Francis Peyton Rous | Biographical Memoirs V.48 | The National Academies Press Cancel Below is the uncorrected machine-read text of this chapter, intended to provide our own search engines and external engines with highly rich, chapter-representative searchable text of each book. Because it is UNCORRECTED material, please consider the following text as a useful but insufficient proxy for the authoritative book pages. ~  :~: At:  .~ ~~ - : :: : - :: ~ - - ~r~~ ~ ~ Ad. .. ::: .~. . ~ ::: . FRANCIS PEYTON ROUS Octo her 5, ~ 879-February ~ 6, 1970 BY RENATO DULBECCO PEYTON ROUS was awarded the Nobel Prize in 1966, when he was eighty-six years old, for discoveries he had made fifty years before. He was born on the 5th of October 1879 in Balti- more, Maryland, to a family that valued humanistic education.

Thus, after the death of his father, when Peyton Rous was a child, his mother rejected the idea of joining her family in Texas and stayed in Baltimore, where excellent education for the children was available. Of the two sisters of Peyton Rous, one became a musicologist, the other a painter; and Peyton himself had a flair for writing. Peyton Rous enrolled in the Medical School of the recently created Johns Hopkins University, which he attended without special distinction. As an undergraduate he showed a naturalist's tendency and published articles about Baltimore's flowers. After graduating in medicine, he went to the University of Michigan, where he began his research career in pathology, which he perfected during a year in Dresden. In 1909 he joined the Rockefeller Institute under Simon Flexner, to engage in cancer research, against the opinion of influential friends who thought it was a hopeless field. There he remained until his death. In 1921 he became co-editor (and later editor) of the Journal for Experimental Medicine. In 1927 he was elected to the National Academy of Sciences.

He died in 1970, at the age of ninety, and 275 276 BIOGRAPHICAL MEMOIRS is survived by his wife, Marion, and three daughters, Marion, Ellen, and Phoebe. In addition to the Nobel Prize, Peyton Rous received many honors and honorary degrees, which are listed at the end of this memoir. The name of Peyton Rous became widely known to biolo- gists in the fifties and sixties for his earlier discovery of a virus causing sarcoma in chickens, which became aptly known as the Rous Sarcoma Virus. At the time he became famous, Peyton Rous appeared as an elderly, highly educated, gentleman with silvery hair. But in his youth he was a very hardworking scientist with a determined, fiery, and highly critical personality. He was a medical man who wished to learn about cancer as a disease and a biologist who did not want to follow the beaten track, and he was willing to hunt for new clues in well-designed but slow experiments.

I, like most of my contemporaries, became acquainted with Peyton Rous's fundamental discovery in the early fifties, when Harry Rubin came to my lab to work with the Rous Sarcoma Virus. He started using a focus technique on the chorioallantoic membrane of the chicken embryo, which Rous had invented many years before. Later I had occasion to meet Peyton Rous several times on the platform as a speaker, or across the discus- sion table, or in his laboratory at the (then) Rockefeller Institute. I remember the manrather small in stature with silvery hair and penetrating eyes. I also remember that before our first meet- ing I was inclined to think of him as a figure of the past, but soon changed my mind at that meeting and even more so at subsequent ones. Clearly, he was very much alive until his very last days, with a keen interest in new developments in virology and cancer research. He was able to discuss his past work with equanimity and to accept new interpretations of his data.

I re- member I suggested to him an explanation of the clonal charac- teristic of the neoplastic transformation of papillomas in terms FRANCIS PEYTON ROUS 277 of somatic cell genetics, a concept that was not part of cancer research in the period of his active work. His interest was im- mediately aroused; he asked me for a thorough clarification of what I meant and then argued, with passion but no animosity. ... ~ ... .. ~ . . . . - rim ~ we parted Able old friends who have found something new to talk about. 4- -' At the time when phage lysogeny was the domain of a very small group of virolo~ists. I suggested to him that it . . might represent a good model for some features of viral cancer. Again his interest was acute, and I had to embark on a detailed discussion of phage integration, immunity, and lysogenic con- version. Peyton Rous discovered the viral etiology of a chicken sar- coma in 1911 through his interest in tumor tr~n~nl~nt~hilirv to 1 , ~ - ~ YE new nosts oy a nitrate.

