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OCR for page 413
CARROLL MILTON WILLIAMS
December 2, ~ 91 6-October ~ I, ~ 99
BY A. M. PAPPENHEIMER, JR.
CARROLL MILTON WILLIAMS was born in Richmond, Virginia,
on December 2, 1916. Even in his early school days he
snowed great interest in science and soon after entering
the University of Richmond at the age of sixteen began
collecting and studying lepicloptera. Upon graduation, he
gave his outstanding collection to the university.
Carroll published his first paper on butterflies in 1937,
when he was twenty, just before graduating from college. In
the fall of that year he became a graduate student at Harvard
University, where he was to remain for the rest of his life.
His thesis adviser was Professor Charles Brues, a well-known
entomologist. Carroll's remarkable and brilliant thesis was
titles] "A Morphological en cl Physiological Analysis of the
Flight of Drosophila, with Special Reference to Factors Con-
trolling Wing Beat" and was written in what was to become
Williams's characteristic and unique style with its humor-
ous overtones. In collaboration with Leigh Chadwick and
with advice from Professor Ecigerton of MTT, Carroll de-
signecl a small apparatus that measured accurately and re-
proclucibly, by a stroboscopic method, the wing beat fre-
quency of both wil(l-type and mutant strains of tiny fruit
flies in flight under a wide variety of conditions, such as
413
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414
BIOGRAPHICAL MEMOIRS
temperature, atmospheric pressure, O2 tension, etc. Tndi-
vidual flies from various inbred strains varier! between 12,000
and 14,000 beats per minute of sustained flight until ex-
haustion set in after as much as three hours, or more than
2 million double wing beats. He measured the glycogen
content of the thoraces of the blowfly (a slightly larger in-
sect) cluring flight to exhaustion ant! determinect the en-
ergy expendecl in terms of glucose consumption. Finally,
he succeeded in demonstrating, where others had failed,
the neuromuscular network in the thorax that controls the
wing beat.
In 1941, after receiving his Ph. D., Carroll was appointed
a junior fellow of the Harvard Society of Fellows. It was
clear from his thesis that he needled a larger experimental
animal than Drosophila to pursue the studies he contem-
plated on insect clevelopment and morphogenesis. He there-
fore selected the giant silkworm, Hyalophra (formerly
Platysamia) cecropia, as his experimental animal and soon
made the important ant] useful observation that insects can
be anesthetized for long periods of time, uncier continuous
flow of carbon dioxicle in a Buchner funnel, thus permit-
ting surgical manipulations without Toss of blood or dam-
age. While still a junior fellow, he decicled to obtain a medi-
cal degree anct in 1946 received his M.D. summa cum laude
from the Harvard Medical School.
The years that follower! were exciting and fruitful ones.
Carroll was appointed assistant professor of biology in 1946,
promoted to associate professor two years later, and be-
came professor of zoology in 1953 at the age of thirty-six.
Finally, in ~ 965, he was appointed the first Bussy Professor
of Biology. As one of his graduate students during the ear-
lier period wrote:
When I think of Carroll's achievements, I am overwhelmed by memories of
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CARROLL MILTON WILLIAMS
415
hilarious events and merry times. And I am sure this was one of the reasons
for Carroll's success in attracting students and bringing out the best in
them: life in his lab was usually such fun and we all shared so many laughs.
Cecropia moths lay their eggs in early summer. After hatch-
ing, the tiny caterpillars grow rapidly and after four molts
attain a length of more than three inches. They then spin a
cocoon inside of which they metamorphose and enter a
prolonged period of pupal diapause over the winter. If pu-
pae that have entered diapause are placed at 3° to 5°C for a
few weeks, adult development may be initiated promptly by
removing them to warm temperatures. Without this period
of chilling, adult development will not begin for many
months, if indeed at all. Carroll began his studies on adult
development by placing (liapausing pupae in (lifferent ori-
entations under temperature gradients with one end kept
at 3° to 5°C and the other at 25° to 30°C. He observed that
although development began in the chilled anter~orend, once
started, the heated end developed faster. It was these initial
observations that led Carroll to publish a long series of
remarkable and highly original papers in the Biological Bul-
letin on the physiology, biochemistry, and hormonal con-
trol of insect diapause and adult clevelooment. Many of
~ ,
these and his subsequent papers and lectures were illus-
trated by the excellent photographs and slides macle by his
wife Muriel.
