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~ " 6 I'd
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OSCAR RIDDLE
September 27, Z877-November 29, 1968
BY GEORGE W. CORNER
OSCAR RIDDLE, zoologist and proponent of the freedom of
science teaching, was born September 27, 1877, in Greene
County, Indiana. His birthplace was a log house near a village
called Cincinnati, twenty miles from the university town of
Bloomington. His boyhood in this countryside of heavily
wooded hills and narrow valleys is well described in autobio-
graphical notes Riddle prepared for the files of the National
Academy of Sciences. The first part of the present account of his
life, scientific career, and writings largely follows his own
narrative.
Oscar Riddle's father, Jonathan Riddle, came from a North-
of-England family that had settled first in Virginia. On his
Indiana land he made a comfortable living by farming and
breeding livestock, though always with the narrow economic
margin characteristic of pioneer life. He kept a racehorse and
was an enthusiastic hunter of deer, wild turkey, and the bears
that were then to be found in the hills of Indiana and neigh-
boring states. It is of interest, in connection with his son's
attitude toward dogmatic religion, that Jonathan Riddle was
. . . .
never active In any re lglOUS sect.
Oscar's mother, Amanda Emeline Carmichael, was born at
Cincinnati, Indiana, of Scottish and northern Irish ancestry.
Her father, relatively prosperous among the villagers, kept a
427
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428
BIOGRAPHICAL MEMOIRS
general store and a flour mill. Something of a philosopher, Mr.
Carmichael wrote a number of unpublished essays, well worded
(according to his grandson) and showing deep interest in the
question of free will and similar religiophilosophical topics.
Sometimes, in the absence of the local Baptist preacher, he took
the pulpit and preached sermons appreciated by the congre-
gat~on.
Jonathan Riddle died in 1882 at the age of fifty-five, leaving
nine children of whom the youngest was six months old. His
wife, then forty-five years of age, raised all of them to maturity
and lived on to the age of eighty-nine. Although during her
husband's lifetime she professed no religious faith, after he died
she joined the local Baptists, often remarking, however, that site
did not believe In eternal punishment nor did she think it
sinful not to be a professed Christian. Her husband's death left
the family in straitened circumstances, and all the older children
had to help with the farm work. In his written reminiscences
Oscar gives a graphic account of his early boyhood and
schooling.
"In our home, and on our farm, there was much work for
even the smallest hands to do. Drinking water had to be carried
up a steep hill from a cold, fast-flowing spring 60 or 70 yards
away; and in summer, to and from the milk-house at this spring
all the milk, some fruits, and vegetables were carried. Each
winter and spring some acres had to be cleared of forest; later
a variety of crops had to be planted and this rough and stub-
born terrain had to be cultivated and harvested.
"In order to obtain some money, it was necessary for the sons
of our family to obtain work on nearby farms or in stores. Thus
during all of my ninth and tenth years, except for the short term
of school, I supported myself by work on a farm two miles from
my home."
Oscar Riddle's first school, a one-room cabin, was a mile
from the Riddle farm by way of a narrow path through woods
. — . . . . .
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OSCAR RIDDLE
429
and across fields. Like other boys of the neighborhood, Oscar
walked barefoot, even in frosty weather, wearing boots only
when snow lay on the ground. The school term was brief, about
seventy days in each year. After two years at the country school,
Oscar attended school in the village, with somewhat longer
terms, as much as one hundred days. To attend school and such
events in the village as spelling bees, debates, and church sup-
pers, the Riddle children walked two miles each way.
When twelve years old, Oscar helped in a store and delivered
newspapers; at thirteen he trapped forbearing animals in
wintertime; and for two years he swept the schoolroom floor and
built the fire, for ten cents a day. From his fourteenth year he
not only supported himself year-round, but like his older
brothers was able to turn over a little money to his mother.
Through hunting and trapping Oscar developed his lifelong
interest in the habits of birds and mammals. As early as the age
of eight his curiosity had been awakened by fossil shells and
imprints he had noticed in the banks and gullies around his
hillside home. These shapes in sandstone and limestone, he was
told, represented animals of kinds that lived only in the sea.
"This seemed to indicate, and led me to suspect, that our
earth must be very old. Yet all the preachers I had heard in-
sisted, and cited the Biblical record in support, that the earth
was created about 6,000 years ago, and that there had been one—
and only one—big and short-lived flood. How could this flood
have brought animals to our high hill from a sea that is almost
a thousand miles away? Even more disconcerting to me were
the dicta of these preachers, again supported by a Heaven-born
Bible, that a hot Hell exists, and that after death all unbelievers
go there and burn everlastingly. And I had to regard myself
as such an unbeliever!"