He commented: "The behaviour of the new growth has been throughout that of a true neoplasm, for which reason the fact of its transmission by means of a cell-free filtrate assumes exceptional importance" (191 1~. He fully realized from the outset that this was "a unique and significant finding" (1911~. He also realized that the signifi- cance of the discovery depended on the true nature of the induced growth. As an experienced pathologist he could see that it was a true cancer: "The (pathological) picture (of the growth) does not in the least suggest a granuloma . . . it exhibits to a special degree, not merely a few, but all those features by which the malignant neoplasms are characterized" (191 1~. For about forty years this momentous discovery had little impact, because the minds of scientists were not prepared to think of viruses as agents of cancer. It was expedient to say that the chicken tumor was not a cancer, but some kind of reaction to the virus more akin to inflammation than neoplasia, and perhaps a peculiarity of chicken biology.

Peyton Rous soon recognized himself that the tumor would not be accepted as a cancer because it was transmitted by a cell-free extract: "A passing reference should perhaps be made to the ill-defined 278 BIOGRAPHICAL MEMOIRS group of pathological products called granulomata, with which this neoplasm of the fowl may by some be classed, owing to its transmission by an agent separable from the tissue cells" (191 1~. Many years later he wrote, "This disclosure (that certain chicken tumors were proved due to viruses), which conflicted with the negative findings in mammalian growths, was determined forth- with as erroneous" (1952~. One wonders how firmly in the early years Peyton Rous himself was convinced that he had demonstrated the induction of a cancer by viruses. The statements he made at the time are very cautious and full of qualifications. At first he used to refer to the "agent" that induced the sarcoma; but a year later, after he discovered a new, different tumor transmissible by filtrate, he proposed that the "agent is probably a living virus" (1912~.

During the years 1911-1914, Peyton Rous worked hard at disproving the objections on the nature of the induced tumors by isolating other viruses that induced tumors in chickens and by carefully studying their pathology. He could show that the tumors induced by the different viruses were capable of invad- ing neighboring tissues and of metastasizing to distant organs; thus they were true cancers. Moreover, each independently isolated virus caused a tumor of a different kind. These facts should have been convincing evidence that the growths were specific responses of the host, yet this conclusion was not gener- ally accepted. However, these discoveries seem to have been convincing for Rous, who wrote, "The findings with the chicken tumors largely demolish the theoretical basis in which objec- tions to an extrinsic cause for cancer have been built up" (1912~. In order to find more generally acceptable evidence, Peyton Rous attempted to extend his observations "especially through carefully devised experiments with the tumors of other species of animals" (1911~. Evidently for the viral etiology to be ac- cepted, similar findings were needed in mammals. The strategy of Rous's future work was determined at that time.

However, FRAN CIS PEYTON ROUS 279 the extension to other species came only many years later with Richard Shope's discovery of the rabbit papilloma virus. In the meantime Peyton Rous studied many features of the cell-free transmission of the tumor. Examining the effect of the age of the host, he showed that the virus induces characteristic foci on the chorioallantoic membrane of the chicken embryo. This result supplied an assay for the virus that was universally employed until the fifties, when it was superseded by the focus formation in tissue culture. In this extensive and careful work, Peyton Rous observed the host resistance to the transmission of the tumor, in the form of either absence of growth, slow growth, or normal growth followed by regression. Other experiments showed how essen- tial the conditions of the host are for the development of a tumor after inoculation of the virus. From this observation Peyton Rous began to recognize the existence of limitations to the expression of the oncogenic potential of the virus: "How does it happen that the sarcoma, though ultimately dependent on an extrinsic agent, is dominated in its behaviour by the cells composing it?" (1912~.

