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1941

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RICHARD EDWIN SHOPE December 25, Z901-October 2, 1966 BY CHRISTOPHER ANDREWES IN THESE DAYS of fashions in research and dependence upon sophisticated equipment, it is refreshing to know of people like Dick Shope. He was a born naturalist: he found his own problems in the field and sought their solutions there and in the laboratory, using simple techniques. He talked to farmers and veterinarians in his native Iowa and learned from what they had to tell him. In the laboratory he usually worked alone, doing essential things, including postmortems of pigs, with his own hands. He was born on Christmas Day 1901 in Des Moines, Iowa. His father was a prominent physician there, and from him and his mother he inherited genes of German, Scottish, English, Pennsylvania Dutch, and Indian origin. He enjoyed an open-air life with hunting and fishing on holidays. From ten onward he earned money by milking cows and looking after farm stock, especially poultry. At seventeen he went to Ames to register in the School of Forestry, but as the registrar's office was not open, he proceeded to Iowa City and registered as a pre-medical student. At medical school he did well both in his studies and in sports, qualifying in 1924. Thus his college education in medicine and his boyhood experiences on the farm combined to produce a man excellently qualified to contribute to knowledge of animal diseases. 353

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354 BIOGRAPHICAL MEMOIRS After qualifying, he became an instructor in pharmacology at the University of Iowa and among other activities did work on the chemotherapy of tuberculosis. Because of what he did in that field he was invited to join the laboratories of the Rocke- feller Institute at Princeton to work under Dr. Paul Lewis. At this time he married a fellow student, Helen Ellis, and the first few years of their marriage were, financially, difficult ones. Nor was the work on tuberculosis particularly rewarding. In 1928, however, he left the field of tuberculosis to work on hog cholera and thus began a career in the field of virology that was to continue for thirty-eight years. While investigating hog cholera in the field, Shope saw his first outbreak of swine influ- enza, and he proceeded to study this, with Paul Lewis, in 1929. They soon isolated a bacterium, Haemophilus inQuenzoe suds, similar to Pfeiffer's bacillus, at one time thought to be the cause of influenza in man. The Haemophilus was regularly present in the bronchial secretions of infected pigs, but cultures failed to reproduce the disease. Shope wrote, "We were at this stage of the game, in almost the identical predicament regarding the role of H. in- puenzue suds that investigators of human influenza had been in regarding the Pfeiffer bacillus at the close of the 1918 pan- demic."~ At this time Paul Lewis died of a yellow fever infection con- tracted in the laboratory, and Shope carried on by himself. He now made sterile filtrates of infectious material and adminis- tered them to swine intranasally. The filtrates did indeed pro- duce symptoms but nothing as severe as real swine flu. A few had fever, some coughed, and most had apathy and loss of appe- tite. Leukopenia was regularly present. Sacrificed animals had changes in the lungs, but whereas swine flu victims showed ex- tensive collapse, bronchitis, and bronchiectasis, those suffering ~ Ricketts Lecture, 1964. See Bibliography.

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RICHARD EDWIN SHOPE 355 from what Shope now called "filtrate disease" had minimal changes. It seemed likely, however, that a virus was concerned. Shope later wrote, "Instead of one agent . . . of possible etiologi- cal importance, here were two such agents." Now, to quote G. W. Corner, "Acting upon an improbable conjecture, Shope administered the bacillus and the virus at the same time, where- upon the animals amazingly came (town with typical influenza characterized by severe pneumonia."! Shope naturally wondered whether his findings had any bearing on the causation of influenza in man. He had not Tong to wait; in 1933, two years after he had described his findings, Wilson Smith, Patrick P. Laidlaw and I reported that a virus from human influenza would infect ferrets. I then visited- Prince- ton and compared notes with Shope, thus beginning a very close friendship that endured until his death. Over the years we ex- changed many long, highly controversial, and often hilarious letters about all aspects of influenza and many other subjects. After 1934 there followed a period of consolidation in the swine flu work. Recovered pigs were found to develop neutral- izing antibodies and to be immune to reinfection. The disease was found to pass readily from pig to pig by contact, but curi- ously enough, though the virus and Haemophilus would pass together from a swine flu-infected pig, only the virus was trans- mitted in subsequent serial contacts. Then, only the filtrate disease appeared unless the contact pig happened to be carrying the Haemophilus. It was soon shown that swine flu virus, like the human one, would infect ferrets, and the further, important fact emerged that when it was given intranasally to anesthetized ferrets, they developed pneumonia instead of only nasal symptoms. Patho- # Ricketts Lecture, 1964. See Bibliography. t A History of the Rockefeller Institute 1901-1953 t`N.Y.: Rockefeller Inst. Press, 1964).

