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FRANK LAPPIN HORSFALL
December 74, 1906-February 19, 1971
BY GEORGE K. HIRST
JR.
FRANK L. HORSFALL, JR. was a clinician and a virologist whose
influential leadership came primarily through his perceptive
scientific experimentation, both in the laboratory and in the
clinic, and also through his vast administrative skill.
He was born December 14, 1906 in Seattle, Washington,
where he spent all his formative years until he was twenty-one.
His father, a native Vermonter, was a prominent surgeon who
maintained a large house on Capitol Hill, and Frank, the first of
four children, was a high-spirited youth whose interests led him
into a wide range of activities. By the time he entered high
school, he had decided to become an engineer, and he spent
afternoons and evenings with a friend rigging up the family
Victrola for radio reception. In the course of his four years at
high school, he participated actively in the student council, the
boys' athletic association, the glee club, and the radio press
association. He was valedictorian of his class.
During four years of college at the University of Washington
he lived in a fraternity house, and during the early part of this
experience he was uncharacteristically erratic in the pursuit of
his studies. It was during this period that he went out for crew,
a major sport at Washington, and made the junior varsity. One
of his outstanding characteristics was a tendency to throw him-
self headlong into every new activity, and it is reported that he
233
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234
BIOGRAPHICAL MEMOIRS
became a passionate oarsman, spending every moment of his
spare time rowing or hanging around the boathouse. However,
many years later, although he kept his oar on the wall, he com-
pletely disavowed any interest in rowing and for that matter in
all other forms of athletics, either as a participant or as a
spectator.
Midway through his college course he gave up the idea of
becoming an engineer and decided to enter the medical pro-
fession. This involved some heroic scrambling to complete the
course-work requirements, but in 1927 he followed in his
father's footsteps by entering McGill University, in Montreal,
Canada, and graduated in 1932, at which time he received the
Holmes Gold Medal for having attained the highest scholastic
record in his class.
After receiving his medical degree, he spent a year as a house
officer in pathology at the Peter Bent Brigham Hospital in
Boston. This was a common preliminary to a postgraduate edu-
cation in surgery. It was during this year that he discovered
that he was exquisitely sensitive to formaldehyde, a fact which
markedly influenced later career decisions.
He responded characteristically to this disability by embark-
ing at once on an extensive study of formaldehyde sensitivity in
experimental animals and coupled this with a long series of
experiments on himself. The result was two substantial papers
on the subject, in which for the first time Horsfall was the sole
or the senior author, and in them he acknowledged the assis-
tance and advice of both H. Zinsser and S. B. Wolbach. This was
the beginning of Horsfall's informal training in immunology,
training which was to continue after another year at the Rocke-
feller Institute.
He returned to Montreal for a year's internship in medicine
at the Royal Victoria Hospital and then signed up for a year in
surgery, but shortly after starting the latter position he found
that it was incompatible with his formaldehyde sensitivity, ant!
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FRAN K LAPPIN HORSFALL, JR.
235
he promptly resigned. Immediately thereafter, in the fall of
1934, he went to the Rockefeller Institute in New York City,
where he was to remain with only minor interruption for the
next twenty-five years. On giving up a surgical career, for which
he was eminently qualified, he showed no outward evidence of
regret and quickly plunged off in another direction, which
involved him in clinical research coupled with basic research.
Horsfall came to New York at a time when the Rockefeller
Institute Hospital was at a peak of its reputation in the academic
medical world. Although it was a very small unit, it had a very
big reputation for pioneering in medical research. The staff
members were both clinical and nonclinical, and there was a
well-developed mystique among them concerning the catalytic
effect which clinical contact had on basic research.
A majority of the young people there were like Horsfall in
that they came from purely medical backgrounds and were to
spend formative postdoctoral years in this very stimulating
environment learning, in an informal manner, the basic disci-
plines such as bacteriology, virology, immunology, and physi-
ology. Through these young people the Hospital provided
excellent medical care, and at the time Horsfall was added to
the staff the pneumonia service had nearly completed a large
and successful series of cases in which type I pneumonia had
been treated with specific antipneumococcal horse serum. These
results had aroused widespread interest, and such sera were
. .
coming Into general use.