This conflict between dogma and observed fact caused the
boy great distress of a kind not uncommon in those days in
youngsters whose inquiring minds were breaking away from the
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430
BIOGRAPHICAL MEMOIRS
rigid beliefs of their elders. From the age of ten until he was
thirteen, Oscar tells us, the threat of hellfire often wrung a
prayer from him and brought frightened tears to his pillow
before he slept at night.
These fears were suddenly brushed away one night when the
boy attended a lecture at the village church, the very place
where he had so often heard the threat of damnation. A college
mate of his elder brother, named Francis Price, was studying
zoology at Indiana University under a twenty-seven-year-old
professor, Carl H. Eigenmann, who later became a member of
the National Academy of Sciences. Price had arranged to give
a talk at the church on the evolution of living things and to
illustrate it borrowed from Eigenmann a collection of fishes
preserved in alcohol, chosen to illustrate the principles of adap-
tation and natural selection. Either Price was very bold for the
time or the current pastor was more liberal than those Oscar
had heard earlier. At any rate the lad was so thrilled by the talk
that he had Price invited to the Riddle house for the night.
Thus enabled to examine the wonderful specimens for him-
self, with Price's kindly guidance, he understood the relics of
ancient life in the hillside strata that had worked so powerfully
upon his youthful mind. "I never prayed or wept upon my
pillow again," he wrote in old age. "Nothing in a long life has
equaled the release, thrill, and resolution obtained from this
message, so simply delivered by a young man from a neigh-
boring farm."
After completing grade school in the village of Cincinnati,
Oscar Riddle attended high school in Bloomfield, the county
seat of Greene County, and entered Indiana University in the
spring of 1896. He began at once the formal study of biology
and spent two summers at the university's biological field
station, then at Turkey Lake, Indiana. In the summer of 1899
his good work on a survey of Winona Lake led Professor
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OSCAR RIDDLE
431
Eigenmann to recommend him to the U.S. Commissioner of
Fisheries for assignment to collect tide-pool and freshwater fishes
of Puerto Rico, which had just become a possession of the
United States. Taking a hasty course in Spanish, Riddle inter-
rupted his college work and left for Puerto Rico in the autumn
of 1899. The island's Commissioner of Education promptly
asked him to teach biology to students of pharmacy and of edu-
cation in the newly established Model and Training School at
San Juan. Early in 1900 he also took over a beginning class in
chemistry.
That summer he was one of five men chosen to conduct
teachers' institutes in the ten largest cities of Puerto Rico. By
this time he was able to lecture in Spanish. Traveling by rail-
way, horse-drawn carriage, ox cart, and steamer, he covered much
of the island and the neighboring smaller isle of Vieques.
During a second year at San Juan Riddle taught classes in
biology in the high school, some of them in Spanish, and
followed up his course in chemistry for pharmacy students by
teaching them zoology and physiology. Several of his class of
fourteen, he learned years later, became physicians, one a
lawyer, one a banker, another a legislator, and one a professor
of Spanish in the new University of Puerto Rico. All this
teaching had left but little time for zoological collecting, but
in 1901 Riddle, at his own expense, made a summer's scientific
expedition to the delta of the Orinoco River, south of
Trinidad.
Returning home in the autumn of 1901, he registered at
Indiana University for the final year required for his bachelor's
degree. In January and February 1902 he accompanied Carl
Eigenmann on a six-week trip to collect blindfishes (a special
interest of Eigenmann's) in the caves and underground streams
of western Cuba. During that year also he prepared an article
on the fishes he had himself collected in Venezuela and Trini-
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432
BIOGRAPHICAL MEMOIRS
dad, but the manuscript that would have yielded his first
publication in zoology was stolen from him. He sold his collec-
tion to the Field Museum in Chicago.
After graduation from college Riddle declined a generous
offer from a family friend of a mercantile position in Indi-
anapolis. He also declined a teaching post at the University of
the Philippines and instead enrolled as a graduate student in the
University of Chicago. There he was under the general leader-
ship of Charles O. Whitman, but also followed (as he had
hoped) the lectures of Jacques Loeb. His plan was to prepare
himself for teaching and research, aiming for a career on the
preclinical side of a medical school. In his first term he took
Loeb's radically planned course in physiology, or rather general
physiology as we would term it today. Although Riddle does
not say so, it is obvious in retrospect that Loeb's departure that
winter for the University of California was an intellectual loss
to the young man, who could have benefited much if he had
gone on to research under Loeb, from the latter's rigorous
analytical thinking, of a kind that the still largely descriptive
methods of zoology did not demand.