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356 BIOGRAPHICAL MEMOIRS genicity of swine flu for mice was also established. Tests on both sides of the Atlantic showed that the swine and human viruses were antigenically related, though not identical. In cross- immunity experiments, only partial protection was produced by the heterologous virus. Swine were found to be susceptible to infection by the hu- man virus; this gave rise to filtrate disease unless the Haemo- philus was present also. During 1937, sera were obtained from pigs at farms near two institutions where flu outbreaks were in progress: the results showed that the pigs too had become in- fected with the human viruses, though no adverse symptoms among them had been observed. These observations suggested that swine influenza, which had first been observed in the Mid- west in 1918, might have originated from the transmission of the pandemic virus from man to pig. This idea was put forward independently by Laidlaw in Britain and by Shope. Remark- able confirmation came from studies of antibodies in people of different ages. Shope found, in 1936, that hardly any human sera from children aged twelve or less would neutralize the swine flu virus while many from older persons did so. This suggested that a virus antigenically related to swine flu had been present in the human population up to 1924 but not later. (One may reasonably suppose that the virus responsible for the 1918- 1919 pandemic persisted for a few years after that catastrophe.) Work in several laboratories has confirmed these suggestions, and the relation of swine flu virus to the pandemic strain is generally accepted as being highly probable. The discovery of the swine influenza virus and the bearing of the findings on human disease remain Shope's greatest con- tribution to knowledge. The next phase of his work, beginning in 1941, is much more controversial. His observations in the field had taught him that the disease was commonly absent during the summer but might break out explosively in October and November. Moreover this might happen, perhaps after the

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RICHARD EDWIN SHOPE 357 onset of inclement weather, in several farms simultaneously. There was no question of direct transmission of virus from one farm to another; it seemed rather that virus had been seeded into the herds beforehand and then activated in many pigs at the same time. After pursuing other clues which proved unre- warding, Shope concluded that swine lung worms were acting as intermediate hosts. Ova laid by these worms are passed in the pigs' feces and taken up by earthworms in which the eggs hatch and undergo further stages in development. Pigs are fond of eating earthworms; the lungworm larvae are thus ingested and eventually reach the pigs' lungs, thus completing the cycle. Shope concluded that lungworms from flu-infected pigs would carry the virus in an inapparent or "masked" state throughout this cycle. On regaining a position in the lungs of fresh swine, the masked virus would not have its pathogenic properties re- stored until some stress to the pig had triggered something off; only after that did respiratory illness result. Shope had no dif- ficultv in infesting earthworms with lungworms from flu-in- fected swine and in passing them back to fresh animals, which he called "prepared" swine. Disease was most readily provoked by repeated injections of these swine with cultures of Haemo- philus: it could also be activated by exposure of the pigs to hard weather. Unfortunately the outcome of the experiments was irregular; success was obtained in only about half the attempts. Moreover, virus could never be demonstrated in lungworms by direct tests. Opinions are much divided as to the validity of Shope's explanation of the facts. Some have accepted it as gospel, others have been wholly skeptical. The technique to test its truth has been beyond the reach of most workers: so few have attempted it. Those who have, have met varying success. I myself was un- successful: Shope gave me infected earthworms to take back to England, but there the attempted provocation did not lead to any disease.

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358 BIOGRAPHICAL MEMOIRS Shope was wont to argue that had it not been for the fortu- nate existence of a suitable, available, intermediate host, swine flu could never have persisted in North America. On the other hand, I used to argue with him that neither the earthworm with which we were most concerned, a species of AlZolobo~phora, nor the pig were native American animals and that I could not believe that a complex biological cycle involving three species could be evolved almost overnight. One can now look at influenza in better perspective. It is now known that strains of influenza A virus infect man, pigs, horses, several other mammals, and many species of wild and domesticated birds. Only among swine in North America is there a suggestion that a complex cycle involving worms is con- cerned. Elsewhere, and particularly in man, outbreaks of influ- enza start mysteriously and explosively: there is, in general, no possibility that a cycle in worms plays any part. One must, I think, conclude that though swine flu virus may well persist in lungworms and earthworms in North America, it probably does so passively and is not of as great epidemiological importance as Shope supposed. In 1930 Shope's attention was drawn to "mad itch," a violent, distressing, and fatal disease of cattle in the Midwest. He showed that it was caused by a virus transmissible to rabbits, and that it was endemic among pigs, in which it was comparatively harm- less. Cattle contracted infection through contact with pigs. He finally proved the identity of mad itch with pseudorabies, a disease prevalent in parts of Europe. Later he studied another disease of pigsswine pox and showed that it could be, though it was not necessarily, transmitted through the agency of pig- lice. He also published evidence that hog cholera virus might persist, as swine flu virus appeared to do, in lungworms. Shope's three most outstanding discoveries followed each other in rapid succession: swine influenza in 1931, the rabbit fibroma in 1932, and the rabbit papilloma in 1933. The infectious fibroma, often referred to as the Shope