In addition to the time spent on clinical responsibilities, all
of the younger staff members were engaged in some basic re-
search project. Those on the pneumonia service often worked
on model infections in mice. Among the older staff members
there were some who did no clinical work at all, and the most
outstanding of these was Oswald T. Avery. The exceptional
thing about the Avery school was that its immediate goals were
so great and still far removed from any clinical application. His
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BIOGRAPHICAL MEMOIRS
very solid intellectual outlook set the tone for the whole depart-
ment.
On entering this mixed atmosphere of clinical and academic
science, Horsfall clearly and consistently showed his marked
preference for those activities for which there appeared to be
obvious relevant application to clinical work. Horsfall's em-
phasis on medical application would be demonstrated again and
again throughout his career, even when he was not taking care
of patients. He tackled his clinical responsibilities with en-
thusiasm. He was an eager therapist, and there are apocryphal
reports about the strong restraints required to prevent him from
testing in patients the enzyme capable of splitting the type III
capsular carbohydrate, which had been developed by O. T.
Avery and Rene Dubos. He was also an enthusiastic advocate of
using type-specific serum in treating lobar pneumonia, and his
influence was an important factor in switching the routine treat-
ment at the Hospital from horse to rabbit antiserum.
This change in treatment was very successful and aroused
wide interest. A large number of cases were treated with rabbit
serum at the Hospital, and Horsfall and his colleagues pub-
lished several extensive papers describing the clinical results.
The use of this new method caught on generally. Commercial
drug houses began producing rabbit antiserum, not only against
types I and II, but also against a myriad of the rarer varieties.
Horsfall, working with Kenneth Goodner, accompanied this
clinical work with a tremendous burst of laboratory activity in
which they studied the comparative immunology of horse and
rabbit sera in detail. These sera were found to differ from each
other in a number of fundamental ways. In the space of a little
more than a year, Hors fall and his colleagues published some
twenty-three papers on these systems. Thirty or more immuno-
logical differences were highlighted, many of which were inter-
preted as suggesting distinct therapeutic advantages for the
rabbit antibody.
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FRANK LAPPIN HORSFALL, JR.
237
Horsfall remained! at the Hospital for three years and during
his last year was Chief Resident Physician, a position which
greatly increased his clinical responsibilities. Avery at the time
was in the midst of his fascinating and fundamental work on the
transforming principle. Horsfall was very familiar with all the
current details of that work, which had been formalized by
Avery into a series of personal lectures known as the Red Seal
records. Frank Horsfall frequently sat at Avery's feet adsorbing
the gospel, which he later was able to repeat ire minute detail
and often did so with great gusto. Yet he never did any research
with Avery, much as he admired his whole approach. Their
scientific style was entirely different, and Horsfall would have
had a most difficult time adjusting to Avery's idiosyncrasies. For
Horsfall, the transforming principle must have seemed far away
from the world of patients.
Although the Rockefeller Hospital was at this time a prime
source of candidates for academic positions in medical schools
throughout the country, positions within the Hospital itself
were at a premium. Nevertheless it became clear within a very
short time that Horsfall's future would be in the Rockefeller
Institute. It is difficult to assess the reasons for this early popu-
larity which lay partly in his ability to see problems clearly, to
apply himself to the problem with enormous energy, and then
finally to present his results in a way which seemed simultane-
ously to be both conservative and expansive, romantic yet con-
vincing. Thomas Rivers, who was a very influential person in
American medicine at that time, was extremely devoted to
Horsfall and played a large part later in promoting him as his
own successor at the Rockefeller Institute Hospital.
It came as no surprise when in 1937 Horsfall resigned from
the Hospital and accepted a staff position with the International
Health Division of the Rockefeller Foundation. The change in
employer was almost imperceptible. The principal Foundation
laboratories were in fact housed at the Rockefeller Institute,
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BIOGRAPHICAL MEMOIRS
and Foundation scientists were treated as part of the family.