Perhaps the agent depends "on a special set of conditions in order that it might produce a neoplastic change" ( 1912~. He returned later to this Point on several occasions. . . , After discovering the second chicken tumor agent, Peyton Rous started wondering about the etiology of cancer in general: "The demonstration that extrinsic agents are the cause of two connective-tissue growths of the fowl which are characteristic malignant tumors renders it necessary to suppose either that such tumors of the fowl have an entirely different etiology from mammalian tumors, or else that the latter are of similar origin" (1912~. This point was also developed to a much greater extent later on. As further evidence for a viral nature of the chicken tumors, the resistance of the host to the tumor cells could be separated 280 BIOGRAPHICAL MEMOIRS from its resistance to the tumor-inducing agent.

Moreover, Rous discovered a third chicken tumor, transmissible by filtrate, markedly different in properties from the two previously de- scribed: "The findings with the three tumor-producing agents have a striking similarity and it is difficult to avoid the con- clusion that the three are of one class, whatever that class may be.... It is perhaps not too much to say that their recognition points to the existence of a new group of entities which cause in chickens neoplasms of diverse characters" (1914~. At the beginning of World War I, Peyton Rous, under the pressure of wartime medical needs, gave up his work with chicken tumor viruses. For the following twenty years until 1934, his interest was in the fields of blood transfusion and attending immune reactions, liver and biliary functions, cellu- lar functions, and vascular permeability. I will return to these · . . activities ater on. A turning point in Peyton Rous's work on cancer was the discovery of the Shope papilloma. In 1933 Richard Shope re- ported his discovery that a mammalian tumor, the papilloma of cottontail rabbits, was transmitted by a virus-like agent.

As in the case of the chicken tumor, Peyton Rous's first concern was whether the papilloma was a true neoplasm. He decided that it was, because, when the papillomas were transplanted deep in- side the body, they developed into carcinomas that grew inva- sively and killed the host. Furthermore, in domestic rabbits the virus-induced papillomas often grew progressively, invading the neighboring tissues and producing metastases, and this malig- nant evolution could be enhanced by exposing the papillomas to various substances, such as Scarlet Red. These findings seem to have been for Rous the decisive argument for the validity of his conclusions concerning the chicken tumors, since in a mammal cancer could also be trans- mitted by a virus. He, therefore, returned to the study of car- cinogenesis using the papilloma virus as a new tool. He focused FRANCIS PEYTON ROUS 281 at first on the malignant evolution of the papillomas.

By careful observations, following small hints, such as the shape of their growths, color, or the degree of pigmentation, he showed that a few cells in a papilloma became cancerous and generated clones, each with different characteristics. In trying to understand how such evolution to cancer occurs, Peyton Rous studied the effects of tar, as both a carcinogen and tumor promoter. He found that tar not only strongly enhanced the induction of papillomas or carcinomas by the Shope virus in domestic rabbits but by itself elicited similar papillomas. Could tar papillomas also be virus-induced? This new phase of Peyton Rous's work, although a natural development of his earlier work, had more ambitious goals, for it aimed at testing the hypothesis that "this disease (cancer) is an infection.... A main attraction of this hypothesis is its accessibility to test." However, he clearly saw that this hypothe- sis could only be true under certain conditions, one of which is that "a living entity responsible for such growths must require for effectiveness a very special basis of predisposition" (1932~.

He sought to possibly disprove the infectious nature of cancer by comparing the frequency of cancer induction by tar in the skin of two groups of mice with different exposure to- the envi- ronment: "The animals of one group have been placed under conditions which would facilitate the entrance into the body of extraneous living agents, whereas those of the others have been sedulously protected" (1932~. The results proved "that the mouse cancer cannot be caused by living entities reaching the body from the surrounding world during adult life" but "fail to exclude the possible activity of entities residing habitually in or upon the body" (1932~. This experiment showed another re- quirement of the hypothesis on the infectious nature of cancer: "The supposition (that tumors in general are due to viruses or other extraneous entities) is tenable only if such entities are widely distributed throughout the animal population, being 282 BIOGRAPHICAL MEMOIRS constantly present in or upon the body, like the colon bacillus or the staphylococcus; and if their opportunity to cause tumors is restricted by the need for very special conditions....

