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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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236 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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238 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

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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

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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 virus of mice (PVM). T. Exp. Med., 96:531-48.

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262 BIOGRAPHICAL MEM OIRS 1953 With I. Tamm. Fractional dilution procedure for precise titration of hemagglutinating viruses and hemagglutination-inhibiting antibodies. J. Immunol., 70:253-59. With D. A. l. Tyrrell. Neutralization of viruses by homologous im- mune serum. I. Quantitative studies on factors which affect the neutralization reaction with Newcastle disease, influenza A and bacterial virus T3. J. Exp. Med., 97:845-61. With I. Tamm and K. Folkers. Inhibition of influenza virus multi- plication by alkyl derivatives of benzimidazole. I. Kinetic aspects of inhibition by 2,5-dimethylbenzimidazole as measured by in- fectivity titrations..~. Exp. Med., 98:219-27. With I. Tamm and K. Folkers. Inhibition of influenza virus multi- plication by alkyl derivatives of benzimidazole. II. Measurement of inhibitory activity by hemagglutination titrations. J. Exp. Med., 98:229~3. With I. Tamm, K. Folkers, C. H. Shunk, and D. Heyl. Inhibition of influenza virus multiplication by alkyl derivatives of benz- imidazole. III. Relationship between inhibitory activity and chemical structure. l. Exp. Med., 98:245-59. 1954 Experiments on chemical alteration of virus infections. In: The Harvey Lectures, Series 98. N.Y.: Academic Press. With I. Tamm, K. Folkers, and C. H. Shunk. Inhibition of in- fluenza virus multiplication by N-glycosides of benzimidazoles. J. Exp. Med., 99:227-50. With F. W. Hartman and I. G. Kidd, eds., The Dynamics of Virus and Rickettsial Infections. (International symposium.) N.Y.: The Blakiston Co. With D. A. J. Tyrrell. Disruption of influenza virus. Properties of degradation products of the virus particle. l. Exp. Med., 99: 321-42. With T. Nadeje and J. Blum. An automatic tool for opening em- bryonated eggs. Rev. Sci. Instrum., 25:293-94. On the reproduction of influenza virus. Quantitative studies with procedures which enumerate infective and hemagglutinating virus particles. J. Exp. Med., 100:135-61.

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FRAN K LAPPIN HORSFALL, JR. 263 With D. A. l. Tyrrell, I. Tamm, and O. C. Forssman. A new count of the allantoic cells of the 10 day chick embryo. Proc. Sac. Exp. Biol. Med., 86:59~98. Chemotherapy of respiratory viral diseases. (Special Reviews edited by Joseph Stokes, Jr., M.D.) Pediatrics, 13: 593-98. 1955 The inhibition of virus reproduction by chemical substances. In: Perspectives and Horizons in Microbiology: Symposium, ed. S. A. Waksman, pp. 152-67. New Brunswick, N.~.: Rutgers Univ. Press. With I. Tamm and I. C. Bugher. Ultracentrifugation studies of a urinary mucoprotein which reacts with various viruses. J. Biol. Chem., 212: 125-33. On the consequences of virus reproduction. Trans. Assoc. Am. Physicians, 68:54-63. Reproduction of influenza viruses. Quantitative investigations with particle enumeration procedures in the dynamics of influenza A and B virus reproduction. J. Exp. Med., 102:441-73. Approaches to the chemotherapy of viral diseases. Bull. N.Y. Acad. Med., 31: 783-93. Virus reproduction and virus disease. Can. Med. Assoc. I., 73:778-82. 1957 Interaction of polysaccharides and viruses. In: Polysaccharides in Biology: Transactions of the Second Conference, April 25, 26, 27, Princeton, NJ., ed. Georg F. Springer, pp. 41-113. N.Y.: Josiah Macy, Jr. Foundation. With I. Tamm. Chemotherapy of viral and rickettsial diseases. Ann. Rev. Microbial., 11: 339-70. Virus Diseases, Public Health Rep., 72:905-6. Virus neutralization tests: Implications and interpretations. Ann. N.Y. Acad. Sci., 69:633~3. 1958 Can viruses be managed? Proc. Amer. Phil. Soc., 102:442~7. Viral multiplication, in poliomyelitis. In: Papers and Discussions Presented at the Fourth International Poliomyelitis Conference (Geneva, 1957), pp. 335~1. Philadelphia: J. B. Lippincott.

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264 BIOGRAPHICAL MEMOIRS 1959 With T. M. Rivers, eds., Viral and Rickettsial Infections of Man, ad ed. Philadelphia: l. B. Lippincott. Our new knowledge of viruses. Worldwide Abstr. Gen. Med., 2: 11-18. Inhibition of multiplication. In: The Viruses, Biochemical, Biologi- cal and Biophysical Properties, vol. 3, Animal Viruses, ed. F. M. Burnet and W. M. Stanley, pp. 195-224. N.Y.: Academic Press. Viral infections of the respiratory tract. (The I. Burns Amberson Lecture.) Amer. Rev. Resp. Dis., 80:315-25. 1960 Viral infection and viral disease in man. (Mayo Foundation Lec- ture.) Proc. Staff Meetings Mayo Clin., 35:269-82. 1961 On the unity of the sciences. Interreactions among the physical and biological sciences show that unification is progressive. Science, 133: 1059-60. Factors contributing to recovery from viral diseases. Can. Med. Assoc. J., 84:1221-26. Summary in poliomyelitis. In: Papers and Discussions Presented at the Fifth International Poliomyelitis Conference, Copenhagen 1960, pp. 425-26. Philadelphia: l. B. Lippincott. On the unity of the sciences. In: Medical Education and Medical Care: Interactions and Prospects. Report of the Eighth Teaching Institute, Association of American Medical Colleges. i. Med. Ed., 36:2~28. Extrinsic factors (environment). In: Congenital Malformations. Papers and Discussions Presented at the First International Con- ference (London 1960J, pp. 124-29. Philadelphia: I. B. Lippin- cott. Outlook for medical progress in this decade. A discussion of the probable effects of medical advances on longevity: Progress in cancer. Trans. Soc. Actuaries, 13:494-98. Dr. Leo Wade. Arch. Environ. Health, 3:496-98.

