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OCR for page 168
C, 1. Fr o ~
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GOTTFRIED SAMUEL FRAENKEL
April 23, 1901-October 26, 1984
BY C. LADD PROSSER, STANLEY FRIEDMAN, AND
JUDITH H. WILLIS
GO T T F R ~ E D F R A E N K E E was elected to the National Acad-
emy of Sciences in 1968 for his contributions to insect
physiology. Although one might attribute his success as a pio-
neer in diverse areas behavior, endocrinology, nutrition,
insect-plant interaction to his living in a period with few
scientists ant} many uncharted fields, a reading of this bio-
graphical sketch reveals that many of his discoveries came
during periods of political upheaval, economic harclship, and
conflict with bosses not conditions generally consiclerect op-
timal for the advancement of basic research. His published
contributions to musicology indicate that he was also adept
at finding treasures in well-minec3 areas. One can only con-
clude that he was an exceptional person with an uncanny
sense of what problems were interesting, important, and solv-
able.
EARLY LIFE
Gottfried Samuel Fraenke! was born in Munich, Germany.
His father was a [ustizrat and the family typically middIe-cIass
Jewish, with interests far from the science that Fraenke] was
later to take up so successfully. As a boy he devoted much
time to music both piano playing and singing but as a
young man his major preoccupation was the Zionist cause.
169
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170
BIOGRAPHICAL MEMOIRS
He continued to pursue these activities throughout his life,
but in his early years his belief in the Zionist movement was
so strong that he made the decision to spend a part of his life
living ant! working in Palestine.
To prepare for this goal, he enrolled in a teaching degree
program at the University of Munich. There he attended
lectures and engaged in laboratory exercises under R. C.
Hertwig, Karl Von Goebel, Richard Martin, Richarc! Will-
statter, Wilhelm Konrad, W. K. Rontgen, and Karl van Frisch.
He became attracted to the field of hydrobiology and-
having cleciclecl to take a doctoral degree began to study
the life histories of certain leeches on fish. When all of his
tank specimens died as the result of a laboratory accident, he
took a short trip to the Zoological Station in Naples to obtain
fresh material. Once there, he was immediately and irrevo-
cably charmed by the enormous variety and beauty of Med-
iterranean invertebrate fauna. Already knowleclgeable about
marine invertebrates from his course with Wolfgang van
Buddenbrock at HeIgoland he began to experiment, in the
short time available, with some jellyfish blown into the Naples
harbor by a storm. Within two weeks he worked out and
successfully tested his iclea that the medusa statocysts func-
tioned as gravity receptors—a theory totally contrary to the
dogma of the time. He returned to Munich, and, being ad-
vised that his discovery was a suitable dissertation thesis, ar-
ranged for Professor O. KoehIer to "direct" it.
His talent for quickly defining and completing a project,
an ability that was to remain with him throughout his life,
was already highly developed at this early stage in his career.
Having receiver! his cloctorate, he returned to Naples on a
Rockefeller Founclation grant and, within a year, produced
six publications on various aspects of sensory physiology and
orientation of marine invertebrates. He also spent a short
period with Alfrec! Kuhn in Gottingen and found time to
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GOTTFRIED SAMUEL FRAENKEL
171
visit the marine stations at Roscoff and Plymouth. The stay
with Kuhn resulted in his first paper on insects, a behavioral
analysis of the response of bees to color. This predilection
for travel and marine stations became a lifelong passion.
After these academic adventures, Fraenke] concluder!
that it was time to fulfill his Zionist commitment. With the
small amount of money remaining from his fellowship, he
boarclect a ship for Palestine. Upon arrival he called on the
clistinguished entomologist F. S. Bodenheimer, who imme-
diately offered him a job as his assistant at the newly founded
Zoology Laboratory of the Hebrew University in Jerusalem.
There were not many young, vigorous, and experienced zo-
ologists at that time ready to work under the conditions pre-
vailing in Palestine. While visiting friends in the perioc!
before the job began, he met and shortly thereafter mar-
ried the Lithuanian-born Rachel Sobol, daughter of a fam-
ily of well-known and politically active settlers. As he later
put it, the family was not overly impressed with this scientist
and "latecomer" to Palestine.