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RICHARD EDWIN SHOPE 359 fibroma, was discovered on a shot cottontail rabbit (Sylvilagus). Minced material from this was inoculated into domestic rabbits (OryctoZagus) and readily produced growths, especially in young animals. Inoculation into the testis was most successful; intra- dermal and intraperitoneal injections gave less constant results. The growths consisted of proliferating fibroblasts; in the over- lying epidermis eosinophilic granules were seen, though only in the cottontails. Shope emphasized that this was a tumor only in the broadest sense of "a local swelling consisting of a mass of new tissue." This was wise, since the tumors normally re- gressed. One persisted as long as seventy-seven days in a cotton- tail, but regressions occurred much earlier in domestic rabbits. It was soon shown that the growths had a filterable cause since an infectious agent passed a Berkefeld V filter. There was, however, no evidence of spread by contact. When infection was transferred, it was evident that the host's cells were being in- fected: it was not a question of transplanting a graft. Recovered rabbits were immune to further infection and developed neu- tralizing antibodies in their sera. The character of the infection suggested to Shope a possible relationship to rabbit myxoma. This infection, of South Ameri- can origin, causes local lesions in the native rabbits, another Sylvilagus species, but in domestic rabbits causes fatal disease. Shope found that rabbits recovered from his fibroma were largely resistant to the myxoma virus: they still developed local lesions, but almost all survived. Some measure of cross-immunity was also apparent in tests with antisera against the two viruses. The fibroma virus has been used subsequently as a practical method of immunizing rabbits against myxomatosis. By analogy with myxomatosis it seems likely that the fibroma is mechanically transmitted by insect bites, but Shope's investi- gation of this in the field was never completed. In 1936 Shope sent me some fibroma material, which I duly Journal of Experimental Medicine, 5~6(1932):793. See Bibliography.

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360 BIOGRAPHICAL MEMOIRS inoculated into rabbits' testes. Most surprisingly there appeared only acute inflammatory lesions instead of the expected pro- liferative changes. Moreover, many inoculated rabbits devel- oped generalized pock-like lesions on their skins. This "inflam- matory" strain was antigenically identical to the original one. Shope noted similar changes in one of his strains, and we col- laborated in work that seemed to indicate that a mutation had occurred in the direction of greater virulence for rabbit cells. Another development of work on the fibroma was the finding in several laboratories that various factors could cause the benign self-limited fibroma to become a generalized, persistent, or even fatal infection. These factors included the simultaneous injection of carcinogens or cortisone, application of X-rays, or the use of very young rabbits. The importance of work on the fibroma is its demonstration that the distinction between infec- tion and neoplasia may be largely artificial: Shope's fibroma is one agent which bridges the gap. Still more important in this connection is Shope's rabbit papilloma. Cottontail rabbits shot in Iowa and Kansas frequently have horns or warts on their skins. Shope found that material from these would readily produce warts on the skins of cotton- tail or tame rabbits when rubbed into the shaved and lightly scarified skin. The warts usually began to appear after six to twelve days: they might regress after a time or persist indefi- nitely as tall, often black, horns. The warts proved to be caused by a virus that gave rise to neutralizing antibodies: recovered animals were immune to reinfection. In cottontails the warts could be passed in series without difficulty, but the warts in domestic rabbits, even though well-developed, commonly failed to be transmitted to further animals. Shope devoted much at- tention to this matter: he did occasionally obtain successful serial transmissions in the tame rabbits, but failures were the rule. The important discovery was then made that the tame rabbit warts, though apparently virus-free, would lead to the production of specific neutralizing antibodies when injected