The more considerable an agent is conditioned in its activity, the more often must it be present if it is to cause disease at all" (1934~. These words were prophetic, as shown by the recent developments in the field; yet they were simply the result of cool, logical assessment of the facts then in hand. However, for Peyton Rous this hypothesis was only a guide for the experiment: "The demonstration of the cause for the generality of tumors, what- ever this is, waits upon the provision by the investigator of the conditions necessary to its effectiveness" (1934~. He tried several new approaches. One of the major tools was still the technique of inducing skin tumors by application of tar. He used it to create favorable cellular conditions for reveal- ing the neoplastic potential of viral agents. Another tool was the immunity of the infected rabbits against the Shope virus. Peyton Rous found no demonstrable antibodies in rabbits with- out papillomas or in those with tar papillomas or Brown-Pierce tumors: these findings "speak decisively against the possibility that these growths are caused by viruses antigenically related to the one causing papillomas.

Yet this does not exclude a virus causation for them, since the sera of fowls with Chicken Tumor I and Fujinami Sarcoma respectively, though possessed of neu- tralizing power for the virus causing the growth carried by the host, have no cross-neutralizing effect whatsoever" (1936~. Shortly afterwards, in taking a bird's eye view of his past work and of the cancer problem, he concluded: "How far should one be led by the assumption that certain tumors may be due to viruses? Only so far as to make tests with these growths. The tumor problem has withstood the most corrosive reasoning. Yet since what one thinks determines what one does in cancer research, as in all else, it is as well to think something. And it may prove worthwhile to think that one or more tumors of FRAN CIS PEYTON ROUS 283 unknown causes are due to viruses" (1936~. He thus recognized that the problem that he so clearly formulated and actively pursued eluded experimental attack and remained unsolved. In fact he later restated the basic question: "What is the papilloma doing in the cancer, if anything?" (1940~.

In a renewed effort to answer this question, Peyton Rous used as a new tool the famous line of transplantable rabbit cancers, derived from a viral papilloma called at first "carcin- oma V2" (1940), and then, after World War II, V X 2 because during the war V2 "came to have another significance" (1952~. This line did not contain infectious papilloma virus, but for many serial transfers in rabbit it continued to elicit the produc- tion of virus-specific antibody. The result suggested that the virus may play a determining role although in "masked or altered form" (1940~. This was a new idea in virology, which had enormous developments many years later. For the next three years, during serial transplanta- tion from one rabbit to another, the V X 2 carcinoma continued to elicit this immune response.

However, when it was retested after an interval of one and a half years, four and a half years after its origin, the tumor was found unable to immunize against the papilloma virus; the loss of this property "was not attended by any perceptible change in the V X 2 carcinoma" (1952~. This "wholly unexpected" result must have been quite shattering; and Peyton Rous was led to rethink the role of the virus in the production of the cancer. In this agonizing re- appraisal he proposed that the virus might have undergone "wider variation" (1952~; but he recognized that "at this un- certain point the problem of the cause for the V X 2 carcinoma must perforce be left" (1952~. In this way the work of Peyton Rous went full circle: from complete ignorance on the role of viruses in cancer to definitely establishing such a role through brilliant discoveries, to postulating a wider and possibly general role of viruses in spontaneous cancers, and ending up again in 284 BIOGRAPHICAL MEMOIRS a condition of uncertainty.

I should not say full circle, but rather one turn of the helix, because the uncertainty was now of a different kind. The emphasis of Peyton Rous's work in the forties and fifties shifted from the viruses to chemical carcinogens. Many articles were dedicated to the potentiating effect of tar and other car- cinogens on virus-induced papillomas. During this work it also became clear that tar alone induces papillomas very similar to those induced by the virus on normal skin or on skin pretreated by tar. In all cases the growth showed progression, i.e., remained benign for some time and then developed into carcinomas, which arose in a few isolated cells. However, many observations also showed that the role of the virus and of the chemicals was different: "The generality of the carcinogens bring about tissue conditions out of which tumors may or may not arise for reasons still undetermined. They may' tee fitly called provocative car- cinogens. The viruses, on the other hand, both initiate tumors and determine their character and behaviour. They are actuat- ing carcinogens" (1943~.

