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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
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 receptorsa 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
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-
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 toolscon- 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-
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
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
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
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 biologya 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
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.
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
GOTTFRIED SAMUEL FRAENKEL SELECTED BIBLIOGRAPHY 1925 Der statische Sinn der Medusen. Z. Vgl. Physiol., 2:658-90. 1927 179 Phototropotaxis bei Meerestieren. Die Naturwissenschaften, 14: 117-22. Beitrage zur Biologie eines Arcturiden. Zoo. Anz., 69:219-22. Beitrage zur Geotaxis und Phototaxis von Littor~na. Z. Vgl. Physiol., 5:585-97. Die Grabbewegungen der Soleniden. Z. Vgl. Physiol., 6:167-220. Uber Photomenotaxis bei Elysia virid~s. Mont. Z. Vgl. Physiol., 6:385-401. Biologische Beobachtungen an Janthina. Morphol. Oekol. Tiere, 7:597-608. With Kuhn, A. Uber das Unterscheidungsvermogen der Bienen fur Wellenlangen im Spektrum. Nachr. Ges. Wiss. Gottingen. Math. Phys. K.:1-6. 1928 Uber den Auslosungsreiz des Umdrehreflexes bei Seestsernen und Schlangensternen. Z. Vgl. Physiol., 7:365-78. 1929 Uber die Geotaxis von Convoluta rosco~ens~s. Z. Vgl. Physiol., 10:237-47. Untersuchungen uber Lebensgewohnheiten, Sinnesphysiologie und Sozialpsychologie der wandernden Larven der afrikan- ischen Wanderheuschrecke Sch~stocerca gregar~a (Forsk.~. Biol. Zentralb., 46:657-80. 1930 Der Atmungsmechanismus des Skorpions. Z. Vgl. Physiol., 11:656-61. Die Orientierung von Schtstocerca gregar~a zu strahlender Warme. Z. Vgl. Physiol., 13:300-13. Beitrage zur Physiologie der Atmung der Insekten. Atti 11. Congr. Int. Zool. Pavoda, pp. 905-21. With F. S. Bodenheimer, K. Reich, and N. Segal. Studien zur
180 BIOGRAPHICAL MEMOIRS Epidemiologie, Okologie und Physiologie der afrikanischen Wanderheuschrecke. (Schistocerca gregaria Forsk.) Z. Angew. Entomol., 15:435-557. 1931 Die Mechanik der Orientierung der Tiere im Raum. Biol. Rev., 6:36-87. 1932 Untersuchungen uber die Koordination von Reflexen und auto- matisch-nervosen Rhythmen bei Insekten. I. Die Flugreflexe der Insekten und ihre Koordination. Z. Vgl. Physiol., 16:371- 93. Untersuchungen uber die Koordination von Reflexen und auto- matisch-nervosen Rhythmen bei Insekten. II. Die nervose Re- gulierung der Atmung wahrend des Fluges. Z. Vgl. Physiol., 16:394-417. Untersuchungen uber die Koordination von Reflexen und auto- matisch-nervosen Rhythmen bei Insekten. III. Das Problem des gerichteten Atemstromes in den Tracheen der Insekten. Z. Vgl. Physiol, 16:418-43. Untersuchungen uber die Koordination von Reflexen und auto- matisch-nervosen Rhythmen bei Insekten. IV. Uber die nervo- sen Zentren der Atmung und die Koordination ihrer Tatigkeit. Z. Vgl. Physiol., 16:444-62. Die Wanderungen der Insekten. Ergeb. Biol., 9: 1-238. 1934 Der Atmungsmechanismus der Vogel wahrend des Fluges. Biol. Zentralbl., 54:96- 101. Pupation of flies initiated by a hormone. Nature, 133:834. 1935 A hormone causing pupation in the blowfly Calliphora erythroce- phala. Proc. R. Soc. London Ser. B., 118:1-12. 1936 Observations and experiments on the blowfly (Calliphora erythroce- phala) during the first day after emergence. Proc. Zool. Soc. London, pp. 893-904.