It was cluring this sojourn in Palestine that he became
involves! with the animals he would study the rest of his work-
ing life. In those clays Jerusalem was considered a "Ion" dis-
tance" from the sea, and Fraenke! was attracted to the only
water around the papyrus pond on the grounds of the uni-
versity and to its myriad insect visitors. This ponc! provicled
subject matter for a number of fundamental studies on insect
tracheal respiration, but it was a major invasion of locusts in
1929 that finally determined Fraenkel's fate. All of the re-
search in the zoology laboratory was turned toward the prob-
lem of locust control, ant! Fraenkel's work in the desert on
locust behavior ant! sensory physiology became the basis of
attempts to hold locusts in check. The investigations are cIas-
sics of their kind and are still widely quoted today. The work
also resulted in a falling out with Bodenheimer over author-
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172
BIOGRAPHICAL MEMOIRS
ship, finally ending in Fraenkel's departure from his job and
from Palestine.
B RITAI N
He returned to Germany in 1932 when the Nazis were
already on the rise. Fraenkel felt himself fortunate to find a
position as Privat Dozent in Frankfurt's Zoology Department.
But as soon as Hitler came to power in 1933, he was dis-
missecl. Fortunately, his reputation was already sufficiently
established that he was offered a position in England. It is
worthy of note that this was one of the many positions
awarded to German refugee scientists through the Academic
Assistance Council, funded by contributions from English
scientists out of their meager, depression-level wages. Fraen-
kel never forgot this help and often spoke about it as having
saved his family. He came to University College, London, as
a research associate in 1933. His life and future scientific
activity were immediately influenced by the fly, Calliphora er-
ythrocephala, that he saw come in through an open window
and deposit its eggs on a small piece of meat. He watched the
larvae emerge from the eggs and grow and amid all the
difficulties of a new language, a new culture, a new family,
almost no salary, and using the most primitive tools—con-
ceived the idea that, within a period of two months, resulted
in the discovery of the blood-borne factor we now know to
be the insect molting hormone, ecdysterone. He submitted
his paper on this to Nature and it was printed three weeks
later his first paper in English. (Twenty-one years after
Fraenkel's discovery, the structure of the molting prehor-
mone was identified by Peter Karlson using Fraenkel's bioas-
say method.)
Fraenkel's encounters with British scientists during these
early years lest to three seminal cooperative ventures. He and
John Pringle showed that the halteres, which replace the sec-
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GOTTFRIED SAMUEL FRAENKEL
173
ond pair of wings on the adult fly, actually function as mini-
ature gyroscopes, or balance organs. Fraenke] and the phys-
ical chemist Kenneth M. Rudall analyzed the strange changes
occurring in the cuticle of the larval fly at pupariation, using,
among other methods, X-ray diffraction. This study pro-
vicled the basis for work on insect cuticle that continues to
this day, forty years later. Finally, at a chance meeting with a
behaviorist, Donald L. Gunn, Fraenke! found a willing au-
dience for his data and ideas on insect behavior informa-
tion that would eventually appear in their classic text, The
Orientation of Animals, publishecl in 1940.
By 1936, Fraenkel's reputation was such that he was of-
fered a post in insect physiology perhaps the first full-time
teaching position ever established in this discipline in the
Department of Zoology and Applied Entomology at Imperial
College of the University of Lonclon. When World War IT
came, the Department was evacuated to Slough, and, to aid
the war effort, the Pest Infestation Laboratory was created.
Professor I. W. Munro wanted Fraenke! to work on insec-
ticicles, but Fraenke] chose to take the view that unclerstand-
ing stored-grain pests would develop the intelligence with
which to deal with them successfully. Published as a series of
detailed diet studies, his findings shower! that insects have
the same nutritional requirements as man, except for the
beetle Tenebrio molitor, that needed! an additional but as yet
undefined! component in its standard! diet. By war's ens!
Fraenke! could conjecture that he had found a new vitamin.
