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OCR for page 403
CARL RICHARD MOORE
December 5, ]892-October 16, 7955
BY DOROTHY PRICE
WHEN THE DISTINGUISHED endocrinologist Carl R. Moore
died, a document of great importance to a biographer lay in
his desk. It was headed "Biographical data prepared for files of
National Academy of Sciences," and it was dated August 28,
1948, seven years before his death. Oddly enough, I remember
that day and that occasion. I had gone to his office at the end
of a particularly hot, muggy, Chicago day and had found him
pecking away at an old typewriter in his inimitable fashion
with one finger of each hand. When he said that he was writing
biographical data for the National Academy of Sciences, I may
have been somewhat surprised because he had been a mem-
ber of the Academy since 1944. In any case, my memory
caught and recorded the incident. I was to see the contents of
those pages after his death and to recall him vividly as I knew
him, the professor under whom I wrote my doctoral thesis in
the Department of Zoology of the University of Chicago, the
scientist with whom I served as a close collaborator for many
years in studies on the physiology of reproduction, the chair-
man of a department in which I later became his colleague.
What he wrote in 1948 is remarkable in that he painted,
quite unknowingly, an extraordinarily revealing picture of
his life and scientific career as he saw them. Viewed through his
eyes, his career was the proverbial one of the farm boy of limited
385
OCR for page 404
386
circumstances who makes good mainly by dint of his own hard
work and ambition. It is a typically American story, and in
an ingenuous and engaging way he showed his satisfaction and
pride that this was his story. He had indeed "made good" and
could look with understandable pride at his position and ac-
complishments, his honors and awards.
A biographer who had never known Carl Moore might have
been puzzled to find that more than one third of the six pages
in his biographical notes of 1948 were devoted to a nostalgic
recounting of his experiences on the farm in the Ozark region
of Missouri where he was born and to details of his life and
early schooling in Springfield. To me, this came as no great
surprise. He had often talked at length about his beloved Ozark
country background, and the subject was always close to the
surface of his mind. But when he wrote these biographical
data his thoughts were certainly resting on a past that was
illuminated for him by a rosy light. He had recently been
granted an honorary degree from Drury College in Springfield,
where he had received his B.S. and M.S. degrees. A visit to
the scenes of his early years and a reunion with his family had
undoubtedly revived old memories. But an additional reason
for his preoccupation with his early vears seems Probable. He
BIOGRAPHICAL MEMOIRS
, , 1
had sometimes spoken to me about returning to his Ozark
O
country when he retired, perhaps to a farm near Springfield.
In August of 1948, at the age of fifty-five years, he mentioned
in his biographical notes that he had nine years before official
retirement. He was apparently looking forward as well as
backward, and a plan to retire to an ideal country spot near
Springfield and its Drury College may well have crystallized
in his mind. His life might then have ended where it began,
but death anticipated him, and he did not live to retirement.
Carl Moore was born on December 5, 1892, on a farm in
Green County, Missouri, twelve miles from the city of Spring-
field. His father, whose family was originally of Scottish an-
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C A R L R I C H A R D M O O R E
387
cestry, had been taken from Tennessee to southwest Missouri at
the age of ten in a covered wagon drawn by oxen; Moore's
mother, whose distant ancestors were English, was born in
Missouri just before the Civil War. The farm on which Carl
was born was cleared land cut from surrounding forest. Here
life was simple, frugal, religious, and relatively primitive, and
the boy learned to work hard and to do all the usual farm
chores. He also learned to hunt and fish, and fishing remained
a favorite outdoor sport and means of relaxation all his life.
When he began school, he went to a one-room country school in
which the teacher was one of his older sisters.
A new period began for him when he reached the age of
nine years and his family removed to Springfield, a town of
20,000 people at that time. In this much less restricted environ-
ment, he went through elementary school and high school and
entered Drury College. His family had discussed whether he
should be "a preacher or a doctor," but the matter must have
been settled by the time he registered in college as a premedical
student. His tuition was paid largely from money he earned
by doing all sorts of odd jobs, such as janitor service, window-
washing, and delivering papers. But it was not all work. He
still had time and abundant energy for tennis, horseback riding,
hunting, and fishing.