Riddle's postgraduate training was interrupted by his ap-
pointment in the spring of 1903 to teach physiology in Central
High School of St. Louis, Missouri. Loeb's department had
been asked to recommend a man capable of introducing ]abora-
tory work into their didactic course. The project interested
Riddle, who moreover needed money to help a sister go to
college in St. Louis. He spent altogether five half-year periods
there (1903-1906) interspersed with other activities, including
participation in the summer course in physiology at Woods
Hole in 1903, a summer assistantship in zoology and biology at
Indiana in 1904, and a similar post at Indiana for eight months
in 1905 while on leave of absence from St. Louis. At St. Louis
he was also principal of one of the city's evening schools and
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OSCAR RIDDLE
433
filled in what was left of his working hours by studying French
and German at the local Berlitz school.
In February 1906, Riddle resigned his St. Louis post and
returned to Chicago to complete his postgraduate studies while
resuming his assistantship in zoology. He found Loeb's suc-
cessor in the chair of physiology, G. N. Stewart, less sympathetic
to the kind of training he wished to obtain than was Professor
Whitman and therefore decided to make zoology, under Whit-
man's tutelage, his major subject for the doctorate. Even
though he had accumulated sufficient credits for a minor in
physiology, with Whitman's approval he chose biochemistry
under Albert P. Matthews as his designated minor subject.
Whitman put him to work for his doctoral dissertation on a
problem of considerable theoretical importance, the cause of
the alternation of light and dark bars seen on the feathers of
many kinds of birds, notably fob and pigeons. Whitman's own
long studies of the evolution of birds, and especially of their
color patterns, had brought him face to face with this question,
which, as he perceived, called for both genetic and biochemical
studies. Thus was the course of Riddle's career as an investi-
gator set by the time he took his Ph.D. in zoology, in June 1907.
The guidance and companionship of Whitman, he says in his
autobiographical statement, provided one of the most profitable
and delightful epochs of his life: "Whitman became nearer to
being a father to me than anyone I have known."
After taking his doctorate, Riddle remained at the Univer-
s~ty of Chicago as an associate (a rank between assistant and
instructor) in zoology and embryology and also as an assistant in
experimental therapeutics (a research post). The next year he
was promoted to instructor in zoology and embryology, and in
the following two years he twice gave the course in embryology
for medical students and organized new courses in vertebrate
zoology and genera] biology and a graduate course in the
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434
BIOGRAPHICAL MEMOIRS
physiology of development, a quite novel topic. From his
laboratory he published several papers on color formation in
feathers, the development of yolk in hens' eggs, and the rate of
digestion in cold-blooded animals. In July 1910, he obtained
leave of absence for a year of travel and study in Europe.
Whitman had assured him that upon his return he would be
made assistant professor of biology and given charge of two of
the three terms of the introductory course in zoology.
Riddle began serious work abroad by settling for a few weeks
In Berlin, where in the university library he wrote a paper on
melanin formation in feathers, which he presented at the Eighth
International Zoological Congress, at Graz. After the Congress
he visited various European countries as a tourist. In Frank-
furt he called on Paul Ehrlich, who advised him about intra-
vitam stains for studying oxidation and reduction in animal
tissues, a topic he intended to investigate in the autumn at the
Naples Zoological Station.
Riddle had not been long at Naples when he received the
distressing news of Whitman's death on December 6, 1910. It
can do no harm now to the memory of the distinguished per-
sonages upon whom Riddle's career depended at that critical
time to say that Whitman's death was very unfortunate for him.
Frank R. Lillie of the Chicago department of zoology, who had
regarded himself as Whitman's heir apparent and in fact suc-
ceeded to the senior chair, was planning a radical redisposition
of the staff. Lillie wrote to Riddle in January 1911 that there
was internal opposition to him (as indeed there had been to
Whitman) and that he would not be reappointed. At about
the same time Whitman's friends in the university wrote of
their fears that the late professor's extensive unpublished
researches on the evolution of pigeons would never be published
under the new regime. Riddle, therefore, with self-sacrificing
loyalty to his late chief and mentor, left Naples and went home
to see what could be done to salvage Whitman's lifework. The
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OSCAR RIDDLE
435
struggle to take on this task, he records, and the labor of com-
pleting it were more formidable than any other efforts of his
lifetime.
Albert P. Matthews, Professor of Biochemistry, managed to
get him a six-month appointment on the payroll of the Labora-
tory of Experimental Therapeutics, a research unit of Matthews's
department. The Sprague Institute gave him $300 toward the
expenses of maintaining ~7hitman's large breeding colony of
pigeons, which was still kept at the late professor's home.