In a new series of experiments, Peyton Rous convincingly demonstrated that the viral and the chemical agents have a cooperative action, producing in combination cancers at much higher frequency and after shorter time than either agent alone. On the basis of this cooperation, Peyton Rous made three important suggestions. One bears on the mechanism of carcino- genesis. He proposed that in utero or at a young age the human or animal body becomes invaded by viruses that "would give no sign of their presence in most instances.... But if a provoca- tive carcinogen happened to work on the cells with which such a virus was associated . . . it might undergo variation and . . . give rise to a tumor. The new pathogenic variant would not be transmitted to other animals . . . but would be a dead-end virus, though the harmless source virus liable to the same or other variation would be passed on" (1943~. This hypothesis is very FRAN CIS PEYTON ROUS 285 similar to some prevalent at the time of this writing, if the dead-end variant is interpreted as a defective integrated provirus that has incorporated a silent oncogene, causing its expression. Another suggestion was that chemical carcinogens might cause the so-called spontaneous tumors.

He added, "A list of human tumors which have been traced to the action of provoca- tive carcinogens is in no small degree a sociological document, reflecting as it does the ways of life, vocation, avocations, habits and environmental stresses of people and individuals" (1943~. The third suggestion was that viruses and chemicals in com- bination might have a continued role in spontaneous cancer: "The recent discovery that viruses of some sorts lie latent for long periods, causing disease only on special occasion, coupled with the realization that some tumors have viruses as their cause, has led to a supposition already mentioned that agents of this sort may reside in animal tissues, perhaps throughout the life- time of the organism, doing no harm unless the cells with which they are associated undergo special pathological changes, when they undergo variation as a result of the new, abnormal milieu and render the cells neoplastic. According to this supposition, tar and methylcholanthrene are carcinogens because they alter the environment of viruses...." (1944~. Such a possibility is very much in the minds of virologists and oncologists today.

During his work on carcinogenic hydrocarbons, Peyton Rous identified important features of the neoplastic process they initiate. One is that their cancer-inducing activity is greatly enhanced by promoters that stimulate cell proliferation, for instance, wounds. Another feature is that "cancers arise by a step-like progression" (1941~. Peyton Rous recognized the im- portance of this observation, because "the cells of not a few tumors attain to their worst by further neoplastic changes which are scarcely less significant than the one primarily responsible for their state. Indeed the practical significance of these changes is often greater as meaning death to the patient" (1955~. 286 BIOGRAPHICAL MEMOIRS In his later years Peyton Rous continued his experimental work, but his contributions declined in number and relevance. During that time tremendous changes were occurring in biol- ogy, especially the great development of genetics and the birth of molecular biology. Although Peyton Rous showed great interest in these developments, he failed to assimilate them. Obviously, even a brilliant mind is subject to the limitations of age. He became attached to old concepts.

The main conse- quence was his rejection of somatic mutations as a possible cause of cancer and his failure to recognize viruses as new genetic material in the cells they infect. It may be said about him what he said about Leo Loeb: "He outlived his era of discoveries about cancer but what he did for science endures" (1960~. The work on cancer is the big basis on which Peyton Rous's fame rests. The other work, which I already mentioned, is permeated by a similar perceptiveness, imagination, and experi- mental ability. I should mention especially the work on blood preservation and substitutes, which Peyton Rous carried out during World War I, because it shows another facet of his personality, i.e., the ability to respond to urgent medical needs of society. For instance, in 1918 he wrote: "There exists at present a great and urgent need for an injection fluid that can be satisfactorily employed instead of blood for transfusion in cases of hemorrhage.

