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Below are the first 10 and last 10 pages of uncorrected machine-read text (when available) of this chapter, followed by the top 30 algorithmically extracted key phrases from the chapter as a whole.
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OCR for page 352
1941
OCR for page 353
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
OCR for page 357
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.
OCR for page 358
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 pigs—swine 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
OCR for page 359
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
OCR for page 361
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-
OCR for page 362
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.
OCR for page 366
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-
OCR for page 367
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.
OCR for page 368
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
Rinderpest—I-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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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.
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1955
An infectious fibroma of deer. Proc. Soc. Exp. Biol. Med., 88:533-35.
With L. G. MacNamara and Robert Mangold. Epizootic hemor-
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The swine lungworm as a reservoir and intermediate host for swine
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With David Bodian et al. Interim Report, Public Health Service
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With David Bodian et al. The monkey safety test for poliomyelitis
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With Michael B. Shimkin. Some observations on cancer research in
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The leech as a potential virus reservoir. J. Exp. Med., 105:373-82.
With Peyton Rous. Presentation of the Kober Medal to Richard
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Influenza: history, epidemiology, and speculation. (The R. E. Dyer
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Dedication Speech, Medical Research Center, University of Iowa.
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Pseudorabies (Aujeszky's Disease, mad itch, infectious bulbar paraly-
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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-
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With Dan H. Moore, Robert S. Stone, and Dorothy Gelber. Ultra-
structure and site of formation of rabbit papilloma virus. Proc.
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With Robert S. Stone and Dan H. Moore. Electron microscope study
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With Norma E. Mettler and Lester G. MacNamara. The propaga-
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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.
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Porcine contagious pleuropneumonia. I. Experimental transmission,
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With David C. White and Grace Leidy. Porcine contagious pleuro-
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The epidemiology of the origin and perpetuation of a new disease.
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With David C. White, Grace Leidy, and James D. iamieson. Porcine
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An antiviral substance from Penicillium funiculosum. IV. Inquiry
into the mechanism by which Helenine exerts its antiviral effect.
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With Ping-Yao Cheng. The presence in Penicillium funiculosum of
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Evolutionary episodes in the concept of viral oncogenesis. (Philip B.
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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.
Representative terms from entire chapter:
richard edwin