It is doubtful whether Fraenkel's work contributed directly
to encling the war, but his experiments and their results
shaped the fields of insect nutrition and applied entomology
for years to come. His nutritional expertise, furthermore,
extended well beyond insects. As a member of a committee
organized by the Fabian Society, he investigated problems of
British agriculture after the war and wrote the chapter on
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174
BIOGRAPHICAL MEMOIRS
Britain's nutrient requirements in the committee report. He
also gained a certain notoriety in Britain as one of the cle-
signers of the British National (bread) Loaf the size, shape,
and composition of which were standardizer! cluring World
War Il.
AME R} C A
In 1947 Fraenke] paid his first visit to the United States
as a lecturer at the University of Minnesota. In 194S, after
meeting with various American entomologists, he accepted
an offer of a position in the Department of Entomology at
the University of Illinois. After his experiences with restric-
tions on research in Palestine and Slough, he later confided,
the freedom to pursue his own objectives in and of itself
justified the move to Illinois.
At Illinois, with its strong chemistry department, he began
a collaboration with Herbert Carter on his new "vitamin" that
led to the isolation, crystallization, and identification of the
Tenebr~o growth factor. He was clisappointec! when the "vita-
min" turned out to be a molecule carnitine that had been
isolated and identified fifty years earlier from mammalian
muscle. Still, no biological role had been assigned to it in the
interim, and the work of Fraenke! and his collaborators suc-
ceedec} in establishing its universality of occurrence and its
importance in Coenzyme A transfer reactions. It is worth
noting that Fraenke! himself was never satisfied that the full
spectrum of its action had yet been elucidated.
Continuing to mine the vein of insect nutrition, he next
posed an important question. If, as he had shown, all insects
had the same dietary requirements, and if, as was well known,
plant leaves generally contained all of the required com-
pounds, why were so many insects restricted in the plants
they wouIcl eat? By 195S, he had examiner! enough of the
literature to recognize that the so-called! "secondary" plant
OCR for page 175
GOTTFRIED SAMUEL FRAENKEL
175
compounds, of many different structures, might provide a
clue to the evolution of host selection. For years, botanists
and chemists hac! been isolating different classes of these
compounds associates} with different plant families but were
unable to establish functions for most of them. Fraenke]
opened a new field insect-plant coevolution based upon
chemical and sensory interactions. He clescribec! the raison
ct'etre of secondary plant compounds "as only . . . to repel ant!
attract insects." Some regarded this as a flash of insight, but
it is, as documented in his 1959 Science paper, the result of a
long, thoughtful process tempered by extensive experience
in nutrition anct behavior. In a comment made much later,
when this paper was chosen as a Citation Classic in 1984, he
described the initial resistance to his idea as follows: "Perhaps
it seemed implausible that such a simple explanation could
be virtually new and at the same time correct."
In 1961, when this idea was beginning to cause ferment
in ecological circles, Fraenke! received one of the few
Research Career Awards ever given in his field by the U. S.
Public Health Service. In collaboration with a number of his
students, he then examined the chemical basis of host selec-
tion, solidifying the theory.
Fraenkel's early work with flies still intrigued him. Using
modern techniques developed long after those halcyon days
of string ant! wax, he reexamined the tanning of adult flies
after emergence from the puparium, and promptly discov-
ered a new hormone, bursicon, which was proven responsible
for post-ecclysial activities. Interest in this hormone grew, and
by 1968 much of Fraenkel's work had been corroborated and
extended to other insects. He was electecl to the National
Academy of Sciences in that year.
By the time he retiree! in 1972 he had also vindicated his
old Calliphora assay. Responding to a challenge by Carroll M.
Williams ant! associates, Fraenke] and a Czech colleague, Jan
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176
BIOGRAPHICAL MEMOIRS
Z6arek, discovered additional factors that accelerate pupar-
· . .
sum formation.
Thanks to the enlightened. policy of the University of
Illinois in supporting the continued research of emeritus
professors, Fraenke} embarked upon a research program
after his retirement that included such topics as interactions
among nutritional states, developmental hormones, behav-
ioral changes accompanying metamorphosis, and aging.
Over the next twelve years he worked with a number of sen-
ior colleagues but most often employed bright undergrad-
uate students as his hands. In this way he helped train a
number of disciplined investigators. Throughout that time
Fraenkel's manual Smith Corona typewriter continued to
pound out research articles on diverse topics. He also main-
tainect a steady flow of correspondence with far-flung col-
leagues until only a few weeks before his death at age eighty-
four.