It is common knowledge that casualty clearing stations, after a 'push', are crowded with men who have lost too much blood to be operated on, who cannot be revived by means of salt solutions and supportive measures, but who would undoubtedly respond to transfusion. For the latter nei- ther time nor donors are available." Quite rapidly, at the be- ginning of the war, Peyton Rous and associates perfected a method for storing human red blood cells, using a weak gelatin solution to protect them during washing and sugars to preserve them. The procedure was used to establish the first blood bank FRANCIS PEYTON ROUS 287 during the war, which was operated by one of Rous's collabora- tors. And the solution for suspending the red cells, known as the Rous-Turner solution, is still in use. Such an accomplishment would be a sufficient reason for fame, because it has both scien- tific and humane values. We know that Peyton Rous was very proud of it. In addition to his experimental work, Peyton Rous had an- other absorbing interest: the editorship of the Journal of Ex- perimental Medicine. It is known that he dedicated to it an immense amount of time and energy.

He was reputed for the accuracy of his editing, both in regard to scientific content and style. I well remember when I, as a prospective author, first encountered him as an editor. He returned my manuscript with many remarks, mostly of style. I remember I was at first baffled, but then, after studying his comments further, came to appre- ciate their reasons, which went beyond the mere words. I realized that for him a word, every word, was a concept, which should be examined not only in its present, but also future, context. He reminded me in his letter that I should think that a certain word might become widely adopted and that I should therefore choose it with deliberate care. He was anticipating the flooding of scientific literature with laboratory slang, which has happened in recent times and which was a trend of which he strongly disapproved. But during his editorship, he succeeded in maintaining the Journal of Experimental Medicine at a high level, both in purity of language and strength of content. Peyton Rous remains in the minds of those who knew him, especially the younger generation, as the image of a man fully dedicated to his work, a scientist with vision, a strong although kind person, with a good sense of humor.

His experiments were always designed to test hypotheses and developed in a logical sequence on the basis of results already secured; they were very methodical and thorough and were reported in detail with ex- 288 BIOGRAPHICAL MEMOIRS treme clarity. He was independent in thought and was against conventional beliefs unsubstantiated by evidence. For instance, he commented: "Not so long ago in the dark ages of medicine, one could think nearly anything about disease because one knew almost nothing. Theoretical system succeeded system, from humours to homeopathy. Opinions strongly held appeared like realities and were acted upon as such. Now for most diseases all this is at an end: fact has killed fancy.... The tumor problem is the last stronghold of metaphysics in medicine" (1936). Although during his career he formulated some penetrating hypotheses not amenable to direct test, he was fundamentally interested in facts: "No explanation of the cause of cancer is worthy of attention that cannot be tested" (1932~. The language of his reports was vivid, full of images from everyday life, as shown by a few examples: "The fowl limps and its wings seem stiff" (1913~.

"The growth gives it (the fowl) a factitious plumpness" (1913~. "During the outward extension of the membrane (of the Kupffer cells), lava-like flows can be seen on its surface, when the light is cut down, and at its edges fimbriated or 'petaloid' extrusions, at times appearing whip-like, which are in constant slow motion" (1934~. Comparing his work with the chicken viruses, which started with a cancer and led to a virus, and with the Shope virus, which started with a virus and led to a cancer: "The trails have met at the same look-out. What does one see from this?" (1936~. His humor was sometimes biting: Speaking of new reactions elicited in the human body by surgery, he saw their positive aspects: "All that surgery has done in such instances is to make plain the relation of effect to cause, as for example in showing that tetanus is due to insufficiency of the parathyroids, and myoxedema to a thyroid lack" (1929~.

FRAN CIS PEYTON ROUS 289 The most visible side of Peyton Rous was his interest in scientific truth, in the younger people, the equanimity of judg- ment and the warmth of human relations he was able to estab- lish both inside his family and outside. I had a hint of this when recently his daughter, Marion, referred to him as "daddy" in an affectionate way; and when several years ago he wrote to me sending his congratulations for the Ehrlich Darmstaedter award (which he nominated me for) and suggesting the nicest Rat- skeller in Frankfurt, with the best food, wine, and atmosphere. Peyton Rous received many official honors in his life, includ- ing the highest. But the paramount recognition was the admira- tion and respect of his younger colleagues, which continues after his death.