In 1912 came a great step forward in Riddle's career when
the Carnegie Institution of Washington made him a salaried
research associate, with funds to continue the pigeon colony,
and undertook to pay for publishing the Whitman papers when-
ever they might be ready for the press. Late in 1913 Riddle
moved, with the birds and the manuscripts, to the Carnegie
Institution's Station for Experimental Evolution, at Cold Spring
Harbor, Long Island. This appointment must have been initi-
ated by Charles B. Davenport, founder in 1904 and director of
the station. Yet Riddle states in his autobiographical notes that
he had a constant struggle to obtain adequate quarters for his
birds and efficient laboratory space for himself and indeed
received little encouragement for his research until, after many
years, Albert F. Blakeslee and later Milislav Demerec succeeded
to the directorship.
Davenport's coolness toward Riddle arose, no doubt, not
only from differences of temperament, but also from Riddle's
devotion to the memory of Whitman, whose scientific ideas as
revealed in the documents that his disciple was editing were
deeply at variance with those of Davenport. The research pro-
gram at Cold Spring Harbor was based on the Mendelian
principles that, since their rediscovery in 1900, had revolu-
tionized genetics. Whitman, on the other hand, had remained
unresponsive to much of the new genetics. He had begun to
study evolution in birds in 1892, at the age of fifty, under the
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436
BIOGRAPHICAL MEMOIRS
influence of an older school of biological thought. To him
recapitulation was the central fact of heredity; and he had,
moreover, chosen as the hereditable factors to be studied in his
hybrid birds three that did not lend themselves easily to
Mendelian analysis: color patterns, which are exceedingly com-
plex in birds, and sex determination and fertility, which are
complex phenomena in all animals. He had never accepted the
Mendelian ideas of unit characters and genetic dominance; he
doubted the importance of mutations for evolution and declared
that he had found evidence for evolution by orthogenesis. The
presence in Davenport's laboratory of an outspoken, enthusiastic
pupil of an anti-Mendelian must have irked the sensitive spirit
of its director.
At any rate, Riddle, while organizing, against what he felt
to be his chief's indifference, a laboratory that never quite
matched his own standards and getting under way a broad
program of research, toiled on and on with \\lhitman's volumi-
nous and, to a large extent, ill-sorted papers. The task was
varied and immense, requiring rearrangement and assemblage
of misplaced portions of chapters, analysis of numerous tables,
and placement of numerous illustrations. In this task also he
did not get all the help he needed, for Mrs. Whitman had for
reasons of her own at times limited his use of the materials.
At last, in 1914, the Carnegie Institution published the Whit-
man papers in three large and handsomely illustrated volumes.
The first two, edited solely by Riddle, present a clear statement
of Whitman's studies on natural and hybrid pigeons and doves,
their growth, and particularly their inheritance of feather
patterns. In the third volume Riddle gathered together Whit-
man's intensive observations of sex behavior and reproductive
activities. Feeling himself not competent to assess this material,
he turned the detailed editing over to Harvey A. Carr, Associate
Professor of Psychology at the University of Chicago. This third
volume, largely free of the conjectural and controversial bias of
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456
BIOGRAPHICAL MEMOIRS
1929
Some interrelations of sexuality, reproduction and internal secre-
tions. I. Am. Med. Assoc., 92:943-50.
The special contribution of developmental mechanics to the thought
and purpose of the man of tomorrow. Proc. Am. Phil. Soc.,
68: 107-17.
With F. G. Benedict. The measurement of the basal heat produc-
tion of pigeons. Part I. Instrumental technique J. Nutrition,
1 :475-95.
With F. G. Benedict. The measurement of the basal heat produc-
tion of pigeons. Part II. Physiological technique. J. Nutrition,
1 :497-536.
Endocrine regulation of reproduction. Endocrinology, 13: 311-19.
The inheritance of thyroid size and the establishment of thyroid
races in ring-do~res. Am. Naturalist, 63: 385~09.
1930
Complete atrophy of kidney in pigeons following section of the
ureter. Proc. Soc. Exp. Biol. Med., 27:1022-24.
The age distribution of mortality in bird embryos and its probable
significance. Am. l. Physiol., 94:535-37.
With G. Christman and F. G. Benedict. Differential response of
male and female ring-doves to metabolism measurements at
higher and lower temperatures.
1931
Am. J. Physiol., 95:111-21.
Factors in the development of sex and secondary sexual character-
istics. Physiological Reviews, 11 :63-106.
With J. Krizenecky. Extirpation of thymus and bursa in pigeons
with a consideration of the failure of thymectomy to reveal
thymus function. Am. J. Physiol., 97:343-52.