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RICHARD EDWIN SHOPE 361 into rabbits intraperitoneally. The virus, or an essential part of it, was still there, perhaps in a masked form. This brought out the point that a virus might be the cause of a neoplastic condition, yet not be directly demonstrable. Critics of the work maintained that the difference between the wild and tame rab- bits' warts was a purely quantitative one, but further work ren- dered this unlikely. Though many may be reluctant to believe in the "masked" swine flu virus in lung-worms, the "masked" papilloma virus seems to be genuine. Shope suggested that it might survive as infectious DNA, but since it gives rise to neutralizing antibodies, there must be more of it remaining than that. The work gained a new dimension when it was found that in many tame rabbits the warts progressed and became carcino- matous. This change, though common in domestic rabbits, was rare in cottontails. Shope, at this time, was busy with many problems, so he generously gave the material to Francis Peyton Rous. What Rous did with the rabbit cancers during the next thirty years is a matter of history. When the war came, there were fears that the enemy might seek to interfere with food production in North America by introducing the very infectious disease rinderpest into American cattle. Shope, attached for this purpose to the Army, was asked to take charge of a joint United States-Canadian project to produce an effective vaccine. Stringent precautions were neces- sary to prevent the escape of the infection, and laboratories were accordingly set up on Grosse Isle, a small island in the St. Law- rence River below the city of Quebec. Here Shope, with a staff of five other scientists, worked in strict isolation, and in the course of nineteen months produced an effective vaccine by growing and attenuating the virus in hens' eggs. This has since been used on a large scale in the field. With this work completed, Shope asked to be transferred back to the Navy, which had been his original wartime assign- ment. T. M. Rivers was in charge of a U.S. Naval Medical Re-

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362 BIOGRAPHICAL MEMOIRS search unit and, in 1944, got together a powerful team including Shope. There was little knowledge of what medical dangers might threaten men attacking islands in the Far East. When the invasion of Okinawa was planned, Shope was a member of a medical team that landed with the assault party in April 1945. A laboratory was established and was at times under fire. Fortu- nately the disease hazards were not found to be great, and before long the group was ordered to return to Guam. Even under war conditions Shope's inquiring mind sought fresh opportunities. He collected in the Pacific a number of molds, hoping to find one that would be of chemotherapeutic value. One of them, from Guam, grew on the cover of a photograph of his wife, Helen, and later this yielded an extract, which he called Hele- nine, having activity against several viruses in vivo. It was proved later that this was due to a nucleoprotein in it capable of stimulating interferon production. Soon after the war, in 1947, fell a severe blow. Shope, essen- tially a country lover, had enjoyed being able to live near Princeton University on a small farm where he could keep a cow and poultry and grow vegetables. Then, with no previous warning to the staff, the Trustees of the Rockefeller Institute decided to close down their Princeton branch, offering the staff the opportunity to work in their main Institute in New York. Shope hated the idea of having to work in a city, especially as he needed facilities for work with large animals. So in 1949 he resigned and accepted a position as Assistant Director of the Merck Institute for Therapeutic Research in Rahway, New Jersey. Here he continued his work, chiefly on Helenine. But work within a commercial organization was quite foreign to his temperament, and in 1952 he returned to the Rockefeller Institute (now the Rockefeller University) in New York, living in an apartment across the street from the Institute but going back whenever possible for a weekend at his "gentleman's farm" near Princeton.

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RICH ARD ED WIN SH OPE 1929 365 Cholesterol in the blood of the horseshoe crab (Limulus polyphe- mus) and the woolly bear caterpillar (Asia isabella). Proc. Soc. Exp. Biol. Med., 26:336. With Paul A. Lewis. A paralytic disease of guinea pigs due to the tubercle bacillus. l. Exp. Med., 50:365-70. With Paul A. Lewis. The cultural and staining reactions of a strain of the tubercle bacillus producing paralysis in guinea pigs. J. Exp. Med., 50:371-76. With Paul A. Lewis. The study of the cells of the blood as an aid to the diagnosis of hog cholera. l. Am. Vet. Med. Assoc., 74:145. With Paul A. Lewis. The blood in hog cholera. I. Exp. Med., 50: 719-37. 1930 Variations in the plasma cholesterol and cholesterol ester content in hog cholera. I. Exp. Med., 51: 179-87. "Mad itch" of cattle. Science, 72 (November 28~: 559. 1931 The etiology of swine influenza. Science, 73(February 20~:214-15. An experimental study of "mad itch" with especial reference to its relationship to pseudorabies. l. Exp. Med., 54:233~8. Swine influenza. I. Experimental transmission and pathology. l. Exp. Med., 54:349-59. With Paul A. Lewis. Swine influenza. II. A hemophilic bacillus from the respiratory tract of infected swine. l. Exp. Med., 54:361-71. Swine influenza. III. Filtration experiments and etiology. l. Exp. Med., 54:373-85. 1932 Studies on immunity to swine influenza. l. Exp. Med., 56:575-85. A transmissible tumor-like condition in rabbits. A filtrable virus causing a tumor-like condition in rabbits and its relationship to Virus myxomatosurn. J. Exp. Med., 56: 793-822. Identity of the viruses causing "mad itch" and pseudorabies. Proc. Soc. Exp. Biol. Med., 30:308-9.