Horsfall's daily contacts with staff members in the Hospital and
the Institute continued as before, including his exceptionally
close relationship with Rivers, a friendship which was now
further cemented by the transformation of Horsfall into a
· .
vlro OglSt.
The Foundation had made a reputation for itself in virology
through its pioneering work on yellow fever, and its leaders
were anxious to continue this record of preeminence through
an attack on the influenza problem. Horsfall became head of a
laboratory section, previously organized by Thomas Francis, Jr.,
and the time seemed really ripe for developing a prophylactic
agent against epidemic influenza. The virus of influenza had
been isolated only four years before, and the Foundation offered
Horsfall a very large technical staff, abundant laboratory space,
generous financial support, and a sizeable professional staff that
was equipped by experience to do large-scale fieldwork and to
tackle epidemic problems on an international scale.
It is somewhat ironic that Horsfall stepped out of the lobar
pneumonia scene just as it was undergoing drastic change. The
elaborate type-specific serotherapy routines which had been very
painstakingly developed in the mid-thirties were briskly and
permanently swept away with the advent of chemotherapy and
antibiotics. This had little to do with Horsfall's change of
employer at this time, but does reflect the sweeping pace of
medical advances. During his three years on the pneumonia
service, he hacl received excellent training in immunology.
Plunging into virology was to be a new experience, and he
quickly became a master of the fundamentals. Once again, he
threw himself wholeheartedly into a new activity, which was to
occupy his attention for several years.
Before leaving the Hospital, Horsfall married Norma Cam-
pagnari, who worked there as a nurse. It was a most successful
marriage. She was a quick, merry, and very lively person and a
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FRANK LAPPIN HORSFALL, JR.
239
gracious hostess. Horsfall's domestic life was always important
to him, and he devoted much time to it in spite of numerous
external demands. They had three children, and he was very
much attached to them all.
The Foundation provided a proper setting for an individual
of Horsfall's brilliant and expansive outlook. In addition to his
major projects, he was able to devote much of his time to some
very fundamental biological problems, and one of these areas
was quantitative biology. He became involved with and obvi-
ously enjoyed the design and execution of mechanical projects
like a low-temperature storage cabinet or a complicated ventila-
tion system for a single room containing a large number of
ferrets that had to be individually housed. The latter project was
so successful that it enabled him to conduct experiments on the
highly contagious distemper virus without danger of spontane-
. ~ .
Ous cross-~ntect~on.
The research was nearly all on influenza virus, and at that
time it was necessary to measure virus concentration by intra-
nasal inoculation of material into mice. To achieve any reason-
able sort of accuracy required the infection of large numbers of
animals, followed by individual isolation during the incubation
period. Nevertheless, Horsfall did very large, highly quantitative
experiments with influenza, in which he attempted to work out
the amount of antibody required to neutralize different amounts
of virus in the inoculum. This was the beginning of a long and
complicated experimental series on neutralization, carried out
with great care and repeated many times in the years which
followed.
The idea of these experiments was to develop some concep-
tion of the mechanism of neutralization. He pushed the problem
as hard as he could with the techniques at his command, but it
was not until many years later that the mystery of neutralization
began to unravel, and it was learned that much simpler host
systems were essential to understand neutralization mechanisms.
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240
BIOGRAPHICAL MEMOIRS
Shortly after starting work faith the Foundation, Horsfall
took time out for a six-month sabbatical with Arne Tiselius at
Uppsala. At that time Tiselius was carrying out his important
pioneer work on the electrophoresis of macromolecules. A1-
though Horsfall published a couple of papers with Tiselius in
the field of physical chemistry, it is clear that he could not use
this experience in virology without wandering far afield from
his customary goals.