GOTTFRIED SAMUEL FRAENKEL 181 Utilization of sugars and polyhydric alcohols by the adult blowfly. Nature, 137:237. 1938 With G. V. B. Herford. The respiration of insects through the skin. J. Exp. Biol., 15:266-80. With J. W. S. Pringle. Halteres of flies as gyroscopic organs of equi- librium. Nature, 141:919. Temperature adaptation and the physiological action of high tem- peratures. Kongressbericht II. 16th Int. Physiol. Congr. Zurich. (Abstract). With I. L. Harrison. Irregular abdomina in Calliphora erythrocephala (Mg.~. Proc. R. Entomol. Soc. London (A), 13:95-96. The evagination of the head in the pupae of cyclorrhaphous flies (Diptera). Proc. R. Entomol. Soc. London (A), 13:137-39. The number of moults in the cyclorrhaphous flies (Diptera). Proc. R. Entomol. Soc. London (A), 13:158-60. 1939 The function of the halteres of flies (Diptera). Proc. Zool. Soc. Lon- don (A), 109:69-78. 1940 Utilization and digestion of carbohydrates by the adult blowfly. I. Exp. Biol., 17: 18-29. With K. M. Rudall. A study of the physical and chemical properties of the insect cuticle. Proc. R. Soc. London (B), 129: 1-35. With H. S. Hopf. The physiological action of abnormally high tem- peratures on poikilothermic animals. I. Temperature adapta- tion and the degree of saturation of the phosphatides. Bio- chem. J., 34:1085-92. With G. V. B. Herford. The physiological action of abnormally high temperatures on poikilothermic animals. II. The respiration at high sublethal and lethal temperatures. }. Exp. Biol., 17:386- 95. With R. A. Davis. The oxygen consumption of flies during flight. J. Exp. Biol., 17:402-7. With D. L. Gun. The Orientation of Animals. Oxford: Clarendon Press. 352 pp.
182 BIOGRAPHICAL MEMOIRS 1941 With I. A. Reid and M. Blewett. The sterol requirements of the larva of the beetle, Dermestes vulpinus Fabr. Biochem. }.,35:712- 20. With M. Blewett. Deficiency of white flour in riboflavin (tested with the flour beetle Tr~bolium confusum). Nature, 147:716. 1942 With M. Blewett. Biotin as a possible growth factor for insects. Nature, 149:301. With M. Blewett. Biotin, B. riboflavin, nicotinic acid, Be, and pan- tothenic acid as growth factors for insects. Nature, 150: 177. 1943 With M. Blewett. Vitamins of the B-group required by insects. Na- ture, 151:703. With M. Blewett. Intracellular symbionts of insects as a source of vitamins. Nature, 152:506. Insect nutrition. I. R. Coll. Sci., 13:59-69. With M. Blewett. The basic food requirements of several insects. I. Exp. Biol, 20:28-34. With M. Blewett. The vitamin B-complex requirements of several insects. Biochem. l., 37:686-92. With M. Blewett. The sterol requirements of several insects. Bio- chem. J., 37:692-95. With M. Blewett. The natural foods and the food requirements of several species of stored products insects. Trans. R. Entomol. Soc. London, 93:457-90. 1944 With M. Blewett. Intracellular symbiosis and vitamin requirements of two insects, Lasioderma serr?corne and Sitodrepa panicea. Proc. R. Soc. London (B), 132:212-21. With M. Blewett. Stages in the recognition of biotin as a growth factor for insects. Proc. R. Entomol. Soc. London (A), 19:30- 35. With M. Blewett. The utilization of metabolic water in insects. Bull. Entomol. Res., 35:127-39.
GOTTFRIED SAMUEL FRAENKEL 1945 183 With M. Blewett. Linoleic acid, ~x-tocopherol and other fat-soluble substances as nutritional factors for insects. Nature, 155:392. 1946 With M. Blewett. The dietetics of the clothes moth, Tineola bissel- liella. Hum. l. Exp. Biol., 22:156-61. With M. Blewett. The dietetics of the caterpillars of three Ephestia species, E. knehniella, E. elutella and E. cautella, and of a closely related species, Plodia interpunctella. ]. Exp. Biol., 22: 162-71. With M. Blewett. Linoleic acid, vitamin E and other fat-soluble substances in the nutrition of certain insects, (Ephestia knehniella, E. elutella, E. cautella and Plodia interpunctella ELep.~. I. Exp. Biol., 22: 172-90. With M. Blewett. Folic acid in the nutrition of certain insects. Na- ture, 157:697. Britain's nutritional requirements. In: Towards a Socialist Agriculture. Studies by a Group of Fabians, ed. F. W. Bateson, London: Victor Gollancz, pp. 42-76. 1947 With K. M. Rudall. The structure of insect cuticles. Proc. R. Soc. London (B), 134:111-43. With M. Blewett. The importance of folic acid and unidentified members of the vitamin B complex in the nutrition of certain insects. Biochem. J., 41 :469-75. With M. Blewett. Linoleic acid and arachidonic acid in the metab- olism of two insects, Ephestia knehniella (Lep.) and Tenebrio molitor (Col.~. Biochem. l., 41 :475-78. With P. Ellinger and M. M. Abdel Kader. The utilization of nico- tinamide derivatives and related compounds by mammals, in- sects and bacteria. Biochem. }., 41:559-68. 1948 BT, a new vitamin of the B-group and its relation to the folic acid group and other anti-anemia factors. Nature, 161:981-83. The effects of a relative deficiency of lysine and tryptophane in the diet of an insect, Tribolium confusum. Biochem. }., 43: Proceedings XIV.