Carroll's experiments were often amusing as well as inge-
nious and revealing. He began with parabiotic experiments
in which he joined together by their heads diapausing pu-
pae ant} chilled diapausing pupae. Almost simultaneously,
both began to develop into aclult moths. He soon found
that removal of the brain from a pupa leads to permanent
diapause but that adult development took place promptly if
the brain from a chilled pupa was clroppe(1 into a brainless
diapausing pupa at 25°C, even if the latter was of a differ-
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416
rat ~ ~ ~ ~ ~ ~ ~ .
BIOGRAPHICAL MEMOIRS
ent species, such as Antheraea (formerly Telea) polyphemus or
Samia cynthia. By means of plastic windows place(1 in either
the face or the tip of the abdomen, development couIc3 be
to~owec~ clay ny clay trom its onset until emergence of the
adult moth twenty-one clays later. Although it was evident
that the chilled brain secretes! a hormone necessary for ini-
tiation of aclult development, it was soon shown that this
was not a sufficient condition. When brainless pupae were
cut in half and chiller! brains were ciroppect into each half-
pupa, only the anterior half went on to clevelop into half an
aclult moth! However, if both a chilled brain and a bit of
prothoracic "glancl" tissue, dissectecl from a normal pupal
Borax, were aroppea Into tne posterior encl, an aclult ab-
domen clevelopec! (see Figure By. Further work showed that
a tropic hormone was synthesized by a set of eleven neuro-
~ 1_ _ ~ ~ · . . ~ . · ~
secretory cells in the anterior part of the chillecl brain that
activates! the prothoracic glands to produce a growth ant!
development factor. In ID54 Peter KarIson, working in
Butenant's laboratory in Germany, isolated 25 mg of the
crystalline growth factor from 500 kg of Bombyx more pupae.
He named it ec~ysone and determined its steroic! structure.
It was obvious that each morphological change from larva
to diapausing pupa ant! finally to an aclult moth must be
accompanied by (dramatic changes in metabolism. During
the next few years these changes were studiecl by Carroll
and his students- R. C. Sanborn, H. A. Schnei(lerman, D.
G. Shappirio, and W. R. Harvey. It came as no surprise to
find that oxygen consumption dropped precipitously upon
entering diapause en c! rose again cluring aclult development.
Nor was the fact that, upon entering diapause, most com-
portents of the cytochrome system were broken clown an(l,
with the exception of the intersegmental muscles of the
pupal abdomen, tissue respiration, including that of the
heart, which continued to beat slowly, became insensitive
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CARROLL MILTON WILLIAMS
A, ~ Act: ~ ~~ : ~
:: ~
417
· .. .
FIGURE 1 Upper: Brain and prothoracic glands obtained from previously
chilled pupae being implanted into an isolated pupal abdomen. Lower:
Implanted endocrine organs have caused adult development of the abdo-
men, which is shown laying eggs.
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418
BIOGRAPHICAL MEMOIRS
to inhibition by cyanide or carbon monoxide. These stucI-
ies led to the discovery of a new cytochrome X (later re-
namecI cytochrome b5) present in the caterpillar midgut
and in the pupal heart during ~liapause.
While these metabolic studies were in progress, Carroll
became interested in juvenile hormones, which V. B.
Wigglesworth showed many years before to be secreted by
the corpora allata, two small glands connected to the brain
of the bug Rhodnius by a pair of tiny nerves. This hormone
opposes metamorphosis. Carroll macle the surprising obser-
vation that excision of the corpora allata from chilled cecropia
pupae hac] no effect on development into normal fertile
aclult moths, although when removed from adults and tested,
they prover! to be more active than at any stage of its life
history. However, during the very early stages of aclult de-
velopment, addition of the hormone caused transforma-
tion into a second pupa with mere traces of adult charac-
teristics. Finally, the most surprising finding was that the
highest concentrations of all were present in the abdomens
of aclult male cecropia moths. Carroll found that the hor-
mone could be extracted from homogenates of mate abcto-
mens by petroleum ether, which yieldect a potent water-
insoluble of! upon evaporation. Even after 50,000-fold
purification, the active component still contained impuri-
ties, but the purest preparations tract the properties of a
terpenoict acid, and certain synthetic derivatives of farnesoic
acid tract potent juvenile hormone activity.
In 1964 Dr. Karel STama came from Czechoslovakia to
work in Carroll's laboratory, bringing with him fertile eggs
from the bug Pyrrhocoris that he had been rearing in Petri
clishes without difficulty in his Prague laboratory for ten
years. In the Harvard laboratory, however, all larvae contin-
ue(1 to experience adclitional larval molts and cliecl without
becoming adults. The clifficulty was finally traced to Scott
OCR for page 419
CARROLL MILTON WILLIAMS
419
paper toweling that had been placed in the rearing jars.