Season of origin as a determiner of age at which birds become sex-
ually mature. Am. i. Physiol., 97:581-87.
With P. F. Braucher. Control of the special secretion of the crop-
gland in pigeons by an anterior pituitary hormone. Am. J.
Physiol., 97:617-25.
With I. Polhemus. Effects of anterior pituitary hormones on gonads
OCR for page 479
O S C A R R I D D L E
457
and other organ weights in the pigeon. Am. I. Physiol., 98:
121-30.
Studies on pituitary functions. Endocrinology, 15:307~14.
lg32
With R. W. Bates and S. W. Dykshorn. A new hormone of the
anterior pituitary. Proc. Soc. Exp. Biol. Med., 29:1211-12.
With S. W. Dykshorn. Secretion of crop-milk in the castrate male
pigeon. Proc. Soc. Exp. Biol. Med., 29:1213-15.
With D. R. Charles and G. E. Cauthen. Relative growth rates in
large and small races of pigeons. Proc. Soc. Exp. Biol. Med.,
29: 1216-20.
With G. C. Smith and F. G. Benedict. Seasonal, endocrine and
temperature factors which determine percentage metabolism
change per degree of temperature change. Am. l. Physiol.,
101:88. (A)
With T. C. Nussmann and F. G. Benedict. Metabolism during
growth in a common pigeon. Am. J. Physiol., 101:251-69.
With G. C. Smith and F. G. Benedict. The basal metabolism of
the mourning dove and some of its hybrids. Am. I. Physiol.,
101 :260-67.
Metabolism and sex. In: Sex and Internal Secretions, ed. by Edgar
Allen, pp. 246-80. Baltimore, The Williams & Wilkins Com-
pany.
With R. W. Bates and S. W. Dykshorn.
Prolactin, a new and third
hormone of the anterior pituitary. Anat. Record, 54:25. (A)
Sex and intersex in pigeons. In: Proceedings of the Sixth Inter-
national Congress of Genetics, ed. by Donald F. tones, Vol. 2,
pp. 165-68. Ithaca, New York. Brooklyn, Brooklyn Botanic
Garden. (A)
1933
With R. W. Bates and S. W. Dykshorn. Thyroid hypertrophy as a
response to the gonad-stimulating hormone of the pituitary.
Proc. Soc. Exp. Biol. Med., 30:794-97.
With R. W. Bates and S. W. Dykshorn. The preparation, identifica-
tion and assay of prolactin a hormone of the anterior pituitary.
Am. J. Physiol., 105: 191-216.
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458
With R. W. Bates.
BIOGRAPHICAL MEMOIRS
Concerning anterior pituitary hormones. En-
docrinology, 17: 689-98.
With G. C. Smith and F. G. Benedict. Basal metabolism and the
temperature factor in brooding ring-doves.
105:428-33.
With T. C. Nussmann.
Am. J. Physiol.,
A sex difference in pituitary size and in-
Anat. Record, 57: 197-204.
testinal length in doves and pigeons.
1934
With G. O. Smith and F. G. Benedict. Seasonal and temperature
factors and their determination in pigeons of percentage metab-
olism change per degree of temperature change. Am. J. Physiol.,
107:333-43.
\Vith P. F. Braucher. Body size changes in doves and pigeons
incident to stages of the reproductive cycle. Am. i. Physiol.,
107:343-47.
With P. F. Braucher.
Hemoglobin and erythrocyte difference ac-
cording to sex and season in doves and pigeons. Am. I. Physiol.,
108:554-66.
With R. W. Bates and E. L. Lahr. On the protein nature of pro-
lactin and of follicle-stimulating hormones. Proc. Soc. Exp.
Biol. Med., 31:1223-24.
With L. B. Dotti. Action of parathyroid hormone in normal and
hypophysectomized pigeons.
507-9.
Proc. Soc. Exp. Biol. Med., 32:
With E. L. Lahr, 11. W. Bates and C. S. Moran. Response of adult
rat testes, sex accessories and adrenals to injections of prolactin.
Proc. Soc. Exp. Biol. Med., 32:509-12.
1935
Contemplating the hormones. Endocrinology, 19:1-13.
With R. W. Bates and E. L. Lahr. Prolactin induces broodiness in
fowl. Am. J. Physiol., 111:352-60.
With E. L. Lahr and R. W. Bates. The gross action of prolactin
and follicle-stimulating hormone on mature ovary and sex
accessories of fowl. Am. l. Physiol., 111:361-68.