184 BIOGRAPHICAL MEMOIRS Evidence for the need, by certain insects, for three chemically un- identified factors of the vitamin B-complex. Br. I. Nutr.2, Abstr. Commun., l:ii. 1950 The nutrition of the meal worm, Tenebrio molitor L. (Tenebrionidae, Coleoptera). Physiol. Zool., 23:92-108. With N. C. Pant. The function of the symbiotic yeasts of two insect species, Lasioderma serricorne F. and Stegobium (Sitodrepa) pani- ceum L. Science, 112:498-500. 1951 With H. R. Stern. The nicotinic acid requirements of two insect species in relation to the protein content of their diets. Arch. Biochem., 30:438-44. Effect and distribution of vitamin BT. Arch. Biochem. Biophys. 34:457-67. Isolation procedures and certain properties of vitamin BT. Arch. Biochem. Biophys., 34:468-77. 1952 With H. E. Carter, P. K. Bhattacharyya, and K. Weidman. The identity of vitamin BT with carnitine. Arch. Biochem. Biophys., 35:241-42. With M. I. Cooper. Nutritive requirements of the small-eyed flour beetle, Palorus ratzeburg' Wissman (Tenebrionidae, Coleoptera), Physiol. Zool., 25:20-28. The role of symbionts as sources of vitamins and growth factors for their insect hosts. Tijdschr. Entomol., 95: 183-95. The nutritional requirements of insects for known and unknown vitamins. Trans. 9th Int. Congr. Entomol. Amsterdam, 1951, 1 :277-80. 1953 The nutritional value of green plants for insects. Trans. 9th Int. Congr. Entomol. Amsterdam, 1951, 2:90-100. With H. E. Gray. Fructomaltose, a recently discovered trisaccharide isolated from honeydew. Science, 118:304-5. Studies on the distribution of vitamin BT (carnitine). BiO1. BU11., 104:359-71.
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.
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.
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.
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.
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.
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 food plants by a polyphagous insect, Prodenia eridania (Cramer). J. Insect Physiol., 12:71 1-30. With T. Ito. The effect of nitrogen starvation on Tenebrio molitor L. I. Insect Physiol., 12:803-17. The heat resistance of intertidal snails at Shirahama, Wakyama- ken, Japan. Publ. Seto Mar. Biol. Lab., 14:185-95. 1967 With C. Hsiao. Calcification, tanning, and the role of ecdyson in the formation of the puparium of the facefly, Musa autumnalis. I. Insect Physiol., 13:1387-94. 1968 With C. Hsiao. Manifestations of a pupal diapause in two species of flies, Sarcophaga arg~rostorna and S. bullata. J. Insect Physiol., 14:689-705. With C. Hsiao. Morphological and endocrinological aspects of pu- pal diapause in a fleshily, Sarcophaga argyrostoma. J. Insect Phys- iol., 14:707-18. The heat resistance of intertidal snails at Bimini, Bahamas; Ocean Springs, Mississippi; and Woods Hole, Massachusetts. Physiol. Zool., 41:1-13. With T. H. Hsiao. The influence of nutrient chemicals on the feed- ing behavior of the Colorado potato beetle, Leptinotarsa decem- lineata (Coleoptera: Chrysomelidae). Ann. Entomol. Soc. Am., 6 1 :44-54. With T. H. Hsiao. Isolation of phagostimulatory substances from the host plant of the Colorado potato beetle. Ann. Entomol. Soc. Am., 61:476-84. With T. H. Asian. The role of secondary plant substances in the