The main attraction of the Foundation position was the
possibility of studying human influenza infection on a vast scale
and attempting prophylaxis against the disease. The Foundation
was well equipped for this sort of venture. With its previous
experience in yellow fever research, it had developed numerous
experienced epidemiologists and other fieldworkers whose value
in executing the experiments that were planned in New York
would be difficult to exaggerate. In his first years with the
Foundation, Horsfall carried out some routine but straightfor-
ward investigations of influenza epidemics. This work, in a very
new field, carried the mark of competence with it.
A ferret colony was an essential resource in carrying out
influenza work at that time, and during the course of some
early experiments, a spontaneous epidemic of distemper broke
out in the Foundation's animal house and nearly all the animals
were lost. To prevent a recurrence of this disaster, all new ani-
mals were immunized, in most cases using a formalinized spleen
from a spontaneously infected animal. Later, some of these im-
munized animals were found to have mysteriously acquired high
anti-influenza titers. It occurred to Horsfall at once that dis-
temper virus infection might have some sort of cooperative
effect in inducing high-level and persistent antibody responses
to influenza virus.
It was a fascinating idea, and a "complex" vaccine containing
both influenza and distemper virus was quickly developed for
use in human beings. The Foundation provided large-scale pro-
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FRANK LAPPIN HORSFALL, JR. 241
auction facilities for the new product, which was made by
growing both viruses in chick embryos and then freeze-drying a
formalinized extract. With the threat of a widespread epidemic
(1940-1941), there was no time to do a lot of testing on prepara-
tions before they could be used in human experiments. There
were some feverishly performed experiments in laboratory ani-
mals and man, but they were not completed in time to modify
the large-scale experiments which were being contemplated in
the field.
Large amounts of vaccine were prepared and shipped to
Great Britain, which was at war, just in case the epidemic
became intolerable. The British examined the preparations in
great detail, but did not utilize an opportunity to try them. In
the United States, however, Horsfall conducted a large-scale
demonstration, vaccinating, under well-controlled circum-
stances, volunteers in a large number of state prisons up and
down the Eastern Seaboard.
The experiment was carried out in magnificent style. E. R.
Rickard, who was Horsfall's chief field marshal, was a master at
this type of human experiment. Not only was the distribution
of inoculations carefully controlled, but a great deal of data was
obtained on individuals through attempts at virus isolation and
by determining pre- and post-infection antibody titers. For-
tunately, the inoculations were completed a relatively short
time before the onset of the epidemic, and the use of a large
number of population groups proved valuable in the final anal-
ysis of the results.
In spite of the excellence of organization and the occurrence
of a large-scale epidemic in the wake of vaccination, the results
were very disappointing. Although antibody rises had been in-
duced in many vaccine recipients, the rise in titer was small and
evanescent, and the reduction in case rate was in most institu-
tions negligible or at best marginal. Only in a couple of places
was there as much as a 50 percent drop in incidence following
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242
BI OGRAPHICAL MEM OIRS
vaccination, ant! it turned out that in these places the batch of
vaccine used had had the highest virus titer before formalini-
zation.
When all the laboratory tests on the complex vaccine were
finally in, the results explained in some degree the disappointing
field tests. In general the virus content of the vaccine was low,
and the initially promising effect of distemper virus in boosting
antigenicity could never be reproduced. It is again rather ironic
that, just as these results were being recorded, new methods of
obtaining high virus titers, as well as high virus purity, were
being developed. It wasn't until several years later that vaccines
made with these improved reagents were tested and gave the
first bona fide protection against influenza in the field. Even
years later the secret of obtaining substantial and prolonged
effects against epidemic influenza is still a formidable problem,
and the earliest experiences with the "complex" vaccine were
by no means unique.
Just as the episode of the vaccine trial was coming to a close,
Horsfall was called back to the Rockefeller Institute Hospital,
where he received a lifetime appointment as a full member.
Essentially he took over Rivers' Department of Virology, but
he peopled it with entirely new personnel, and in his new pro-
gram he once again confined his attention to respiratory disease.