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366 BIOGRAPHICAL MEMOIRS 1933 Modification of the pathogenicity of pseudorabies virus by animal passage. J. Exp. Med., 57: 925-31. Infectious papillomatosis of rabbits. T. Exp. Med., 58:607-24. 1934 Swine influenza. V. Studies on contagion. l. Exp. Med., 59:201-11. The infection of ferrets with swine influenza virus. l. Exp. Med., 60:49-61. Pseudorabies as a contagious disease in swine. Science, 80July): 102-3. 1935 Serial transmission of virus of infectious papillomatosis in domestic rabbits. Proc. Soc. Exp. Biol. Med., 32:830-32. Experiments on the epidemiology of pseudorabies. I. Mode of trans- mission of the disease in swine and their possible role in its spread to cattle. J. Exp. Med., 62:85-99. Experiments on the epidemiology of pseudorabies. II. Prevalence of the disease among middle western swine and the possible role of rats in herd-to-herd infections. l. Exp. Med., 62: 101-17. The infection of mice with swine influenza virus. T. Exp. Med., 62: 561-72. With Marion L. Orcutt. The distribution of swine influenza virus in swine. J. Exp. Med., 62:823-26. 1936 Infectious fibroma of rabbits. III. The serial transmission of Virus myxomatosum in cottontail rabbits, and cross-immunity tests with the fibroma virus. l. Exp. Med., 63: 33-41. Infectious fibroma of rabbits. IV. The infection with Virus myxoma- tosum of rabbits recovered from fibroma. l. Exp. Med., 63:43-57. A change in rabbit fibroma virus suggesting mutation. II. Behavior of the variant virus in cottontail rabbits. l. Exp. Med., 63: 173-78. With C. H. Andrewes. A change in rabbit fibroma virus suggesting mutation. III. Interpretation of findings. J. Exp. Med., 63: 179-84. With Thomas Francis, in Neutralization tests with sera of conva-

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RICHARD EDWIN SHOPE 367 lescent or immunized animals and the viruses of swine and hu- man influenza. I. Exp. Med., 63:645-53. The incidence of neutralizing antibodies of swine influenza virus in the sera of human beings of different ages. J. Exp. Med., 63:669- 84. Immunization experiments with swine influenza virus. J. Exp. Med., 64:47-61. With Thomas Francis, fir. The susceptibility of swine to the virus of human influenza. J. Exp. Med., 64: 791-801. The influenzas of swine and man. In: The Harvey Lectures, 1935- 1936, p p. 183-213. N.Y.: The Harvey Society. 1937 Immunization of rabbits to infectious papillomatosis. J. Exp. Med., 65:219-31. Recent knowledge concerning influenza. Ann. Intern. Med., 11 (no. 1~:1-12. The effect of Hemophilus influenzoe suds vaccines on swine influ- enza. l. Exp. Med., 66:169-75. Immunological relationship between the swine and human influenza viruses in swine. i. Exp. Med., 66: 151-68. 1938 Protection of rabbits against naturally acquired infectious myxoma- tosis by previous infection with fibroma virus. Proc. Soc. Exp. Biol. Med., 38:86-89. Serological evidence for the occurrence of infection with human influenza virus in swine. .T. Exp. Med., 67:739~8. 1939 With Carlos T. Rosenbusch. The antibody response to swine influ- enza. l. Exp. Med., 69:499-505. An intermediate host for the swine influenza virus. Science, 89:441- 42. Serological studies of swine influenza viruses. I. Exp. Med., 69:847- 56. Complex infections. (The Middleton Goldsmith Lecture, read before the New York Pathological Society, Dec. 7, 1938.) Arch. Pathol., 913-32.