GOTTFRIED SAMUEL FRAENKEL 191 food specificity of the Colorado potato beetle. Ann. Er~tomol. Soc. Am., 61:485-93. With T. H. Hsiao. Selection and specificity of the Colorado potato beetle for solanaceous and nonsolanaceous plants. Ann. Ento- ~nol. Soc. Am., 61:493-503. Decorative Music Title Pages. New York: Dover Publications. 230 pp. 1969 With W. Fogal. Melanin in the puparium and adult integument of the fleshily, Sarcophaga bullata. ]. Insect Physiol., 15: 1437-47. With W. H. Fogal. The role of bursicon in melanization and en- docuticle formation in the adult fleshily, Sarcophaga bullata. J. Insect Physiol., 15: 1235 -47. With M. Seligman and S. Friedman. Hormonal control of turnover of tyrosine and tyrosine phosphate during tanning of the adult cuticle in the fly, Sarcophaga bullata. ]. Insect Physiol., 15: 1085- 101. With M. Seligman and S. Friedman. Bursicon mediation of tyro- sine hydroxylation during tanning of the adult cuticle of the fly, Sarcophaga bullata. ]. Insect Physiol., 15:553-62. With T. H. Hsiao. Properties of leptinotarsin: A toxic hemolymph protein from the Colorado potato beetle. Toxicon, 7:119-30. With P. Berreur. Puparium formation in flies: Contraction to pu- parium induced by ecdysone. Science, 164: 1182-83. With I. Z6arek. Correlated effects of ec~ysone and neurosecretion in puparium formation (pupariation) of flies. Proc. Natl. Acad. Sci. USA, 64:565-72. Evaluation of our thoughts on secondary plant substances. Ento- mol. Exp. Appl., 12:473-86. 1970 With E. Zlotkin. Acceleration of puparium formation in Sarcophaga argyrostoma by electrical stimulation or scorpion venom. I. Insect Physiol., 16:549-54. With l. Zdarek. The evaluation of the "Calliphora test" as an assay for ecdysone. Biol. Bull., 139: 138-50. With W. Fogal. Histogenesis of the cuticle of the adult flies, Sarco- phaga bullata and S. argyrostoma. ]. Morphol., 130: 137-50. With l. Zdarek. Overt and covert effects of endogenous and
192 BIOGRAPHICAL MEMOIRS exogenous ecdysone in puparium formation of flies. Proc. Natl. Acad. Sci. USA, 67:331-37. 1971 With I. Zdarek. Neurosecretory control of ecdysone release during puparium formation of flies. Gen. Comp. Endocrinol.,17:483- 89. With E. Zlotkin, F. Miranda, and S. Lissitzky. The effect of scorpion venoms on blowfly larvae a new method for the evaluation of scorpion venoms potency. Toxicon, 9: 1-8. 1972 With D. L. Denlinger and J. H. Willis. Rates and cycles of oxygen consumption during pupal diapause in Sarcophaga flesh flies. J. Insect. Physiol., 18:871-82. With I. Zdarek. The mechanism of puparium formation in flies. J. Exp. Zool., 179:315-24. With S. Friedman. Carnitine. In: The Vitamins, ed. W. H. Sebrell and R. S. Harris, New York and London: Academic Press, vol. 5., pp. 329-55. 1973 With D. M. DeGuire. The meconium of Aedes aegypti (Diptera: Cul- icidae). Ann. Entomol. Soc. Am., 66:475-76. With G. Bhaskaran. Pupariation and pupation in cyclorrhaphous flies (Diptera): terminology and interpretation. Ann. Entomol. Soc. Am., 66:418-22. With N. P. Ratnasiri. Inhibition of purpariation in Sacrophaga bul- lata. Nature, 243:91-93. With N. Ratnasiri. Anterior inhibition of pupariation in ligated larvae of Sarcophaga bullata and other fly species: Incidence and expression. Ann. Entomol. Soc. Am., 67:195-203. 1974 With N. Ratnasiri. The physiological basis of anterior inhibition of puparium formation in ligated fly larvae. l. Insect Physiol., 20: 105-19. With P. Sivasubramanian and H. S. Ducoff. Effect of X-irradiation on the formation of the puparium in the fleshfly, Sarcophaga bullata.~. Insect Physiol., 20:1303-17.