Rivers, who was now Director of the Rockefeller Institute
Hospital, played a very important role in shaping developments
within that organization. The return of Horsfall, with the pos-
sibility that he might become the Hospital's next Director,
brought Rivers' planning to a climax.
By now the United States had entered the war, and Rivers
organized a naval unit at the Hospital, most of which later went
to Guam under his command. The remainder of this unit stayed
on at Rockefeller Institute with Horsfall as the commanding
officer, and as a group they worked on respiratory diseases, espe-
cially those that they felt might have a virus etiology such as
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FRAN K LAPPIN HORSFALL, JR.
1945
~7
With E. C. Curnen, G. S. Mirick, l. E. Ziegler, Jr., and L. Thomas.
Studies on primary atypical pneumonia. I. Clinical features and
results of laboratory investigations. i. Clin. Invest., 24:209-26.
With L. Thomas, G. S. Mirick, E. C. Curnen, and l. E. Ziegler, Jr.
Studies on primary atypical pneumonia. II. Observations con-
cerning the relationship of a non-hemolytic streptococcus to tl~e
disease. l. Clin. Invest., 24:227~0.
1946
With E. C. Curnen. Studies of pneumonia virus of mice. I. The
precision of measurements in vivo of the virus and antibodies
against it. i. Exp. Med., 83:25~.
With E. C. Curnen. Studies of pneumonia virus of mice. II. Im-
munoIogical evidence of latent infection with the virus in
numerous mammalian species. I. Exp. Med., 83:43-64.
With E. C. Curnen. Studies of pneumonia virus of mice. III.
Hemagglutination by the virus: The occurrence of combination
between the virus and a tissue substance. .T- Exp. Med., 83:
105-32.
Primary Atypical Pneumonia. N.Y. State l. Med., 46: 1810-14.
1947
With E. C. Curnen and E. G. Pickels. Centrifugation studies on
pneumonia virus of mice (PVM). ]. Exp. Med., 85:23-38.
With E. C. Curnen. Properties of pneumonia virus of mice (PVM) in
relation to its state. l. Exp. Med., 85:39-53.
With J. E. Ziegler, fir., E. C. Curnen and G. S. Mirick. Diagnosis of
acute respiratory infections. Am. l. Med. Sci., 213:268-81.
With M. McCarty. The modifying effects of certain substances of
bacterial origin on the course of infection with pneumonia virus
of mice (PVM). J. Exp. Med., 85:623~6.
Primary atypical pneumonia. Ann. Intern. Med., 27:275-81.
Virus och virussjukdomar has manniskan. Nord. Med., 34:1333~1.
Latenta virus och deras relation till infektioner. Nord. Med., 35:
1611-18.
Hemagglutination och forekomsten av bundet virus. Nord. Med.,
35: 1739-46.
With M. Volkert, C. Pierce, and R. J. Dubos. The enhancing effect
OCR for page 258
258
BIOGRAPHICAL MEMOIRS
of concurrent infection with pneumotropic viruses on pulmonary
tuberculosis in mice. l. Exp. Med., 86:203-13.
With M. Volkert. The effect of sulfhydryl groups on pneumonia
virus of mice (PVM). J. Exp. Med., 86:383-91.
\Vith M. Volkert. Studies on a lung tissue component which com-
bines with pneumonia virus of mice (PVM). if. Exp. Med., 86:
393-407.
With H. S. Ginsberg and W. G. Goebel. Inhibition of mumps virus
multiplication by a polysaccharide. Proc. Soc. Exp. Biol. Med.,
66:99-100.
With M. McCarty. The antagonistic effect of certain substances of
bacterial origin on the course of infection with pneumonia virus
of mice (PVM). Trans. Assoc. Am. Physicians, 60:18-21.
1948
With H. S. Ginsberg and W. G. Goebel. The inhibitory effect of
polysaccharide on mumps virus multiplication. l. Exp. Med.,
87:385-410.