At Drury College, Carl Moore found the teacher who un-
questionably shaped the pattern of his future life in scientific
research and teaching. Moore should speak for himself on this
point just as he did in his biographical data:
"In college, biology in addition to being a rather natural
interest from earlier farm experiences, became a favorite sub-
ject largely because of the commanding personality of the
teacher, Charles Haddon Spurgeon; he was a self-made, jolly,
fat man, of large physical stature who inspired youngsters by
providing opportunity for work outside the regular curriculum.
Being considerable of a critic, his encouragement and com-
OCR for page 406
388 BIOGRAPHICAL MEMOIRS
mendations were vitalizing. Preparation of slides for histology,
serial sectioning of embryos, and many additional activities in
the laboratory occupied many all night sessions as well as those
on Saturdays and Sundays."
When Moore spoke of Spurgeon's "power over youngsters"
he knew whereof he spoke. Spurgeon's example was firmly im-
printed on him as a young, eager student. He, too, became just
such an inspiring and enthusiastic teacher, and he, too, gave
undergraduate students an opportunity to work on projects
outside the regular curriculum (often in his own research,
with joint publication as an additional bonus). As for dedica-
tion to night and weekend work in the laboratory, Moore was
a convert, and so were those of us who worked with him. In-
deed, it was often necessary for our experiments, but it became
a habit and, ultimately, almost a compulsion. It stayed with
him in the last years of his life during his failing health, and I
would find him in his office on Sunday mornings when he had
hardly strength enough to lift his packed briefcase.
At Drury College, extracurricular laboratory work was a
joy to the young Carl Moore. It held opportunities for ad-
venture and exploration and a chance to solve the problems he
met by some method of his own devising, be it orthodox or un-
orthodox. I can supplement his own biographical notes with a
significant anecdote. When he had difficulty in obtaining good
serial sections of embryos because the paraffin blocks crushed,
he met his problem in a direct fashion by opening all the
windows to let in cold air, putting on overcoat and muffler, and
cranking the microtome around to cut again (with better suc-
cess). There were much simpler and less heroic ways to obtain
perfect serial sections, but he did not know them. It is doubt-
ful that Spurgeon knew much about the matter either, but he
seems to have given his students free rein to use their own
initiative with whatever simple equipment was available and
come up with the best results they could.
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CARL RICHARD MOORE
389
Moore learned these lessons well. In his own direction of
the research of his graduate students, he allowed them free rein
for months at a time to let them "find their own feet" (unless
they came to him for help and advice). Some floundered hope-
lessly and dropped out: some completed their research project
. . .
In a pecrestr~an way on well-worn paths; but some learned the
priceless benefit of being free to develop independence, initia-
tive, and imagination as Moore had learned earlier. They
learned, too, to use relatively simple equipment with only the
minimum of refined instruments. Money was not wasted on
new and showy gadgets—good research did not depend upon
such things. Moore was wont to conduct our distinguished
guests through our research laboratories, operating rooms, and
animal quarters and report to me afterward that he liked to
show them what research could come out of a "setup" like
ours. And he was right in large measure. An ever-increasing
volume of good research was done under relatively primitive
conditions; he did not ask for more. In later years, of course,
he recognized the obvious advantages of modernization and
. . . .
air cone .ltlonlng.
We may be sure that there were no special refinements in
the Biology Department of Drury College when Moore was
studying biology under Spurgeon, learning embryology from
Lillie's Development of the Chick, and enthusiastically carry-
ing on extracurricular projects. He obtained his B.S. degree
in the spring of 1913. His family had no money to send him to
medical school, but another opportunity opened. He was of-
fered a position at Drury as an assistant in biology. The posi-
tion carried a munificent salary of $100 for the year, and he
could work under Spurgeon for an M.S. degree (incidentally,
one of the very few ever granted by that institution, as Moore
states in 1948~. Moore snapped at the chance. But before he
began his fifth and final year at Drury, he came to the University
of Chicago and registered for summer quarter courses in the
OCR for page 408
390
Department of Zoology. Lillie's textbook had caught his inter-
est and piqued his curiosity about the department where the
famous embryologist was chairman. Of course, Lillie was not
there—he spent every summer at the Marine Biological Labora-
tory at Woods Hole—but there was much to see and much to
learn at the University of Chicago, and it was Moore's first trip
away from home.