With E. L. Lahr and R. W. Bates. Maternal behavior induced in
virgin rats by prolactin. Proc. Soc. Exp. Biol. Med., 32:730-34.
With E. L. Lahr and R. W. Bates. Demonstration of prolactin
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OSCAR RIDDLE
45Q
induced activities which express maternal behavior in virgin
rats. Am. I. Physiol., 113:110.
With l. P. Schooley. Absence of follicle-stimulating hormone in
pituitaries of young pigeons. Proc. Soc. Exp. Biol. Med., 32:
1610-14.
With G. C. Smith and C. S. Moran. Effects of complete and incom-
plete hypophysectomy on the basal metabolism of pigeons. Proc.
Soc. Exp. Biol. Med., 32:1614-16.
Aspects and implications of the hormonal control of the maternal
instincts. Proc. Am. Phil. Soc., 75:521-25.
With R. W. Bates and E. L. Lahr. An assay of three hormones
present in the anterior pituitaries of seven types of cattle classi-
fied for age, sex, and stage of reproduction. Am. l. Physiol.,
113:259-64.
With G. C. Smith. The effect of temperature on the calorigenic
action of dinitrophenol in normal and thyroidectomized pigeons.
i. Pharmacol. Exp. Therap., 55:173-78.
With R. W. Bates. The preparation of prolactin. l. Pharmacol.
Exp. Therap., 5~:365-71.
.,
With R. W. Bates and T. Laanes. Evidence from dwarf mice
against the individuality of growth hormone. Proc. Soc. Exp.
Biol. Med., 33:446-50.
1936
With G. C. Smith, R. W. Bates, C. S. Moran and E. L. Lahr. Action
of anterior pituitary hormones on basal metabolism of normal
and hypophysectomized pigeons and on a paradoxical influence
of temperature. Endocrinology, 20: 1-16.
The confusion of tongues. Science, 83:41-4b, 69-74.
With E. L. Lahr and R. W. Bates. Histological changes induced in
the testes of immature doves and pigeons by gonadotropic hor-
mone. Am. l. Physiol., 116:94-95. (A)
With R. W. Bates.
of prolactin. Proc. Soc. Exp. Biol. Med., 34:847~9.
With E. L. Lahr. Temporary suppression of estrous cycles in the
rat by prolactin. Proc. Soc. Exp. Biol. Med., 34:880-83.
With L. B. Dotti. Blood calcium in relation to anterior pituitary
and sex hormones. Science, 84: 557-59.
Effect of route of administration on the bioassay
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BIOGRAPHICAL MEMOIRS
With l. P. Schooley. Reciprocal weight changes in liver and testis
of pigeons during reproduction. Anat. Record, 67:51. (A)
1937
With i. P. Schooley and R. W. Bates. Effective stimulation of crop-
sacs by prolactin in hypophysectomized and in adrenalectomized
pigeons. Proc. Soc. Exp. Biol. Med., 36:408-11.
The relative claims of natural science and of social studies to a
core place in the secondary school curriculum. Science Edu-
cation, 21: 65-71.
With R. W. Bates, E. L. Lahr and l. P. Schooley. Aspects of
splanchnomegaly associated with the action of prolactin. Am.
J. Physiol., 19:603-9.
With R. W. Bates and E. L. Lahr. The mechanism of the anti-
gonad action of prolactin in adult pigeons. Am. i. Physiol.,
19:610-14.
Physiological responses to prolactin. Cold Spring Harbor Symp.
Quant. Biol., 5:218-28.
Carbohydrate metabolism in pigeons. Cold Spring Harbor Symp.
Quant. Biol., 5:362-74.
1938
With J. P. Schooley. The morphological basis of pituitary function
in pigeons. Am. I. Anat., 62: 313~9.
Science, 87: 375-80.
With G. E. Cauthen. Erythrocyte number in young pigeons and
its relation to heredity, growth and metabolism. Am. l. Physiol.,
122:480-85.
Educational darkness and luminous research.
Prolactin, a product of the pituitary, and the part it plays in vital
processes. Sci. Monthly, 47: 97-113.
With E. L. Lahr. Proliferation of crop-sac epithelium in incubating
and in prolactin injected pigeons studied with the colchicine
technique. Am. J. Physiol., 123:614-19.
1939
With R. W. Bates and E. L. Lahr. The racial factor in the pigeon
crop-sac method of bioassay. Am. i. Physiol., 125:722-29.
The opportunity and obligation of the National Association of
Biology Teachers. Am. Biol. Teacher, 1:115-21.
With R. Miller. Stimulation of the adrenal cortex of pigeons by
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OSCAR RIDDLE
461
anterior pituitary hormones and by their secondary products.