When replaced by Whatman filter paper, all larvae devel-
oped normally into adult bugs. The following is a quotation
from a paper by Slama and Williams published in the Pro-
ceedings of the National Academy of Sciences in 1965:
Indeed, pieces of American newspapers and journals (New York Tames, Wall
Street fournal, Boston Globe, Science, and Scientific American) showed extremely
high juvenile hormone activity when placed in contact with Pyrrhocoras lar-
vae. The London Times and Nature were inactive.
The active factor could be extracted from Scott paper
towels with organic solvents and was found to be a petro-
leum ether-soluble oil that was highly active as a juvenile
hormone when tested on Pyrrhocoris but that had no effect
on metamorphosis of Cecropia. The factor could easily be
extracted from American balsa fir, but only traces were present
in European paper pulp. Even in his first paper on juvenile
hormone, which appeared! in 1956 as a letter to Nature,
Carroll realized its potentiality as a pesticide and wrote as
follows:
In addition to the theoretical interest of the juvenile hormone, it seems
likely that the hormone, when identified and synthesized, will prove to be
an effective insecticide. This prospect is worthy of attention because insects
can scarcely evolve a resistance to their own hormone.
By the mid-1970s this prediction had been verified. Three
closely related juvenile hormones had been synthesized, and
several substances that cleraiT clevelopment of certain insect
species by turning off secretion of juvenile hormone by the
corpora allata hacl been isolated from plants in other labora-
tories. The chemical industry was engaged in synthesizing
hundreds of cheaper and more stable analogs of these com-
pounds. in a number of cases, these analogs have shown
highly specific activity for certain insect species. For example,
the juvenile hormone analog Methoprene is now in use in
.
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420
BIOGRAPHICAL MEMOIRS
controlling the floodwater mosquito Cedes nigromaculus, a
species that hac! become resistant to conventional pesticides.
Less than 5 g spread ner acre gives Hooch control.
About this time, Carroll found that he was unable to
mate polyphemus moths in the laboratory. The larva of this
moth feed on oak. With Lynn Ricicliford, he soon found
that, after acicling a few oak leaves to the cages, the females
secreted a pheromone that attracted mates and stimulatecl
.
. . .
mating. (! cannot resist quoting here from a short note
published in Science by Rictdiford ant! Williams: "The action
of oak factor on the female can be masker! by other volatile
agents including Chanel No. 5."
In 1970 or 1971 Carroll abanclonect his favorite experi-
mental animal, cecropia, and switched to Manduca sepia (the
tobacco hornworm). The great advantage of Manduca is
that it can be raisecl, even during the winter, on a simple
artificial diet, making possible the study of the larval stages
all year round.
Manduca larvae cluring their final fifth instar increase
their weight within four to five days from about ~ g to as
much as 10 g, after which they stop fee(ling and purge their
gut, reducing their weight to 5 g. After the purge the larvae
enter a "wandering stage" that soon ends in an abrupt on-
set of negative phototaxis. In the wiTcI, larvae wouIc! then
lig into the grouncl before pupation. The experiments at
the Harvard laboratory, however, were carried out on a tweIve-
hour light/ciark cycle. Most of the experiments on hormonal
control of Manduca development were carried out by
postdoctoral fellows ant! students cluring the decade before
Carroll's retirement to emeritus status. Manduca has now
become one of the most important mocle! systems for the
stucly of insect physiology, development, neurobiology, en cl
molecular biology.
During the last few years before his final illness, Carroll
OCR for page 421
CARROLL MILTON WILLIAMS
42
clid very little at the bench himself. Nevertheless, he kept
up with current literature and realizecI that many of the
questions raised by his experiments on insect development
ant! the regulation of insect hormone expression could be
answered by the techniques of present-clay molecular biol-
ogy. After departmental colloquia he often rose to ask visit-
ing lecturers important en c! penetrating questions relating
to the biological significance of their molecular finclings,
even though the subject might be quite remote from his
own fielcI.
Carroll enjoyed teaching and was not only an entertain-
ing and popular lecturer but also stimulates! many students
to become interested! permanently in biology and often to
seek to clo graduate study uncler his guidance. But he never
clirectect a large team of graduate students and postdoctoral
fellows as is so often the case in molecular biology today.