To obtain his M.S. degree at Drury, which he did in June
of 1914, Moore assisted in courses and made what he later
termed "an attempt at research" on the origin of the vena cave
in bat embryos. He prepared slides of serial sections of embryos,
projected the sections on melted beeswax, and by cutting out
the projected sections and stacking them he produced models.
He had never seen a wax model, but he made some in his own
way (undoubtedly with the window open). This research
problem might not seem the most interesting one for a young
student. For him, it was an exciting foretaste of biological re-
search, and he was thenceforth lost to medicine.
He had applied to some universities for support for graduate
study and received offers of fellowships from three. The one he
accepted was from the University of Chicago, and in choosing
this fellowship he made one of the most important decisions
of his life. He would have made a name for himself wherever
he had gone, but it is open to doubt whether he would have
advanced as rapidly and his name have loomed so large if he had
gone elsewhere. Moore came to work with the right man-
Frank R. Lillie—at just the right time in the development of
Lillie's research program. The Department of Zoology was an
ideal environment in which Carl Moore could mature; when he
received his Ph.D. degree he was to step into that department
as a member just before the beginning of the 1920's, a decade
of great and brilliant advances in endocrinology. And Moore
was to be in the middle of it all.
However, Moore could not gaze into the crystal ball. When
BIOGRAPHICAL MEMOIRS
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CARL RICHARD MOORE
391
he went to Woods Hole for the summer session in 1914 and met
Frank R. Lillie for the first time, the young man fresh from
the Ozark hills was in a state of "uncertainty and trepidation"
as he described it. Others, too, had found the dignified, reticent,
soft-spoken embryologist overawing. But this first meeting for
Carl Moore was a successful one, and he was assigned a doctoral
problem in Lillie's large program of studies on the fertilization
of the eggs of marine invertebrates.
Lillie became his mentor, and Moore began to learn eagerly
the elements of sophisticated scientific research and criticism
from one of the best possible teachers. Moore always remained,
in a sense, Lillie's protege, and Moore repaid him with profound
admiration, respect, and affection. Fortunately, the young stu-
dent did not try to emulate too closely the middle-aged man
of great distinction who patiently directed him. That would
have been disastrous; their backgrounds and personalities were
very different. But when Moore later gave the seminars for
which Lillie had been famous, "Biology of Sex" and "Physiology
of Reproduction," he followed almost exactly not only the
lucid organization but much of the subject matter of Lillie's
brilliant introductions.
Moore completed his doctoral thesis on fertilization and
parthenogenesis in the eggs of a sea urchin in record time and
received his Ph.D. degree from the University of Chicago in
1916. He was immediately appointed an associate in the De-
partment of Zoology for the period from 1916 to 1918. He spent
half the time in teaching an embryology course designed
primarily for premedical students and the remainder in re-
search. In 1918 he became instructor, and in the ensuing years
he advanced rapidly, reaching a full professorship in 1928 and
the chairmanship of the department in 1934.
In the period from 1919 to 1920 an event of great importance
in Carl Moore's life and in his scientific career occurred. A
student named Edith Naomi Abernethy caught his attention in
OCR for page 410
392
BIOGRAPHICAL MEMOIRS
a laboratory section he was teaching. Soon his interest was more
than academic, and they were married in July of 1920. He
acquired not only an attractive and charming wife, but a hostess
who presided with grace and competence on the many occasions
when they entertained students and scientific visitors from many
places in America (in the broad sense) and abroad. She under-
stood his need to consider his laboratory also a "home," and
she shared his love of nature and the outdoors. Their summer
home in Michigan was a haven for him and a beloved spot for
her. Their honeymoon was spent on a float trip on a river in
the Ozarks (where else could it have being. Two of their three
children survived—Harris Mason and Ellen Abernethy.
