed going to “out-of-the-way” places
with me, and staying in the “economical” hostels I tended to choose.
Equally, he introduced me to some “upscale” places his life took him:
lunches in the House of Lords and dinners at some of the best restaurants in London. I was at his maiden speech in the Lords, though he
didn’t even know I was in England until I suddenly appeared. Richard
was a most generous and understanding friend. I enjoyed his company
immensely in all circumstances. I experienced and learned much that I
would have missed had he not been my friend.
I close by repeating, in honor of Richard Adrian, some words that
Benjamin Franklin penned in honor of one of his own dear friends and
colleagues,
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whose virtue and integrity, in every station of life, public and private,
will ever make his Memory dear to those who knew him, and who
knew how to value him.1
Elected 1987
Lee D. Peachey
Professor of Biology
University of Pennsylvania
Acknowledgments
The writer would like to express his deep appreciation to Ramsey
Bronk, Richard Keynes, Peter Matthews, Barry Till, Richard’s sisters
Anne Keynes and Jennet Campbell, and most especially, to Lucy Adrian,
for providing insight and information about Richard. Without their
help I would not have been able to speak even as imperfectly as I have
about his remarkable life.
Selected bibliography for Lord Adrian
1956. The effect of internal and external potassium concentration on the membrane
potential of frog muscle. J. Physiol. 133: 631–58.
1960. Potassium chloride movement and the membrane potential of frog muscle. J.
Physiol. 151: 154–85.
1969. The kinetics of mechanical activation in frog muscle (with W. K. Chandler and
A. L. Hodgkin). J. Physiol. 204: 207–30.
1969. Radial spread of contraction in frog muscle fibres (with L. L. Costantin and L. D.
Peachey). J. Physiol. 204: 231–57.
1970. Voltage clamp experiments in striated muscle fibres (with W. K. Chandler and A. L.
Hodgkin). J. Physiol. 208: 607–44.
1973. Reconstruction of the action potential of frog sartorius muscle (with L. D.
Peachey). J. Physiol. 235: 103–31.
1976. Charge movement in the membrane of striated muscle (with W. Almers). J. Physiol. 254: 339–60.
1979. Charge movement and membrane capacity in frog muscle (with A. Peres). J.
Physiol. 289: 83–97.
1983. Electrical properties of striated muscle. Pp. 275–300 in Handbook of Physiology, Section 10, Skeletal Muscle, ed. L. D. Peachey, assoc. ed. R. H. Adrian. American Physiological Society, Bethesda, Md.
1984. Experimental analysis of the relationship between charge movement components in skeletal muscle of Rana temporaria (with C.L.-H. Huang). J. Physiol.
353: 419–34.
1988. The crisis in British Universities. The Henry LaBarre Jayne Lecture. Proc. Amer.
Philos. Soc. 132: 237–46.
1 Whitfield J. Bell, Jr., Patriot-Improvers: Biographical Sketches of Members of the
American Philosophical Society, vol. 1 (Philadelphia: American Philosophical Society, 1997),
12–13.