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368 BIOGRAPHICAL MEMOIRS 1940 Swine pox. Arch. Gesamte Virusforsch., 1:457-67. 1941 Problems in the epidemiology of virus diseases. (Illinois Conference on Public Health, Springfield, Dec. 6, 1940.) The Illinois Health Messenger, 13: 30-35. The influence of host and intermediate reservoir host in determining the epidemiologic pattern of bovine pseudorabies and swine in- fluenza. In: Problems and Trends in Virus Research, University of Pennsylvania, Bicentennial Conference, pp. 55-66. Phila- delphia: Univ. of Pennsylvania Press. The swine lungworm as a reservoir and intermediate host for swine influenza virus. I. The presence of swine influenza virus in healthy and susceptible pigs. J. Exp. Med., 74:41~7. The swine lungworm as a reservoir and intermediate host for swine influenza virus. II. The transmission of swine influenza virus by the swine lungworm. J. Exp. Med., 74:49-68. 1943 Swine influenza. In: Virus Diseases: the Messenger Lectures, 1942, pp. 85-109. Ithaca: Cornell Univ. Press. The swine lungworm as a reservoir and intermediate host for swine influenza virus. III. Factors influencing transmission of the virus and the provocation of influenza. l. Exp. Med., 77: 111-26. The swine lungworm as a reservoir and intermediate host for swine influenza virus. IV. The demonstration of masked swine influ- enza virus in lungworm larvae and swine under natural condi- tions. l. Exp. Med., 77: 127-38. 1944 Old, intermediate, and contemporary contributions to our knowl- edge of pandemic influenza. Medicine, 23:415-55. 1946 RinderpestI-XVI. Am. J. Vet. Res., 7: 133-237. The problem of certain animal diseases prevalent in Iowa as related to human medicine. l. Iowa State Med. Soc., 36:419-25.

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RICHARD EDWIN SHOPE 1948 369 An unfamiliar mechanism of disease transmission. Proc. Am. Phil. Soc., 92:289-93. Propagation of the virus of cattle plague in the developing egg. Proc. 4th Internat. Cong. Trop. Med. and Malaria, Washington, 1948. Wash., D.C.: U.S. Govt. Printing Once, 2: 1351-57. With F. R. Selbie and R. H. M. Robinson. Shope papilloma virus: Reversion of adaptation to domestic rabbit by passage through cottontail. Brit. I. Cancer, 2: 375-80. 1950 "Masking," transformation, and interepidemic survival of animal viruses. In: Viruses of 1950, pp. 79-92. Pasadena: Division of Biology of the California Institute of Technology. The spread of viruses from infected to susceptible hosts. In: The Pathogenesis and Pathology of Viral Diseases, New York Acad- emy of Medicine, Section on Microbiology, Symposium Number Three, pp. 6-18. N.Y.: Columbia Univ. Press. 1951 Factors involved in the transmission and propagation of viruses. Cornell Vet., 41: 181-89. The provocation of masked swine influenza virus by infection with human influenza virus. Tschr. diergeneesk, 76:414-20. 1952 Swine and human health. Proc. Book Am. Vet. Med. Assoc. (89th Ann. Meet., 1952), 97:381-84. Role of swine diseases in diseases of man. Pub. Health Rep., 67:983- 84. With Oscar Sussman and Ralph A. Hendershott. Administrative considerations of garbage feeding with reference to vesicular exanthema and trichinosis. United States Livestock Sanitary As- sociation (56th Annual Meeting, 1952), 218-22. 1953 An antiviral substance from Penicillium funiculosum. I. Effect upon infection in mice with swine influenza virus and Columbia SK encephaIomyelitis virus. l. Exp. Med., 97:601-25.

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370 BIOGRAPHICAL MEMOIRS An antiviral substance from Penicillium funiculosum. II. Effect of Helenine upon infection in mice with Semliki Forest virus. J. Exp. Med., 97:627-38. An antiviral substance from Penicillium funiculosum. III. General properties and characteristics of Helenine. l. Exp. Med., 97:639- 50. Public health regulations concerning brucellosis. Public Health News, New jersey State Department of Health, 34:183-88. 1954 Ecology and virus reservoirs. In: The Dynamics of Virus and Rickettsial Infections, pp. 125-41. N.Y.: Blakiston. Biographical memoir of Raymond Alexander Kelser. In: Biographi- cal Memoirs, 28:199-221. N.Y.: Columbia Univ. Press for the National Academy of Sciences. Infectious ectromelia of mice (mouse pox). l. Lab. Clin. Med., 44: 333-35. Also in: J. Nat. Cancer Inst., 15:405-8. Foreword. In: M. N. Dreguss and L. S. Lombard, Experimental Studies in Equine Infectious Anemia, pp. vii-viii. Philadelphia: Univ. of Pennsylvania Press. 1955 An infectious fibroma of deer. Proc. Soc. Exp. Biol. Med., 88:533-35. With L. G. MacNamara and Robert Mangold. Epizootic hemor- rhagic disease of deer. New Jersey Outdoors, State of New Jersey Division of Fish and Game, 6(no. 5~:17-21. The swine lungworm as a reservoir and intermediate host for swine influenza virus. V. Provocation of swine influenza by exposure of prepared swine to adverse weather. J. Exp. Med., 102:567-72. With David Bodian et al. Interim Report, Public Health Service Technical Committee on Poliomyelitis Vaccine. J. Am. Med. Assoc., 159: 1444 47. Epizootiology of virus diseases. In: Advances in Veterinary Science, 2: 1-46. N.Y.: Academic Press. 1956 Swine influenza. In: U.S. Department of Agriculture Yearbook, 1956, pp. 366-68. Washington, D.C.: U.S. Govt. Printing Once. With Colin M. MacLeod et al. The United States medical mission