CLOSING REMARKS
Fraenkel's travels, both to meetings and for research pur-
poses, took him all over the world. He studied inter-tidal
snails on Bimini, leather pests in Yemen, rice leaf folders in
Sri Lanka, and silkworm nutrition in Japan. He had a collec-
tor's eye for art objects and with his love of music" the
decorative title pages of sheet music. In 1968 he published a
book deriving from this avocation, Decorative Music Title Pages
(Dover Press). He also turned up a rare and instructive edi-
tion of Hector Berlioz's Les Troyens ant! published a paper on
its significance. He was a skilled pianist and made a practice
of seeking other musicians wherever he went.
But Fraenkel's first love was biology—a love he commu-
n~cated to his two sons. Gideon Fraenke! is now professor of
chemistry at Ohio State University and Dan, professor of mi-
crobiology and molecular genetics at Harvard Medical
OCR for page 177
GOTTFRIED SAMUEL FRAENKEL 177
School. Their father diect a few months after the death of his
clevoteci wife of fifty-six years, Rachel Sobe! Fraenkel, herself
an accomplished sculptor.
Gottfried Fraenke] had that rare ability to recognize im-
portant questions and solve them with direct ant! simple tech-
niques. Ever ready to exploit the materials at hand, his work
was seminal to diverse areas of insect biology that have since
become major fielcts of study.
W E W ~ S H T O T H A N K Robert Metcalf for his help in discussing
Professor Fraenkel's scientific contributions.
OCR for page 178
178
1955
BIOGRAPHICAL MEMOIRS
HONORS AND DISTINCTIONS
1926-1927 Fellow, International Education Board, Rockefeller
Foundation
Honorary Fellow, Royal Entomological Society, Lon-
don
1962-1972 Research Career Awardee, U. S. Public Health Ser-
v~ce
1968
1972
1980
1982
1984
Member, National Academy of Sciences
Fellow, American Association for the Advancement
of Science
Honorary Doctor, Francois Rabelais University, Tours
Honorary Fellow, The Linnean Society, London
Honorary Doctor, The Hebrew University, Jerusalem
OCR for page 185
GOTTFRIED SAMUEL FRAENKEL
185
With S. Friedman and P. K. Bhattacharyya. Function of carnitine
(BT) Fed. Proc. Fed. Am. Soc. Exp. Biol., 12:414-15.
With V. I. Brookes. The process by which the puparia of many
species of flies become fixed to a substrate. Biol. Bu11.,105:442-
49.
1954
With E. W. French. Carnitine (vitamin BT) as a nutritional require-
ment for the confused flour beetle, Tribolium confusum Duval.
Nature, 173: 173.
With H. E. Gray. The carbohydrate components of honeydew.
Physiol. Zool., 27:56-65.
With P. I. Chang. Manifestations of a vitamin BT (carnitine) defi-
ciency in the larvae of the meal worm, Tenebrio molitor L. Physiol.
Zool., 27:40-56.
With G. E. Printy. The amino acid requirements of the confused
flour beetle, Tribolium confusum Duval. Biol. Bull., 106: 149-57.
With H. H. Moorefield. The character and ultimate fate of the
larval salivary secretion of Phormia regina Meigen (Diptera, Cal-
liphoridae). Biol. Bull., 106:178-84.
With H. Lipke and I. E. Liener. Effect of soybean inhibitors on
growth of Tribolium confusum. Agric. Food Chem., 2:410-14.
With P. I. Chang. Histopathology of vitamin BT (carnitine) defi-
ciency in larvae of meal worm, Tenebrio molitor L. Physiol. Zool.,
27:259-67.
The distribution of vitamin BT (carnitine) throughout the animal
kingdom. Arch. Biochem. Biophys., 50:486-95.
With S. C. Rasso. The food requirements of the adult female
blowfly, Phormia regina (Meigen), in relation to ovarian devel-
opment. Ann. Entomol. Soc. Am., 47:636-45.