With H. S. Ginsberg and W. G. Goebel. The effect of polysaccha-
rides on the reaction between erythrocytes and viruses, with
particular reference to mumps virus. l. Exp. Med., 87:411-24.
Primary atypical pneumonia and influenza~iagnosis, prevention,
treatment. Bull. N.Y. Acad. Med., 24:431-46.
With S. E. Bjorkman. The production of persistent alteration in
influenza virus by lanthanum or ultraviolet irradiation. l. Exp.
Med., 88 :445-61.
With P. H. Hardy, fir. Reactions between influenza virus and a
component of allantoic fluid. J. Exp. Med., 88:463-83.
With F. M. Davenport. Assocative reactions of pneumonia virus of
mice (PVM) and influenza viruses: The effects of pH and electro-
lytes upon virus host-cell combinations. l. Exp. Med., 88:621~4.
1949
With H. S. Ginsberg. Concurrent infection with influenza virus and
mumps virus or pneumonia virus of mice (PVM) as bearing on
the inhibition of virus multiplication by bacterial polysaccha-
rides. J. Exp. Med., 89:37-52.
With O. Lahelle. Multiplication of influenza virus in dead chick
embryos. Proc. Soc. Exp. Biol. Med., 70:547-51.
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FRAN K LAPPIN HORSFALL, JR.
259
With H. S. Ginsberg. Chemotherapy of viral infections. In: Evalua-
tion of Chemotherapeutic Agents, pp. 170-80. N.Y.: Columbia
Univ. Press.
Prospects for the control of viral diseases by chemical agents. Can.
Med. Assoc. I., 60:439~7.
With P. K. Olitsky, l. Casals, D. L. Walker, and H. S. Ginsberg.
Preservation of viruses in a mechanical refrigerator at 24°.
J. Lab. Clin. Med., 34:1023-26.
With E. D. Kilbourne. A chemical method for the detection of virus
infection of the chick embryo. Proc. Soc. Exp. Biol. Med., 71:
708-13.
With O. Lahelle. Hemagglutination with the GDVIII strain of
mouse encephalomyelitis virus. Proc. Soc. Exp. Biol. Med., 71:
713-18.
With H. S. Ginsberg. A resistant variant of mumps virus. Multi-
plication of the variant in the presence of inhibitory quantities
of Friedlander bacillus polysaccharide. I. Exp. Med., 90:393~07.
With H. S. Ginsberg. A labile component of normal serum which
combines with various viruses. Neutralization of infectivity and
inhibition of hemagglutination by the component. l. Exp. Med.,
90:475-95.
1950
With F. M. Davenport. Further studies on the associative reactions
of pneumonia virus of mice (PVM) and influenza viruses. Com^
bination with various animal tissues and absorbents. J. Exp.
Med., 91:53-64.
With D. L. Walker. Lack of identity in neutralizing and hemag-
glutination-inhibiting antibodies against influenza viruses. J.
Exp. Med., 91:65-86.
Advances in the diagnosis of virus infections. Bull. N.Y. Acad. Med.,
26:3-19.
With I. Tamm. Characterization and separation of an inhibitor of
viral hema~lutination present in urine. Proc. Soc. Exo. Biol.
Med., 74:108-14.
.~, .
Chemotherapy in viral infections. (The Musser Lecture) Am. l.
Med. Sci., 220:91-102.
Approaches to the control of viral diseases. Bact. Rev., 14:219-24.
With I. Archetti. Persistent antigenic variation of influenza A
OCR for page 260
260
BIOGRAPHICAL MEMOIRS
viruses after incomplete neutralization in eve with heterologous
immune serum. J. Exp. Med., 92:441-62.
With I. Tamm and E. D. Kilbourne. Comparison of influenza B
strains from the 1950 epidemic with strains from earlier epi-
demics. Proc. Soc. Exp. Biol. Med., 75:89-92.
With H. S. Ginsberg. The implications of chemical interruption of
viral multiplication. Trans. Assoc. Am. Physicians, 63:118-21.