Almost all} of his students came to his laboratory because
they fell under his spell while listening to his lectures or
because they were fascinated by his publishecI experimental
work. Carroll's five o'clock laboratory teas were legenciary
and were attenclecl by everyone, from undergracluates to
visiting professors. The following are direct quotations from
letters written to me by three of his former graduate stu-
dents, now tenured professors of biology at the State Uni-
versity of New York at Stonybrook, the University of Wash-
ington, ant! the Universitv of Michigan respectively:
~ , ~
Carroll was also exceptional—certainly by the standards of today in his
willingness to point students to problems that were quite remote from the
work he did himself. He did not hesitate to launch students on projects
that required techniques he had never used and which were founded on
principles about which he had little knowledge. Carroll would learn along
with the student and seemed always to contribute the needed experimental
trick or flash of insight.
OCR for page 422
422
BIOGRAPHICAL MEMOIRS
Carroll was always full of ideas and tried to instill into his graduate stu-
dents and postdoctoral associates the importance of doing experiments to
test ideas particularly those that seemed far-fetched. pie had little patience
with students who would find various theoretical reasons why something
might not work and would not go to the lab to test an idea. He also,
though, was a hard task master in ensuring that experimental results were
repeatable and that further experiments necessary to explain the results
were done before they were published. Hence, many of his papers talk
about work done over years.
.
In the years I was a graduate student, we had tea each afternoon. Under-
graduates, graduate students, postdoctoral fellows and visitors regularly at-
tended. Of all the insect hormones then known (and perhaps now known),
juvenile hormone was the most mysterious and fascinating. At the tea-table
I heard what I think were his first statements about using it and perhaps
other insect hormones as insecticides of the future or third generation
. .
pesticides.
it will come as no surprise to learn that Carroll was in
much clemanct as a gifted lecturer. He was invite ct to deliver
more than forty named lectures, among which, to mention
only a few, were the Lowell lectures in Boston (19481; the
Harvey Lecture in New York (19521; the AAAS Holiday Lec-
ture, University of Chicago (19701; and the CSTRO Lec-
tures in Australia ~ ~ 973) .
Carroll was elected a fellow of the American Acaclemy of
Arts and Sciences in 1951 and server! on its council from
1952 to 1955 ant! again from 1974 to 1977. He was elected
to the National Academy of Sciences in 1~960 and was a
member of its council from 1973 to 1976 and again from
1985 to 1988. He was chairman of the Section on Biological
Sciences from 1981 to 1984. He also became a member of
the Philosophical Society in 1969 ant! was a member of
numerous other Earned societies, inclucling the Pontifical
Academy of Rome.
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CARROLL MILTON WILLIAMS
423
~ AM GREATLY INDEBTED to Professors Lynn M. Riddiford, William G.
Van der Kloot, and David G. Shappirio for sending me their remi-
niscences of Carroll and to Daniel Branton and Fotis Kafatos for
critical reading of the manuscript.
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424
BIOGRAPHICAL MEMOIRS
HONORS AND DISTINCTIONS
Borden Research Award, Harvard Medical School, 1946
AAAS-Newcomb Cleveland Prize, 1950
Guggenheim Fellowship (Cambridge University), 1955-56
Founders Memorial Award, Entomological Society of America,
1958
Boylston Medalist, Harvard Medical School, 1961
Trustee of Radcliffe College, 1961-64
George Leslie Award, Harvard University, 1967
Howard Taylor Ricketts Award, University of Chicago, 1969
Chief scientist to Alpha Helix expedition to the upper Amazon
OCR for page 425
CARROLL MILTON WILLIAMS
SELECTED BIBLIOGRAPHY
1937
425
With A. H. Clark. Records of Argynnis diana and of some other
butterflies from Virginia. [. Wash. Acad. Sci. 27:209-13.
1942
The effects of temperature gradients on the pupal-adult transfor-
mation of silkworms. Biol. Bull. 82:347-55.
With S. C. Reed and L. E. Chadwick. Frequency of wing-beat as a
character for separating species, races and geographic varieties
of Drosophila. Genetics 27:349-61.
1943
With L. A. Barness and W. H. Sawyer. The utilization of glycogen by
flies during flight and some aspects of the physiological aging of
Drosophila. Biol. Bull. 84:263-72.
With M. V. Williams. The flight muscles of Drosophila releta. i. Morphol.
72:589-99.
With L. E. Chadwick. Technique for stroboscopic studies of insect
flight. Science 98:522-24.
1944
With S. C. Reed. Physiological effects of genes: the flight of Droso-
phila considered in relation to gene mutations. Am. Nat. 78:214-
23.
1946
Continuous anesthesia for insects. Science 103:57-59.
Physiology of insect diapause: the role of the brain in the produc-
tion and termination of pupal dormancy in the giant silkworm
Platysamia cecropia. Biol. Bull. 90:234-43.
1947
Physiology of insect diapause. II. Interaction between the pupal
brain and prothoracic glands in the metamorphosis of the giant
silkworm Platysamia cecropia. Biol. Bull. 93:89-98.