With N. C. Pant. Studies on the symbiotic yeasts of two insect spe-
cies, Lasioderma serricorne F. and Stegobium paniceum L. Biol.
Bull., 107:420-32.
With N. C. Pant. On the function of the intracellular symbionts of
Oryzoephilus surinamensis L. (Cucujidae, Coleoptera). l. Zool.
Soc. India, 6: 173-77.
1955
Inhibitory effects of sugars on the growth of the mealworm, Tene-
brio molitor L. J.. Cell Comp. Physiol., 45:393-408.
OCR for page 186
186
BIOGRAPHICAL MEMOIRS
With P. K. Bhattacharyya and S. Friedman. The effect of some
derivatives and structural analogues of carnitine on the nutri-
tion of Tenebrio molitor. Arch. Biochem. Biophys., 54:424-31.
With S. Friedman, T. Hinton, S. Laszlo, and I. L. Noland. The
effect of substituting carnitine for choline in the nutrition of
several organisms. Arch. Biochem. Biophys., 54:432-39.
With H. Lipke. The toxicity of corn germ to the meal worm, Tene-
brio molitor. ],. Nutr., 55: 165-78.
With M. Brust. The nutritional requirements of the larvae of a
blowfly, Phormia regina Meig. Physiol. Zool., 28: 186-204.
With S. Friedman. Reversible enzymatic acetylation of carnitine.
Arch. Biochem. Biophys., 59:491-501.
With S. Friedman, J. E. McFarlane, and P. K. Bhattacharyya. Quan-
titative separation and identification of quaternary ammonium
bases. Arch. Biochem. Biophys., 59:484-90.
1956
.
With H. Lipke. Insect nutrition. Annul Rev. Entomol., 1:17-44.
Insects and plant biochemistry. The specificity of food plants for
insects. Proc. 14th Int. Congr. Zool. Copenhagen, pp. 383-87.
With I. Leclercq. Nouvelles recherches sur les besoins nutritifs de
la larve du Tenebrio molitor L. (Insecte, Coleoptere) Arch. Int.
Physiol. Biochim., 64:601-22.
With K. Bloch, R. G. Langdon, and A. t. Clark. Impaired steroid
biogenesis in insect larvae. Biochim. Biophys. Acta., 21:176.
1957
The Tenebrio assay for carnitine. In: Methods of Enzymology, ed. S. P.
Colowick and N. O. Kaplan, New York: Academic Press, vol. 3,
pp. 662-67.
With S. Friedman and A. B. Galun. Isolation and physiological
action of (+~-carnitine. Arch. Biochim. Biophys., 66:10-15.
With T. Ito. y-butyrobetaine as a specific antagonist for carnitine
in the development of the early chick embryo. l. Gen. Physiol.,
41 :279-88.
With S. Friedman. Carnitine. Vitam. Horm., 15:73-118.
With R. Galun. Physiological effects of carbohydrates in the nutri-
tion of a mosquito, Aedes aegypti and two flies, Sarcophaga bullata
and Musca domestica. J. Cell. Comp. Physiol., 50: 1-Y3.
OCR for page 187
GOTTFRIED SAMUEL FRALNKEL
187
1958
With V. I- Brooks. The nutrition of the larva of the housefly Musca
domestica L. Physiol. Zool., 31:208-23.
The effect of zinc and potassium in the nutrition of Tenebrio molitor,
with observations on the expression of a carnitine deficiency. I.
Nutr., 65:361-96.
The basis of food selection in insects which feed on leaves. Ab-
stracts of Invitational Papers. 18th Annul Meet. Entomol. Soc.
of Japan. 5 pp.
1959
A historical and comparative survey of the dietary requirements of
insects. Ann. N.Y. Acad. Sci., 77:267-74.
The raison d'e^tre of secondary plant substances. Science, 129: 1466-
70.
The chemistry of host specificity of phytophagous insects. In: Bio-
chemistry of Insects, 4th Int. Congr. Biochem., London: Perga-
mon Press, vol. 12, pp. 1-14.
With T. Ito and Y. Horie. Feeding on cabbage and cherry leaves by
maxillectomized silkworm larvae. I. Seric. Sci. Jpn., 28: 107-13.