After publishing his thesis, Moore completed a second paper
on the sea urchin, Arbacia, and then turned abruptly to an
entirely different line of research in which he was occupied
for the rest of his scientific career. The reason was clear, but it
requires explanation. Lillie published in 1916 the first of his
classic papers on the freemartin, a bovine intersex that resulted
from cases of heterosexual twinning when there were anasto-
moses of blood vessels in the fused fetal membranes. The type
of intersexuality and the sterility usually found in freemartins
posed intriguing problems. Lillie's observations and his master-
ful analysis resulted in a theory to explain freemartinism on the
basis of masculinization of the female by male hormone pro-
duced in the testes of the male co-twin and crossed to the female
through the vascular anastomoses. The ovaries were inhibited
(antagonized), and the duct systems and glands were masculin-
ized (stimulated). He then proposed that normal sex differentia-
tion in the mammalian fetus might be controlled by bloodborne
substances, hormones, secreted by fetal testes and ovaries. How-
ever, he cautioned that the theory was only tentative. Nothing
was known about the possible effects of female hormone on the
male fetus. Fetal gonadectomy—all-important for an under-
OCR for page 411
CARL RICHARD MOORE
393
standing of normal sex differentiation—had never been accom-
plished and should be done as a critical test of the theory.
Lillie made "a mild suggestion" (in Moore's words) that
Moore try to produce freemartins by experimental means. A
mild suggestion was all that was required, and Moore plunged
into the problem, or rather into the problems, for his research
did not follow any straight path. Thirty-eight years and some
one hundred publications later, he still had not produced free-
martinism experimentally (nor, indeed, had anyone else). But
by then he had developed a theory of his own to explain normal
sex differentiation. This proposed that sex hormones from fetal
gonads were not controlling normal sex differentiation. The
evidence that he presented negated, in his opinion, Lillie's
hormonal theory for freemartinism and its extension to normal
sex differentiation. However, before Moore was led to this
conclusion he made many outstanding contributions in the
physiology of reproduction even as he digressed from his original
purpose.
Moore began his attempts to subject fetuses to male hormone
and produce experimental freemartinism by transplanting
testicular tissue onto the membranes of rat and guinea pig
fetuses. This failed dismally. Then he transplanted testes into
young females in the hope of obtaining freemartins in the litters
when these females were ultimately bred. This, too, failed, but
he was far from discouraged. He had succeeded in obtaining
well-developed testicular grafts in young females. The field
of gonad transplantation with its postulates of sex gland an-
tagonism lay open before him.
The Viennese scientist E. Steinach had first reported in
1910 that young spayed female rats and guinea pigs were
masculinized when given grafts of testes, and young castrated
males were feminized by grafts of ovaries but the hosts must be
gonadectomized prior to receiving grafts. He therefore pro-
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394
BIOGRAPHICAL MEMOIRS
posed and strongly defended, then and later, a concept of sex
gland antagonism involving a direct antagonism between testis
and ovary.
The question of whether sex gland antagonism really existed
was an important one. The validity of such a concept might
have a direct bearing on Lillie's theory of freemartinism. Moore
promptly sought confirmation or refutation of the concept and
soon completely disproved it. He was successful In maintaining
testis grafts in young females possessing ovaries and ovarian
grafts in young males with testes. In these early experiments
of Moore's there was a fortuitous circumstance that undoubtedly
favored unusually successful "takes" of the grafts. He routinely
exchanged one gonad each between young males and females
at the time of grafting, apparently for efficient utilization of
animals. Thus the grafts were placed in hosts that had just
been hemispayed or hemicastrated. About ten years later, he
and I were to propose a theory of balanced control between
gonadal hormones and nituitarv ~onadotronins.
, c7
rid --, =_
The experi-
mental design that he used in 1919 had a sound rationale of
which he never dreamed at the time. The grafts that he ob-
tained differed so materially from those described by Steinach
that he was led into his next research problems.
Moore's grafts of testes had far better-developed tubules and
fewer interstitial cells between them than those of Steinach.
The two investigators agreed that spermatozoa were not present
in the grafts. Contrary to Steinach, Moore found no evidence
for increased hormone secretion and no basis for a contention
that testis grafts might effect rejuvenation in senile animals and
men.
This fata morgana of warding off all changes of aging in
man, or rejuvenating the senile, had appeared again. Testis-
secreted hormone was the miraculous cure-all. In France, the
Russian-French surgeon S. Voronoff was rejuvenating senes-
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CARL RICHARD MOORE
BIBLIOGRAPHY
403
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OCR for page 425
CARL RICHARD MOORE
407
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OCR for page 431
Representative terms from entire chapter:
sex differentiation