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426
BIOGRAPHICAL MEMOIRS
1948
Physiology of insect diapause. III. The prothoracic glands in the
Cecropia silkworm, with special reference to their significance in
embryonic and post-embryonic development. Biol. Bull. 94:60-65.
With R. C. Sanborn. The cytochrome system in relation to diapause
and development in the Cecropia silkworm. Biol. Bull. 95:282-83.
Extrinsic control of morphogenesis as illustrated in the metamor-
phosis of insects. Growth Symposium 12:61-74.
1949
With P. C. Zamecnik, R. B. Loftfield, and M. L. Stephenson. Bio-
logical synthesis of radioactive silk. Science 109:624-26.
With L. E. Chadwick. Effects of atmospheric pressure and composi-
tion on the flight of Drosophila. Biol. Bull. 97:115-37.
1950
With R. C. Sanborn. The cytochrome system in the Cecropia silk-
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33:300-330.
The metamorphosis of insects. Sci. Am. 182:24-37.
1951
With M. I. Watanabe. Mitochondria in the flight muscles of insects.
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34:675-89.
Biochemical mechanisms in insect growth and metamorphosis. Fed.
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1952
With A. M. Pappenheimer, Jr. The effects of diphtheria toxin on
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1953
With W. Van der Kloot. Cocoon construction by the Cecropia silk-
-
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CARROLL MILTON WILLIAMS
427
worm. I. The role of the external environment. II. The role of
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With E. S. Schmidt. Physiology of insect diapause. V. Assay of the
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Insect breathing. Sci. Am. 188:28-32.
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1954
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With A. M. Pappenheimer, Jr. Cytochrome b5 and the dibydrocoenzyme
I-oxidase system in the Cecropia silkworm. J. Biol. Chem. 209:915-
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1955
With H. A. Schneiderman. An experimental analysis of the discon-
OCR for page 428
428
BIOGRAPHICAL MEMOIRS
sinuous respiration of the Cecropia silkworm. Biol. Bull. 109:123-
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1956
With L. Levenbook. Mitochondria in the flight muscles of insects.
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1957
With D. G. Shappirio. The cytochrome system of the Cecropia silk-
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1958
With W. R. Harvey. Physiology of insect diapause. XI. Cyanide-sensi-
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The juvenile hormone. Sci. Am. 198:67-75.
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1959
With L. V. Moorhead and T. F. Pulis. Juvenile hormone in thymus,
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1960
With W. FI. Telfer. The effects of diapause, development, and in-
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CARROLL MILTON WILLIAMS
429
jury on the incorporation of radioactive glycine into the blood
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1961
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1963
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1964
With B. Bowers. Physiology of insect diapause. XIII. DNA synthesis
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With F. C. Kafatos. Enzymatic mechanism for the escape of certain
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1965
With R. A. Lockshin. Programmed cell death. I. Cytology of degen-
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Neural control of the breakdown of the intersegmental muscles
of silkworms. V. Cytolytic enzymes in relation to the breakdown
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With l. H. Law. The juvenile hormone. IV. Its extraction, assay, and
purification. J. Insect Physiol. 11 :569-80.
With V. I. Brookes. Thymidine kinase and thymidylate kinase in
relation to the endocrine control of insect diapause and develop-
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With P. L. Adkisson and C. Walcott. Physiology of insect diapause.
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430
BIOGRAPHICAL MEMOIRS
XV. The transmission of photoperiod signals to the brain of the
oak silkworm, Antherea pernyi. Biol. Bull. 128:497-507.
With K. Slama. Juvenile hormone activity for the bug Pyrrhocoris
apterus. Proc. Natl. Acad. Sci. U.S.A. 54:411-14.
1966
With J. H. Law and Y. Ching. Synthesis of a material with high
juvenile hormone activity. Proc. Natl. Acad. Sci. U.S.A. 55:576-78.
With K. Slama. The juvenile hormone. N1. The sensitivity of the bug,
Pyrrhocoras apterus, to a hormonally active factor in American pa-
per-pulp. The juvenile hormone. VI. Effects of the "paper factor"
on the growth and metamorphosis of the bug, Pyrrhocoris apterus.
Biol. Bull. 130:235-46; 130:247-53.
With D. R. Walters. Reaggregation of insect cells as studied by a
new method of tissue and organ culture. Science 154:516-17.
With A. Spielman. Lethal effects of synthetic juvenile hormone on
larvae of the yellow fever mosquito, Aedes egypti. Science 154:1043-
44.
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s
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Representative terms from entire chapter:
juvenile hormone