With R. T. Yamamoto. Common attractant for the tobacco horn-
worm, Protoparce sexta ~ Nathan.) and the Colorado potato beetle,
Leptinotarsa decemlineata (Say). Nature, 184:206-7.
1960
With R. T. Yamamoto. The specificity of the tobacco hornworm,
Protoparce sexta to solanaceous plants. Ann. Entomol. Soc. Am.,
53:503-7.
With R. T. Yamamoto. Assay of the principal gustatory stimulant
for the tobacco hornworm, Protoparce sexta from solanaceous
plants. Ann. Entomol. Soc. Am., 53:499-503.
With R. T. Yamamoto. The suitability of tobaccos for the growth
of the cigarette beetle, Lasioderma serricorne. J. Econ. Entomol.,
53:381-84.
Lethal high temperatures for three marine invertebrates, Limulus
polyphemus, Littorina littorea and Pagurus longicarpus. Oikos,
11: 171-82.
With S. Friedman, I. E. McFarlane, and P. K. Bhattacharyya.
OCR for page 188
188
BIOGRAPHICAL MEMOIRS
(-~-Carnitine chloride. In: Biochemical Preparations, 7:26-30.
New York: John Wiley & Sons.
196
A new type of negative phototropotaxis observed in a marine iso-
pod, Eurydice. Physiol. Zool., 34:228-32.
Resistance to high temperatures in a Mediterranean snail, Littorals
neritoides. Ecology, 42:604-6.
Quelques observations sur le comportement de Convoluta roscoffen-
s~s. Cab. Biol. Mar., 2:155-60.
With G. P. Waldbauer. Feeding on normally rejected plants by max-
illectomized larvae of the tobacco hornworm, Protoparce sexta
(Lepidoptera, Sphingidae). Ann. Entomol. Soc. Am., 54:477—
85.
With D. L. Gunn. The Orientation of Animals. Kineses, Taxes and
Compass Reactions. New York: Dover Publications, Inc.376 pp.
With Galun, R. The effect of low atmospheric pressure on adult
Aedes aegypti and on housefly pupae. J. Insect Physiol., 7:161-
76.
Die biologische Funktion der sekundaren Pflanzenstoffe im Allge-
meinen und solcher Stoffe in Solanaceen im Besonderen. In:
Chemie und Biochemie der Solanum-Alkaloide, Tagungsberichte 27,
Int. Symp. Deutsch. Akad. Landw., Berlin, pp. 297-307.
1962
The physiology of insect nutrition. (Atti del Simposio Internazion-
ale di Biologi Sperimentale, Celebrazione Spallanzaniana, Reg-
gio Emilia-Pavia, May 2-7, 1961.) Symp. Genet. Biol. Ital.,9:3-
11.
With J. Nayar, O. Nalbandov, and R. T. Yamamoto. Further inves-
tigations into the chemical basis of the insect-host plant rela-
tionship, XI. Int. Congr. Entomol. Vienna, Verhandlungen,
3: 122-26.
With R. T. Yamamoto. The physiological basis for the selection of
plants for egg-laying in the tobacco hornworm, Protoparce sexta
(Johan.~. XI. Inter. Congr. Entomol. Vienna, Verhandlungen,
3: 127-133.
With J. K. Nayar. The chemical basis of host plant selection in the
silkworm, Bombyx mor? (L.). J. Insect Physiol., 8:505-25.
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GOTTFRIED SAMUEL FRAENKEL
189
With C. Hsiao. Hormonal and nervous control of tanning in the
fly. Science, 138:27-29.
1963
With I. K. Nayar. The chemical basis of host selection in the Catalpa
sphinx, Ceratomia castalpae (Lepidoptera, Sphingidae). Ann. En-
tomol. Soc. Am., 56:119-22.
With I. K. Nayar. The chemical basis of the host selection in the
Mexican bean beetle, Epilachna varivestis (Coleoptera, Coccinel-
lidae). Ann. Entomol. Soc. Am., 56:174-78.
With l. K. Nayar. Practical methods of year-round laboratory rear-
ing of the silkworm, Bombyx mori (L.) (Lepidoptera, Bombyci-
dae). Ann. Entomol. Soc. Am., 56:122-23.