290 BIOGRAPHICAL MEMOIRS HONORS AND DISTINCTIONS Honorary degrees from the universities of Cambridge, Michigan, Yale, Birmingham, McGill, Chicago, and Zurich Member of the National Academy of Sciences Member of the American Philosophical Society Foreign Member of the Royal Society Member of the Royal Danish Academy of Sciences National Medal of Science Cleveland Medal of the American Cancer Society Gold headed cane of the Association of Pathologists and Bacteriolo- g~sts United Nations Prize Gold Medal of the Royal Society of Medicine Albert Lasker Award Landsteiner Award of the American Society of Blood Banks Distinguished Service Award of the American Cancer Society Kovalenko Award of the National Academy of Sciences Ehrlich Darmstaedter Prize Kober Medal of the Association of American Physicians Benter Medal and Award of the University of Texas Walker Prize of the Royal College of Surgeons John Scott Medal and Award of the City of Philadelphia Nobel Prize for Medicine (shared with Charles Huggins) FRANCIS PEYTON ROUS BIBLIOGRAPHY 291 KEY TO ABBREVIA TIONS Am. J. Med. Sci. = American Journal of Medical Sciences Cancer Res. = Cancer Research I. Am. Med. Assoc.

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Med., 10: 238-70. The effect of pilocarpine on the output of lymphocytes through the thoracic duct. l. Exp. Med., 10: 329~2. Some differential counts on the cells in the lymph of the dog: their bearing on problems in haematology. l. Exp. Med., 10:537-47. The teaching of physiological pathology at the University of Michi- gan. Johns Hopkins Hospital Bulletin, 19:336-38. 1909 The resistance to a specific hemolysin of human erythrocytes in health and disease. i. Exp. Med., 11 :763-85. Parabiosis as a test for circulating antibodies in cancer. l. Exp. Med., 11 :810-14. 292 BIOGRAPHICAL MEMOIRS On the reaction of tumor mice to the injections of tumor emulsion. Zeitschrift fur Immunitatsforschung und Experimentelle Thera- pie, 4:238~2. 1910 The fate of embryonic tissue implanted in the mother.

Proc. Soc. Exp. Biol. Med., 7:71-72. The behaviour of transplanted mixtures of tumor and embryo. Proc. Soc. Exp. Biol. Med., 7:73-74. The experimental production of secondary union between normal and carcinomatous epithelium pseudometaplasia. J. Am. Med. Assoc., 54: 1939. An experimental comparison of transplanted tumor and a trans- planted normal tissue capable of growth. i. Exp. Med., 12:344- 705. A transmissible avian neoplasm (sarcoma of the common fowl). l. Exp. Med., 12:696-705. Metastasis and tumor immunity. Observations with a transmissible avian neoplasm. l. Am. Med. Assoc., 55: 1805-15. 1911 Transmission of a malignant new growth by means of a cell-free filtrate. J. Am. Med. Assoc., 56: 198. The relations of embryonic tissue and tumor in mixed grafts.

l. Exp. Med., 13:239~7. The effect of pregnancy on implanted embryonic tissue. J. Exp. Med., 13: 248-62. With J. B. Murphy. Tumor implantations in the developing em- bryo. Experiments with a transmissible sarcoma of the fowl. I. Am. Med. Assoc., 56:741~2. The rate of tumor growth in underfed hosts. Proc. Soc. Exp. Biol. Med., 8: 128-30. A sarcoma of the fowl transmissible by an agent separable from the tumor cells. J. Exp. Med., 13:397~11. 1912 With J. B. Murphy and W. H. Tytler. Transplantable tumors of the fowl; a neglected material for cancer research. J. Am. Med. Assoc., 58:1682-83. FRANCIS PEYTON ROUS 293 With l. B. Murphy and W. H. Tytler.

The role of injury in the production of a chicken sarcoma by a filterable agent. l. Am. Med. Assoc., 58: 1751. With I. B. Murphy and W. H. Tytler. The relation between a chicken sarcoma's behaviour and the growth's filterable cause. I. Am. Med. Assoc., 58: 1840-41. With l. B. Murphy. The nature of the filterable agent causing a sarcoma of the fowl. l. Am. Med. Assoc., 58:1938. An avian tumor in its relation to the tumor problem. Proceedings of the American Philosophical Society, 51 :201-5. With i. B. Murphy. The behaviour of chicken sarcoma implanted in the developing embryo. J. Exp. Med., 15: 119-32. With J. B. Murphy. The histological signs of resistance to a trans- missible sarcoma of the fowl. l. Exp. Med., 15:270-86. With l. B. Murphy and W. H.

