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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 5C 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 5C and the other at 25 to 30C. 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 25C, 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

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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

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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 exceptionalcertainly 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.

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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

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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- worm with special reference to a new component. [. Gen. Physiol. 33:300-330. The metamorphosis of insects. Sci. Am. 182:24-37. 1951 With M. I. Watanabe. Mitochondria in the flight muscles of insects. I. Chemical composition and enzymic content. i. Gen. Physiol. 34:675-89. Biochemical mechanisms in insect growth and metamorphosis. Fed. Proc. 10:546-52. 1952 With A. M. Pappenheimer, Jr. The effects of diphtheria toxin on the Cecropia silkworm. {. Gen. Physiol. 35: 727-40. Physiology of insect diapause. IV. The brain and prothoracic glands as an endocrine system in the Cecropia silkworm. Biol. Bull., pp. 120-38. Morphogenesis and metamorphosis of insects. Harvey Lect. 47:126- 55. 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 the internal environment. Behavior5:141-56; 5:157-74. With W. H. Telfer. Immunological studies of insect metamorphosis. I. Qualitative and quantitative description of the blood antigens of the Cecropia silkworm. {. Gen. Physiol. 36:389-413. With E. S. Schmidt. Physiology of insect diapause. V. Assay of the growth and differentiation hormone of lepidoptera by the method of tissue culture. Biol. Bull. 105:174-87. With H. A. Schneiderman and M. Ketchel. Physiology of insect dia- pause. VI. Effects of temperature, oxygen tension and metabolic inhibitors on in vitro spermatogenesis in the Cecropia silkworm. Biol. Bull. 105: 188-99. With M. I. Watanabe. Mitochondria in the flight muscles of insects. II. Effects of the medium on the size, form and organization of isolated sarcosomes. {. Gen. Physiol. 37:71-90. Insect breathing. Sci. Am. 188:28-32. With H. A. Schneiderman. Physiology of insect diapause. VII. Respi- ratory metabolism of the Cecropia silkworm during diapause and adult development. Biol. Bull. 105:320-34. 1954 With W. Van der Kloot. Cocoon construction by the Cecropia silk- worm. III. The alteration of spinning behavior by chemical and surgical techniques. Behavior 6:233-55. With H. A. Schneiderman. Physiology of insect diapause. VIII. Qualitative changes in the metabolism of the Cecropia silkworm during dia- pause and development. IX. The cytochrome oxidase system in relation to the diapause and development of the Cecropia silk- worm. Biol. Bull. 106:210-29; 106:238-52. With W. H. Telfer. Immunological studies of insect metamorphosis. II. The role of a sex-limited blood protein in egg formation by the Cecropia silkworm. i. Gen. Physiol. 37:539-58. With A. M. Pappenheimer, Jr. Cytochrome b5 and the dibydrocoenzyme I-oxidase system in the Cecropia silkworm. J. Biol. Chem. 209:915- 29. 1955 With H. A. Schneiderman. An experimental analysis of the discon-