With C. Hsiao. Tanning in the adult fly: A new function of neu-
rosecretion in the brain. Science, 141:1057-58.
Berlioz, the princess and 'Les Troyens.' Mus. Let., 44:249-56.
1964
With O. Nalbandov and R. T. Yamamoto. Insecticides from plants.
Nicandrenone, a new compound with insecticidal properties,
isolated from Nicandra physalodes. Agric. Food Chem., 12 :55-
59.
With C. F. Soo Hoo. The resistance of ferns to the feeding of Pro-
denia eridania larvae. Ann. Entomol. Soc. Am., 57:788-90.
With C. F. Soo Hoo. A simplified laboratory method for rearing
the Southern armyworm, Prodenia eridania for feeding experi-
ments. Ann. Entomol. Soc. Am., 57:798-99.
1965
With C. Hsiao. Bursicon, a hormone which mediates tanning of
the cuticle in the adult fly and other insects. }. Insect Physiol.,
11:513-56.
A brief survey of the recognition of carnitine as a substance of
physiological importance. In: Recent Research on Carnitine. Its
Relation to Lipid Metabolism, ed. G. Wolf, Cambridge: The MIT
Press, pp. 1-3.
1966
With C. Hsiao and M. Seligman. Properties of bursicon: An insect
hormone that controls cuticular tanning. Science, 151:91-93.
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190
BIOGRAPHICAL MEMOIRS
With R. D. Pausch. The nutrition of the larva of the oriental rat
flea, Xenopsylla cheopis (Rothschild). Physiol. Zool., 39:202-22.
With C. Hsiao. Neurosecretory cells in the central nervous system
of the adult blowfly, Phormia regina Meigen (Diptera, Caliphor-
idae). J. hIorphol., 119:21-38.
With C. F. Soo Hoo. The consumption, digestion and utilization of
food plants by a polyphagous insect, Prodenia eridania (Cramer).
I. Insect Physiol., 12:71 1-30.
With C. F. Soo Hoo. The consumption, digestion and utilization of
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GOTTFRIED SAMUEL FRAENKEL 191
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With E. Zlotkin. Acceleration of puparium formation in Sarcophaga
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OCR for page 192
192
BIOGRAPHICAL MEMOIRS
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With D. L. Denlinger and J. H. Willis. Rates and cycles of oxygen
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With D. M. DeGuire. The meconium of Aedes aegypti (Diptera: Cul-
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With G. Bhaskaran. Pupariation and pupation in cyclorrhaphous
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With N. P. Ratnasiri. Inhibition of purpariation in Sacrophaga bul-
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With N. Ratnasiri. The physiological basis of anterior inhibition of
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GOTTFRIED SAMUEL FRAENKEL
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With P. Sivasubramanian and S. Friedman. Nature and role of pro-
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Interactions between ecdysone, bursicon, and other endocrines
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Molting and development in undersized fly larvae. In: The Insect
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With A. Blechl, I. Blechl, P. Herman, and M. Seligman. 3':5'-cyclic,
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With M. Hollowell. Actions of the juvenile hormone, 20-hydroxy-
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The proposed vitamin role of carnitine. In: Carnitine Biosynthesis,
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OCR for page 194
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BIOGRAPHICAL MEMOIRS
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1981
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Food conversion efficiency by fleshily larvae, Sarcophaga bullata.
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~ _ ~ 1 _ _ . . ~ ~
~ _ C~
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1984
This week's citation classic. The raison d'etre of secondary plant sub-
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1986
With I. Zdarek and S. Reid. How does an eclosing fly deal with
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1987
With I. Zdarek. Pupariation in flies: A tool for monitoring effects
of drugs, venoms and other neurotoxic compounds. Arch. In-
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GOTTFRIED SAMUEL FRAENKEL
195
With S. N. M. Reid and S. Friedman. Extrication, the primary event
in eclosion, and its relationship to digging, pumping and tan-
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With S. N. M. Reid and S. Friedman. Extrication, the primary event
in eclosion, and its neural control in Sarcophaga bullata. J. Insect
Physiol., 33:481-486.
Representative terms from entire chapter:
sarcophaga bullata