1951
With H. S. Ginsberg. The dependence of the pathological lesion
upon the multiplication of pneumonia virus of mice (PVM).
Kinetic relation between the degree of viral multiplication and
the extent of pneumonia. it- Exp. Med., 93:139-50.
With H. S. Ginsberg. Characteristics of the multiplication cycle of
pneumonia virus of mice (PVM). ]. Exp. Med., 93:151-60.
With H. S. Ginsberg. Therapy of infection with pneumonia virus
of mice (PVM). Effect of a polysaccharide on the multiplication
cycles of the virus and on the course of the viral pneumonia.
J. Exp. Med., 93: 161-71.
Studies on non-hemolytic streptococci isolated from the respiratory
tract of man. The antigenic basis for type specific reactions with
Streptococcus salivarius and non-levan-forming streptococci.
J. Exp. Med., 93:229-45.
With E. D. Kilbourne. Increased virus in eggs injected with corti-
sone. Proc. Soc. Exp. Biol. Med., 76:116-18.
With H. S. Ginsberg. An improved CO2 cabinet for low temperature
storage of infectious agents with gaseous CO2 excluded from the
specimen compartment. J. Bacterial., 61:443-51.
With E. D. Kilbourne. Lethal infection with Coxsackie virus of
adult mice given cortisone. Proc. Soc. Exp. Biol. Med., 77:135-38.
With E. D. Kilbourne. Primary herpes simplex virus infection of
the adult: With a note on the relation of herpes simplex virus
to recurrent aphthous stomatitis. Arch. Int. Med., 88:495-502.
With E. D. Kilbourne. Studies of herpes simplex virus in newborn
mice. l. Immunol., 67:321-29.
With H. S. Ginsberg. Interference between mumps virus and pneu-
monia virus of mice (PVM). Fate of mumps virus in the mouse
lung. J. Immunol., 67:369-77.
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FRANK LAPPIN HORSFALL, JR.
261
With E. D. Kilbourne. Mouse-egg neutralization. Neutralization in
the mouse of influenza viruses not adapted to the mouse. J.
Immunol., 67:43 1-36.
With E. D. Kilbourne and H. C. Anderson. Concurrent infection
with influenza A and B viruses in a single epidemic of influenza.
I. Immunol., 67:547-58.
1952
With I. Tamm. A mucoprotein derived from human urine which
reacts with influenza, mumps and New Castle disease viruses.
J. Exp. Med., 95:71-97.
With G. E. Perlman and I. Tamm. An electrophoretic examination
of a urinary mucoprotein which reacts with various viruses. I.
Exp. Med., 95:99-104.
With H. S. Ginsberg. Quantitative aspects of the multiplication of
influenza A viruses in the mouse lung. Relation between the
degree of viral multiplication and the extent of pneumonia. l.
Exp. Med., 95: 135-45.
Factors concerned with viral proliferation in vivo. Ann. N.Y. Acad.
Sci., 54:926-35.
With I. Tamm and K. Folkers. Inhibition of influenza virus multi-
plication by 2,5-dimethlbenzimidazole. Yale l. Biol. Med., 24:
559-67.
With M. R. Hilleman. Comparison of the antigenic patterns of
influenza A virus strains determined by in ovo neutralization
and hemagglutination inhibition. J. Immunol., 69: 343-56.
With D. A. J. Tyrrell. A procedure which eliminates nonspecific
inhibitor from human serum but does not affect specific anti-
bodies against influenza viruses. I. Immunol., 69:563-74.
With I. Tamm. Variation demonstrable by methods other than
serological. Bull. N.Y. Acad. Med., 28:765-68.
With I. Tamm and H. S. Ginsberg. Antigenic similarity between
the first and later egg passage strains of freshly recovered in-
fluenza A viruses. Proc. Soc. Exp. Biol. Med., 81:94-98.
With W. i. Mogabgab. Effect of sodium monofluoroacetate on the
multiplication of influenza viruses, mumps virus and pneumonia
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Representative terms from entire chapter:
pneumonia virus