Proc. Soc. Exp. Biol. NIed., 41:518-22.
With R. W. Bates. The preparation, assay and actions of the lacto-
genic hormone. Chapter XX in: Sex and Internal Secretions,
2d ea., ed. by Edgar Allen, pp. 1088-1 1 17. Baltimore, The
Williams 8c Wilkins Company.
With R. W. Bates and E. L. Lahr. The role of sex, estrogenic hor-
mone, fasting and diuresis in the response of crop-sacs of pigeons
to prolactin. Am. J. Physiol., 127:422-29.
With SKI. W. Johnson. An undescribed type of partial sex-reversal
in dove hybrids from a sub-family cross. Anat. Record, 75:509-
27.
With M. W. Johnson. Tests of mammalian gonad-stimulating
hormones on gonads of fishes. Proc. Soc. Exp. Biol. Med., 42:
260-62.
1940
Lactogenic and mammogenic hormones. I- Am. \ted. Assoc., 115:
2276-81.
1941
With E. L. Lahr and L. H. Elwell. Mitosis observed under col-
chicine in crop-sac tissue after subcutaneous and intramuscular
injection of prolactin. Archives of International Pharmaco-
dynamics and Therapeutics, 65:278-82.
With E. L. Lahr and R. W. Bates. The response of testes of im-
mature pigeons to gonadotrophins. Endocrinology, 28:681-93.
With J. P. Schooley and R. W. Bates. Replacement therapy in
hypophysectomized juvenile pigeons. Am. J. Anat., 69:123-54.
Endocrine aspects of the physiology of reproduction. Annual Re-
vie`N of Physiology, 3: 573-616.
Preliminary impressions and facts from a questionnaire on secondary
school biology. Am. Biol. Teacher, 3:151-59.
With R. W. Bates. Annual variation in the crop-sac response by
prolactin. I. Biol. Chem., 1 1 1: cxliii-iv. (A)
With R. A. Miller. Cellular response to insulin in suprarenals of
pigeons. Proc. Soc. Exp. Biol. Med., 47:449-53.
Recognition and removal of barriers to effective teaching of sec-
ondary school biology. Bulletin of the Department of Science
Instruction of the National Education Association, pp. 20-27.
7
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462
BIOGRAPHICAL MEMOIRS
With R. W. Bates and E. L. Lahr. A strain difference in responsive-
ness of chick thyroids to thyrotrophin and a step-wise increase
during three years in thyroid weights of Carneau pigeons. En-
docrinology, 29:492-97.
With R. W. Bates. Annual variation in the response of crop-sacs
and viscera of pigeons to prolactin. Endocrinology, 29:702-9.
1942
With H. H. Dunham. Effects of a series of sterols on ovulation and
reproduction in pigeons. Physiol. Zool., 15: 383-95.
With H. H. Dunham. Transformation of males to intersexes by
estrogen passed from blood of ring-doves to their ovarian eggs.
Endocrinology, 30:959-68.
With R. W. Bates, T. Laanes and E. C. MacDowell. Growth in
silver dwarf mice, with and without injections of anterior pitui-
tary extracts. Endocrinology, 31: 53-58.
With B. B. Wells and H. N. Marvin. The Bomskov reports on
thymus mediation of pituitary function. Proc. Soc. Exp. Biol.
Med., 49:473-76.
Amount and nature of biology teaching in secondary schools. Sec-
tion VI in: The Teaching of Biology in Secondary Schools of the
United States, ed. by Oscar Riddle, pp. 54-76. Lancaster, Penn-
sylvania, Science Press.
Cyclic changes in blood calcium and phosphorus in relation to egg
laying and estrogen production. Endocrinology, 31:498-506.
General relationship of hormones to growth and development. Cold
Spring Harbor Symp. Quant. Biol., 10:7-14.
With R. A. Miller. The cytology of the pigeon adrenal cortex in
experimentally induced atrophy and hyperactivity. Am. l.
Anat.,71:311-35.
The preparation of high school science teachers. Am. Biol.
Teacher, 5: 63-65.
Hormone therapy viewed by the research physiologist. In: Proceed-
ings of the American Pharmaceutical Manufacture's' Association
New York City, pp. 82-89.
1943
With E. L. Lahr and R. W. Bates. Non-specific results obtained
with the micromethod for assay of prolactin. Endocrinology,
32:251-59.
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OSCAR RIDDLE
Edith R. A. Miller.
463
Effects of prolactin and cortical hormones on
body weight and food intake of adrenalectomized pigeons. Proc.
Soc. Exp. Biol. Med., 52:231-33.