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FRAN K LAPPIN HORSFALL, JR. 1962 265 Heritance of acquired characters. A unifying concept is developed in relation to the genesis of cancer. Science, 136:472-76. A look to the future. Cancer. Worldwide Abstr. Gen. Med., b: 13-14. Some new concepts in biology. Arch. Biochem., l(Suppl.~:63-67. 1963 Summary of the conference. In: Congenital Defects. Papers and Dis- cussions Presented at the First Inter-American Conference (Los ~lugeles 1962~' pp. 249-51. Philadelphia: J. B. Lippincott. Viruses and cancer. Acta Unio Int. Contra Cancrum, 19~1-2~:247~9. Role of infectious agents in the pathogenesis of neoplasia. In: Recent Progress in Microbiology, VIII International Congress for Microbiology (Montreal 1962J, pp. 458-66. Toronto: Univ. Of Toronto Press. A unitary concept of the origin of cancer. Bull. Un. Int. Cancer, 1 (1 August 1963~: 1. Foreword in honor of Thomas Milton Rivers. In: Perspectives in Virology III, ed. M. Pollard, pp. xvii-xix. N.Y.: Harper & Row. The role of viruses in relation to cancer in animals and man. In: Viruses, Nucleic Acids, and Cancer, Seventeenth Annual Sym- posium on Fundamental Cancer Research (Houston 1963J. Baltimore: Williams & Wilkins. Current concepts of cancer. Can. Med. Assoc. T., 89: 1224-29. 1964 Citation and presentation of the Academy Medal to Gilbert Dall- dorf, M.D. Bull. N.Y. Acad. Med., 40:267-68. Introductory and concluding remarks. In: Congenital Malforma- tions, Papers and Discussions Presented at the Second Inter- national Conference (New York 1963J, ed. M. Fishbein, pp. 3~. N.Y.: International Medical Congress. Some facts and fancies about cancer. (George Kober Memorial Lec- ture.) Georgetown Med. Bull., 18: 37-45. Committee on Science and Public Policy. Federal Support of Basic

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266 BIOGRAPHICAL MEMOIRS Research in Institutions of Higher Learning. Pub. No. 1185. Wash., D.C.: National Academy of Sciences-National Research Council. The President's Commission on Heart Disease, Cancer and stroke. In: Report to the President: ~4 National Program to Conquer Heart Disease, Cancer and Stroke, vol. 1. Wash., D.C.: U.S. Gov- ernment Printing Office. 1965 Some facts and fancies about cancer. Perspect. Biol. Med., 8:167-79. The President's Commission on Heart Disease, Cancer and stroke. Report of the subcommittee on cancer. In: Report to the Presi- dent: A National Program to Conquer Heart Disease, Cancer and Stroke, vol. 2, pp. 103-17. Wash., D.C.: U.S. Government Printing Once. The President's Commission on Heart Disease, Cancer and stroke. Report of the subcommittee on research. In: Report to the Presi- dent: a National Program to Conquer Heart Disease, Cancer and Stroke, vol. 2, pp. 137-61. Wash., D.C.: U.S. Government Printing Once. Federal support of biomedical sciences. In: Basic Research and National Goals. A report to the Committee on Science and Astronautics, U.S. House of Representatives, by the National Academy of Sciences, March 1965, pp. 111-25. Thomas Milton Rivers. In: Biographical Memoirs, 38:263-94. New York: Columbia Univ. Press for the National Academy of Sciences. Fatti e fantasie sul cancro. Rassegna Medica e Culturale (Milano), 42:10-17, N.3. 1966 Cancer and viruses. Bull. N.Y. Acad. Med. 42:167-81. Leukemia in man and mouse. Acta Med. Scand. (Suppl. 445), 179: 304-11. Unifying concept of the origin of cancer. Med. Clin. of N. Amer. 50:869-74. Concepto unificador del origien deI cancer. In: Clinicas medicas de Norteamerica, Mexico, D. F. Editorial Interamericana, s.a., ed. D. A. Karnofsky and R. W. Rawson, pp. 869-74.

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FRANK LAPPIN HORSFALL, JR. 267 Introductory remarks. In: Macromolecules and Cancer, Sloan- Kettering Institute for Cancer Research Symposium (New York 1965J. Cancer Res., 26:1957. Peyton Rous, Nobel Laureate 1966. The Rockefeller University Review, 4: 1~. Introductory remarks. Transformation mechanism. In: Subviral Carcinogenesis, ed. Y. Ito, 1st International Symposium on Tumor Viruses (Nagoya, 1966), p. 98. 1968 Immunology in 1968. (Presidential address before the American Association of Immunologists.) l. Immunol., 101:183-86.