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RICHARD EDWIN SHOPE 371 on microbiology and epidemiology to the Soviet Union. J. Am. Med. Assoc., 162: 656-57. With David Bodian et al. The monkey safety test for poliomyelitis vaccine. Am. J. Hyg., 64: 104-37. With Michael B. Shimkin. Some observations on cancer research in the Soviet Union. Cancer Res., 16:915-17. 1957 The leech as a potential virus reservoir. J. Exp. Med., 105:373-82. With Peyton Rous. Presentation of the Kober Medal to Richard Shope. Acceptance of the Kober Medal for 1957. Trans. Assoc. Am. Physicians, 70:29-40. Discussion: A. B. Sabin. Properties of attenuated polioviruses and their behavior in human beings. In: Cellular Biology Nucleic A cids and viruses' 5: 1 39~0. N.Y.: New York Academy of Sciences. The provocation of masked hog cholera virus in lungworm-related swine by ascaris larvae. National Academy of Sciences Autumn Meeting, Nov. 18-20, 1957. Science, 126: 1236. 1958 Influenza: history, epidemiology, and speculation. (The R. E. Dyer Lecture.) Public Health Reports, 73(no. 2~:165-78. Dedication Speech, Medical Research Center, University of Iowa. Medical Bulletin (State Universiy of Iowa, Iowa City), Winter 1957-58: 33-40. The swine lungworm as a reservoir and intermediate host for hog cholera virus. I. The provocation of masked hog cholera virus in lungworm-infested swine by ascaris larvae. J. Exp. Med., 107: 609-22. Foreword. In: F. R. Beaudette, Progress in Psittacosis Research and Control, pp. v-vii. New Brunswick, Ad.: Rutgers Univ. Press. Virus reservoirs. In: Progress in Psittacosis Research and Control, 19-23. New Brunswick, N.~.: Rutgers Univ. Press. The swine lungworm as a reservoir and intermediate host for hog cholera virus. II. Attempts to demonstrate the presence of hog cholera virus in Iungworms derived from swine with cholera. J. Exp. Med., 108: 159-69.

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372 BIOGRAPHICAL MEMOIRS Swine influenza (flu, hog flu, swine flu). In: Diseases of Swine, pp. 81-98. Ames Iowa State College Press. Pseudorabies (Aujeszky's Disease, mad itch, infectious bulbar paraly- sis>. In: Diseases of Swine, pp. 219-28. Ames Iowa State College Press. With Robert Mangold, Lester G. MacNamara, and Keith R. Dumbell. An infectious cutaneous fibroma of the Virginia white- tailed deer (Odocoileus virginianus). J. Exp. Med., 108:797-802. The role of latency in the epidemiology of virus diseases. In: Com- parative Medicine in Transition, Proceedings of the First Insti- tute of Veterinary Public Health Practice, Oct. 6-9, 1958. The Univ. of Michigan School of Public Health, Ann Arbor. Lord Baltimore Press. 1959 Hog cholera eradication, an editorial. l. Am. Vet. Med. Assoc., 134 (Feb. 1, 1959~:143-45. Latent virus infections in animals. Recent Progress in Microbiology, Symposia Held at VII Intern. Congr. for Microbiology, 1959, 230-37. With Dan H. Moore, Robert S. Stone, and Dorothy Gelber. Ultra- structure and site of formation of rabbit papilloma virus. Proc. Soc. Exp. Biol. Med., 101:575-78. With Robert S. Stone and Dan H. Moore. Electron microscope study of the development of the papilloma virus in the skin of the rabbit. J. Exp. Med., 110:543~6. The story on E. E. E.: Eastern Equine Encephalomyelitis. New Jersey Outdoors, State of New Tersev Division of Fish and Game. 10(no. 5~:12-15. Comparative medicine. Occasional Paper No. 7. N.Y.: The Rocke- feller Institute Press. The roles of virus and host in determining the host reaction to the fibroma-myxoma virus complex. In: Genetics and Cancer, Thir- teenth Annual Symposium on Fundamental Cancer Research at the Univ. of Texas M.D. Anderson Hospital and Tumor Insti- tute, Bertner Foundation Lecture, pp. 311-23. Austin: Univ. of Texas Press. \Vith Keith R. Dumbell, Robert Mangold, and L. G. MacNamara. J ~