With R. A. Miller. Ability of adrenal cortical hormones, prolactin
and thyroxin to maintain weight of body and viscera of hypo-
physectomized pigeons. Endocrinology, 32:463-74.
With W. F. Hollander. The inheritance of "scraggly" plumage
and of ataxia in the pigeon. journal of Heredity, 34:167-72.
1944
With i. P. Schooley. Effect of light upon time of ovi-position in
ring-doves. Physiol. Zool., 16: 187-93.
Charles Benedict Davenport. Science, 99:441-42.
With G. C. Smith and R. A. Miller. The effect of adrenalectomy
on heat production in pigeons. Am. i. Physiol., 141:151-57.
With G. C. Smith. The effects of fasting on heat production in
normal and hypophysectomized young pigeons. Am. J. Physiol.,
141 :303-11.
With E. L. Lahr. On broodiness of ring-doves following implants
of certain steroid hormones. Endocrinology, 35:255-60.
With E. L. Lahr. The action of steroid hormones on the mature
dove testis. Endocrinology, 35:261-66.
With E. L. Lahr. Relative ability of various steroid hormones to
promote growth in the oviduct of immature ring-doves. Yale
journal of Biology and Medicine, 17:259-68.
With V. M. Rauch and G. C. Smith. Changes in medullary bone
during the reproductive cycle of female pigeons. Anat. Record,
90:295-305.
1945
With V. M. Rauch and G. C. Smith. Action of estrogen on plasma
calcium and endosteal bone formation in parathyroidectomized
pigeons. Endocrinology, 36:41-47.
With M. R. McDonald. The partition of plasma calcium and in-
organic phosphorus in estrogen-treated normal, parathyroidec-
tomized and hypophysectomized pigeons. Endocrinology, 36:
48-~.
With M. R. McDonald and G. C. Smith. Action of thyroxin on
estrogen-induced changes in blood chemistry and endosteal bone.
Endocrinology, 37: 23-28.
.
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464
BIOGRAPHICAL MEMOIRS
With W. F. Hollander. On partial melanism associated with para-
thyroid enlargement in pigeons. Am. Naturalist, 78:456-63.
With M. R. McDonald. The effect of reproduction and estrogen
administration on the partition of calcium, phosphorus and
nitrogen in pigeon plasma. i. Biol. Chem., 159:445-64.
With W. F. Hollander and I. P. Schooley. A race of hermaphrodite-
producing pigeons. Anat. Record, 92:401-23.
With E. L. Lahr. Intersexuality in male embryos of pigeons. Anat.
Record, 92 :425-31.
With W. F. Hollander. Goiter in domestic pigeons. Poultry
Science, 25:20-27.
With L. B. Dotti. Pituitary ant! sex hormones which increase
plasma calcium in birds and mammals. Proc. Am. Phil. Soc.,
89:499-516.
1947
With seven associates. Studies on Carbohydrate and Fat Metabolism
with Especial Reference to the Pigeon. Washington, D.C., Car-
negie Institution of Washington. Publication No. 569. iv +
128 pp.
Endocrines and Constitution in Doves and Pigeons. Washington,
D.C., Carnegie Institution of Washington. Publication No. 572.
xi + 306 pp.
1948
Charles Benedict Davenport (1866-1944~. In: National Academy
of Sciences, Biographical Memoirs, 25:75-110. Washington, D.C.,
National Academy of Sciences.
1954
High schools and biological literacy in the United States. Am.
Biol. Teacher, 16: 179-84.
The Unleashing of Evolutionary Thought.
Press, Inc. xxi + 414 pp.
1958
New York, Vantage
The new national interest in high school science. Am. Biol.
Teacher, 20: 1 ~ 1-53.
With Eugene F. Dubois.
Francis Gano Benedict (1870-1957) . In:
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OSCAR RIDDLE
465
National Academy of Sciences, Biographical Memoirs, 32:67-99.
New York, Columbia University Press.
Is organized religion the source of today's moral impasse? Unity
144: 37-40.
1959
The origin of good and evil. Unity, 145:5-12.
Must we fail in science education? Physiologist, 2:55-57.
Must the humanities perpetuate supernaturalism? Realist, 2
1960
:1-7.
The spreading spark of life. Am. Biol. Teacher, 22:228-32.
Editorial. A real religious issue in the campaign. American
Rationalist, 5:3-4.
1961
The nature of sex. In: Encyclopedia of Sexual Behavior, ed. by A.
Ellis and A. Abarbanel, pp. 757-68. New York, Hawthorn
Books, Inc.
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Representative terms from entire chapter:
oscar riddle