GOTTFRIED SAMUEL FRAENKEL 193 With P. Sivasubramanian and S. Friedman. Nature and role of pro- teinaceous hormonal factors acting during puparium forma- tion in flies. Biol. Bull., 147: 163-85. 1975 Interactions between ecdysone, bursicon, and other endocrines during puparium formation and adult emergence in flies. Am. Zool. 15 Suppl. 1, 15:29-48. 1976 Molting and development in undersized fly larvae. In: The Insect Integument, ed. H. R. Hepburn, Amsterdam: Elsevier, pp. 323- 38. 1977 With A. Blechl, I. Blechl, P. Herman, and M. Seligman. 3':5'-cyclic, AMP, and hormonal control of puparium formation in the fleshily Sarcohpaga bullata. Proc. Natl. Acad. Sci. USA, 74:2182- 86. With C. Pappas. Nutritional aspects of oogenesis in the flies Phor- mia regina and Sarcophaga bullata. Physiol. Zool., 50:237-46. With C. Pappas. Hormonal aspects of oogenesis in the files Phormia regina and Sarcophaga bullata. ]. Insect Physiol., 24:75-80. With M. Seligman, A. Blechl, I. Blechl, and P. Herman. Role of ecdysone, pupariation factors, and cyclic AMP in formation and tanning of the puparium of the fleshily Sarcophaga bullata. Proc. Natl. Acad. Sci. USA, 74:4697-701. 1979 With M. Hollowell. Actions of the juvenile hormone, 20-hydroxy- ecdysone and the oostatic hormone during oogenesis in the flies Phormia regina and Sarcophaga bullata. J. Insect Physiol.,25:305- 10. With }. Zdarek and K. Slama. Changes in internal pressure during puparium formation in flies. J. Exp. Zool., 207:187-95. 1980 The proposed vitamin role of carnitine. In: Carnitine Biosynthesis, Metabolism and Functions, ed. R. E. Fraenkel and l. D. McGarry, New York: Academic Press, pp. 1-6.
194 BIOGRAPHICAL MEMOIRS Foreword and overview. In: Neurohormonal Techniques in Insects, ed. T. A. Miller, Heidelberg: Springer-Verlag, pp. IX-XV. 1981 Importance of allelochemics in plant insect relations. In: The lEcol- ogy of Breuchids Attacking Legumes (Pulses), ed. V. Labeyrie, Hingham, Mass.: Junk Publishers, pp. 57-60. With F. Fallil. The spinning (stitching) behavior of the rice leaf folder, Cnaphalocrocis medinalas. Entomol. Exp. Appl., 29:138- 46. With F. Fallil and K. S. Kumarasinghe. The feeding behavior of the rice leaf folder, Cnaphalocrocis medinalis. Entomol. Exp. Appl. 29:147-61. With B. Bennetova. What determines the number of ovarioles in a fly ovary? I. Insect Physiol., 27:403-10. Food conversion efficiency by fleshily larvae, Sarcophaga bullata. Physiol. Entomol., 6:157-60. With I. Zdarek, R. Rohlf, and I. Blechl. A hormone enacting im- ~ _ ~ 1 _ _ . . ~ ~ ~ _ C~ o['lllzauon In puparlaung ny larvae. J. Exp. Biol., 93:51-63. 1984 This week's citation classic. The raison d'etre of secondary plant sub- stances. C. C. Life Sci., 11: 18. With I. Zdarek, I. Zavidilova, and I. Su. Post-eclosion behaviour of flies after emergence from the puparium. Acta Entomol. Bo- hemoslov., 81: 161-70. With l. Su. Hormonal control of eclosion of flies from the pupar- ium. Proc. Natl. Acad. Sci. USA, 81:1457-59. With J. Su and J. Zdarek. Neuromuscular and hormonal control of post-eclosion processes in flies. Arch. Insect Biochem. Physiol., 1 :345-66. 1986 With I. Zdarek and S. Reid. How does an eclosing fly deal with obstacles? Physiol. Entomol., 11: 107-14. 1987 With I. Zdarek. Pupariation in flies: A tool for monitoring effects of drugs, venoms and other neurotoxic compounds. Arch. In- sect Biochem. Physiol., 4:29~6.
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- ning in Sarcophaga bullata. J. Insect Physiol., 33 :339-348. 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.