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428 BIOGRAPHICAL MEMOIRS sinuous respiration of the Cecropia silkworm. Biol. Bull. 109:123- 43. 1956 With L. Levenbook. Mitochondria in the flight muscles of insects. III. Mitochondrial cytochrome c in relation to the aging and wing beat frequency of flies. J. Gen. Physiol. 39:497-512. Physiology of insect diapause. X. An endocrine mechanism for the influence of temperature on the diapausing pupa of the Cecropia silkworm. Biol. Bull. 110:201-18. The juvenile hormone of insects. Nature 178:212-13. 1957 With D. G. Shappirio. The cytochrome system of the Cecropia silk- worm. I. Spectroscopic studies of individual tissues. II. Spectro- scopic studies of oxidative enzyme systems in the wing epithe- lium. Proc. R. Soc. Lond. B. 147:218-32; 147:233-46. 1958 With W. R. Harvey. Physiology of insect diapause. XI. Cyanide-sensi- tivity of the heartbeat of the Cecropia silkworm, with special refer- ence to the anaerobic capacity of the heart. XII. The mechanism of carbon monoxide sensitivity and insensitivity during the pupal diapause of the Cecropia silkworm. Biol. Bull. 114:23-35; 114:36- 53. The juvenile hormone. Sci. Am. 198:67-75. Hormonal regulation of insect metamorphosis. In Chemical Basis of Development, ed. W. D. McElroy and B. Glass. Baltimore: Johns Hopkins Press. 1959 With L. V. Moorhead and T. F. Pulis. Juvenile hormone in thymus, human placenta and other mammalian organs. Nature 183:405. The juvenile hormone. I. Endocrine activity of the corpora allata of the adult Cecropia silkworm. Biol. Bull. 116:323-38. 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 proteins of the Cecropia silkworm. [. Insect Physiol. 5:61-72. 1961 The juvenile hormone. II. Its role in the endocrine control of molt- ing, pupation, and adult development in the Cecropia silkworm. Biol. Bull. 121:572-85. 1963 The juvenile hormone. III. Its accumulation and storage in the ab- domen of certain male moths. Biol. Bull. 124:355-67. Differentiation and morphogenesis in insects. In The Nature of Bio- logical Diversity, ed. I. M. Allen, pp. 243-60. New York: McGraw- Hill. 1964 With B. Bowers. Physiology of insect diapause. XIII. DNA synthesis during the metamorphosis of the Cecropia silkworm. Biol. Bull. 126:205-19. With F. C. Kafatos. Enzymatic mechanism for the escape of certain moths from their cocoons. Science 146:538-40. With P. L. Adkisson. Physiology of insect diapause. XIV. An endo- crine mechanism for the photoperiodic control of pupal diapause in the oak silkworm, Antherea pernyi. Biol. Bull. 127:511-25. 1965 With R. A. Lockshin. Programmed cell death. I. Cytology of degen- eration in the intersegmental muscles of the Pernyi silkworm. III. Neural control of the breakdown of the intersegmental muscles of silkworms. V. Cytolytic enzymes in relation to the breakdown of the intersegmental muscles of silkmoths. J. Insect Physiol. 11:123- 34; 11:601-10; 11:831-44. 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- ment. Proc. Natl. Acad. Sci. U.S.A. 53:770-77. 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. 1967 With L. M. Riddiford. The effects of juvenile hormone analogues on the embryonic development of silkworms. Proc. Natl. Acad. Sci. U.S.A. 57:595-601. With L. M. Riddiford. Volatile principle in oak leaves: role in sex life of the polyphemus moth. Science 155:589-90. With L. M. Riddiford. Chemical signaling between polyphemus moths and between moths and host plant. Science 156:541. Third-generation pesticides. Sci. Am. 217:13-17. 1968 Ecdysone and ecdysone analogues: their assay and action on dia- pausing pupae of the Cynthia silkworm. Biol. Bull. 134:344-55. With T. Ohtaki and R. D. Milkman. Dynamics of ecdysone secretion and action in the fleshily Sarcophaga peregrina. Biol. Bull. 135:326- 33. 1969 Photoperiodism and the endocrine aspects of insect diapause. In

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CARROLL MILTON WILLIAMS 431 Dormancy and Survival, ed. H. W. Woolhouse, pp. 285-300. Cam- bridge: Cambridge University Press. Nervous and hormonal communication ire insect development. Dev. Biol. (Suppl.) 3:133-50. 1971 With L. M. Riddiford. Role of the corpora allata in the behavior of saturnid moths. I. Release of sex pheromone. Biol. Bull. 140:1-7. With M. P. Kambysellis. In vitro development of insect tissues. I. A macromolecular factor prerequisite for silkworm spermatogen- esis. Biol. Bull. 141:527-40. With M. P. Kambysellis. In vitro development of insect tissues. II. The role of ecdysone in the spermatogenesis of silkworms. Biol. Bull. 141:541-52. 1972 With F. C. Kafatos. Theoretical aspects of the action of juvenile hormone. In Insect Juvenile Hormones, ed. l. l. Menn and M. Beroza, pp. 29-41. New York: Academic Press. 1974 With H. F. Nijhout. Control of molting and metamorphosis in the tobacco hornworm, Manduca sexta. I. Growth of the last instar larva and the decision to pupate. J. Exp. Biol. 61:481-92. With H. F. Nijhout. Control of molting and metamorphosis in the tobacco hornworm, Manduca sexta. II. Cessation of juvenile hor- mone secretion as a trigger for pupation. [. Exp. Biol. 61:493-502. 1976 Juvenile hormone . . . In retrospect and in prospect. In The Juvenile Hormones, ed. L. I. Gilbert, pp. 1-14. New York: Plenum Press. 1977 With P. and L. Cherbas. Induction of acetylcholine esterase activity by 13-ecdysone in a Drosophila cell line. Science 197:275-77. 1979 How basic studies on insects have helped man. In The Biological

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432 BIOGRAPHICAL MEMOIRS Revolution: Applications of Cell Biology to Public Welfare, ed. G. Weissmann, pp. 65-78. New York: Plenum Press. 1980 With L. Safranek. Studies of the prothoracicotropic hormone in the tobacco hornworm, Manduca sexta. Biol. Bull. 158:141-53. With L. Cherbas, C. D. Yonge, and P. Cherbas. The morphological response of Kc-H cells to ecdysteroids: hormonal specificity. Roux's Archives Dev. Biol. 189: 1 -15. 1981 With G. M. Carrow and R. L. Calabrese. Spontaneous and evoked release of prothoracicotropin from multiple neurohemal organs of the tobacco hornworm. Proc. Natl. Acad. Sci. U.S.A. 78:5866-70. 1984 With G. M. Carrow and R. L. Calabrese. Architecture and physiol- ogy of insect cerebral neurosecretory cells. J. Neuroscience 4:1034- 44. 1986 With L. Safranek and C. R. Squire. Precocious termination of dia- pause in neck- and abdomen-ligated pupal preparations of the tobacco hornworm, Manduca sexta. Biol. Bull. 171:126-34. s

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