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RICHARD EDWIN SHOPE 373 An infectious fibroma of the Virginia white-tailed deer (Odocoileus virginianus). Acta Unio Internat. Contra Cancrum, 15:799-800. 1960 With Lester G. MacNamara and Robert Mangold. A virus-induced epizootic hemorrhagic disease of the Virginia white-tailed deer (Odocoileus virginianus). J. Exp. Med., 111: 155-70. Eastern viral encephalomyelitis, an editorial. J. Am. Vet. Med. Assoc. 136(no. 7~:339~0. Summarization of symposium on Eastern encephalomyelitis. Phila- delphia Medicine, 56(no. 14~:431-34. CommentsSven Gard's detection of viruses by chemical and bio- Iogical means. Cancer Res., CO(no. 5~:742-43. Summary of informal discussionsA consideration of virus-host re- lationships in neoplasia at the level of the whole animal. (Sympo- sium sponsored by the American Cancer Society The Possible Role of Viruses in Cancer.) Cancer Res. 20:784-95. Koch's postulates and a viral cause of human cancer: guest editorial. Cancer Res., 20: 1119-20. 196 Contributions of animal disease to research. Public Health News, New jersey State Dept. of Health, 42(no. 1 ): 9-1 2. An immunizing substance for Semliki Forest virus in the livers of immune mice. I. The initial observation together with a con- sideration of the attending conditions. l. Exp. Med., 113:511-15. Consideration of the possible etiological role of viruses in cancer. I. Am. College Dentists, June: 128-41. 1962 Are animal tumor viruses always virus-like? I. Gen. Physiol., 45: 143- 54 (Supp. to March issue). With Norma E. Mettler and Lester G. MacNamara. The propaga- tion of the virus of epizootic hemorrhagic disease of deer in new- born mice and HeLa Cells. i. Exp. Med., 116:665-78. Viruses and cancer. Annals of Dentistry, 21:96-101.

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374 BIOGRAPHICAL MEMOIRS 1963 With Lester G. MacNamara and Norma E. Mettler. The attenuation of the virus of epizootic hemorrhagic disease of door by its serial passage in the brains of newborn mice. l. Exp. Med., 118:421-24. 1964 Porcine contagious pleuropneumonia. I. Experimental transmission, etiology, and pathology. J. Exp. Med., 119:357-68. With David C. White and Grace Leidy. Porcine contagious pleuro- pneumonia. II. Studies of the pathogenicity of the etiological agent, Hemophilus pleurotneumoniae. J. Exp. Med., 119:369-76. The epidemiology of the origin and perpetuation of a new disease. (Howard Taylor Ricketts Lecture presented May 31, 1963, at Univ. of Chicago Medical School.) Perspectives in Biol. and Med., 7(Spring): 263-78. With David C. White, Grace Leidy, and James D. iamieson. Porcine contagious pleuropneumonia. III. Interrelationship of Hemo- philus pleurotneumoniae to other species of Hemophilus: Nutri- tional, metabolic, transformation, and electron microscopy studies. J. Exp. Med., 120: 1-12. The birth of a new disease. In: Newcastle Disease Virus: An Evolv- ing Pathogen, ed. Robert Paul Hanson. Madison The Univ. of Wisconsin Press. 1965 Transmission of viruses and epidemiology of viral infections. In: Viral and Rickettesial Infections of Man, ed. Igor Tamm and Frank L. Horsfall, in Philadelphia: l. B. Lippincott. 1966 An antiviral substance from Penicillium funiculosum. IV. Inquiry into the mechanism by which Helenine exerts its antiviral effect. J. Exp. Med., 123:213-27. With Ping-Yao Cheng. The presence in Penicillium funiculosum of an inhibitor to the antiviral agent Helenine. l. Exp. Med., 123: 505-8. Evolutionary episodes in the concept of viral oncogenesis. (Philip B.

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RICHARD EDWIN SHOPE 375 Price Lecture presented June 3, 1965, at Univ. of Utah College of Medicine.) Perspectives in Biol. and Med., 9:258-74. With Michael W. Rytel and Edwin D. Kilbourne. An antiviral sub- stance from Penicillium funiculosum. V. Induction of interferon by Helenine. J. Exp. Med., 123:577-84. An antiviral substance from Penicillium funiculosum. VI. Preven- tion of the establishment of passive immunity to Semliki Forest virus infection in mice by Helenine. l. Exp. Med., 124:15-32. An antiviral substance from Penicillium funiculosum. VII. An at- tempt to determine whether the material responsible for the anti- passive immunity effect exhibited by mice injected with Helenine is an interferon. I. Exp. Med., 124:915-19.