Tytler. A filterable agent the cause of a second chicken-tumor: an osteochondrosarcoma. l. Am. Med. Assoc., 59: 1793-94. 1913 With i. B. Murphy. Variations in a chicken sarcoma caused by a filterable agent. J. Exp. Med., 17:219-31. False transitions between normal and cancerous epithelium. I. Exp. Med., 17:494-96. The growth of tissue in acid media. J. Exp. Med., 18:183-86. Resistance to a tumor-producing agent as distinct from resistance to the implanted tumor cells. Observations with a sarcoma of the fowl. J. Exp. Med., 18:416-27. With Linda B. Lange. The characters of a third transplantable chicken tumor due to a filterable cause. A sarcoma of intra- canalicular pattern. J. Exp. Med., 18:651-64. 1914 With i. B. Murphy.

On the causation by filterable agents of three distinct chicken tumors. i. Exp. Med., 18:52-69. On certain spontaneous chicken tumors as manifestations of a single disease. I. Spindle-celled sarcomata rifted with blood sinuses. J. Exp. Med., 19:570-76. With F. S. Jones. On the cause of the localization of secondary tu- mors at points of injury. J. Exp. Med., 20:404-12. 294 B I O G RA P H I C A L M E M O I R S With Linda B. Lange. On the greater susceptibility of an alien variety of host to an avian tumor. l. Exp. Med., 20:413-18. With J. B. Murphy. On immunity to transplantable chicken tumors. J. Exp. Med., 20:419-32. The influence of diet on transplanted and spontaneous mouse tu- mors. l. Exp. Med., 20:433-51. 1915 With l. R. Turner.

The resistance to mechanical injury of the erythrocytes of different species. Proc. Soc. Exp. Biol. Med., 12: 106-7. With J. R. Turner. The protection of fragile erythrocytes against mechanical injury. Proc. Soc. Exp. Biol. Med., 12:107-8. With l. R. Turner. On the preservation in vitro of living erythro- cytes. Proc. Soc. Exp. Biol. Med., 12: 122-24. With I. R. Turner. A rapid and simple method of testing donors for transfusion. i. Am. Med. Assoc., 64:1980-86. 1916 With .T. R. Turner. The preservation of living red blood cells in vitro. I. Methods of preservation. l. Exp. Med., 23:219-37. With i. R. Turner. The preservation of living red blood cells in vitro. II. The transfusion of kept cells. l. Exp. Med., 23:239-48.

With F. S. tones. A method of obtaining suspensions of living cells from the fixed tissues, and for the plating out of individual cells. J. Exp. Med., 23:549-55. With F. S. ~ones. The protection of pathogenic micro-organisms by living tissue cells. J. Exp. Med., 23:601-12. 1917 With F. S. ~ones. The phagocytic power of connective tissue cells. J. Exp. Med., 25: 189-93. With O. H. Robertson. The normal fate of erythrocytes. I. The findings of healthy animals. J. Exp. Med., 25:651-63. With O. H. Robertson. The normal fate of erythrocytes. II. Blood destruction in plethoric animals with a simple anemia. l. Exp. Med., 25: 665-73. FRANCIS PEYTON ROUS 1918 295 With G. W. Wilson. Fluid substitutes for transfusion after hemor- rhage.

First communication. l. Am. Med. Assoc., 70:219-22. Method for intravenous injection of guinea-pigs. l. Exp. Med., 27: 459-62. With O. H. Robertson. Free antigen and antibody circulating to- gether in large amounts (hemagglutinin and agglutinogen in the blood of transfused rabbits). I. Exp. Med., 27:509-17. With O. H. Robertson. Autohemagglutination experimentally in- duced by the repeated withdrawal of blood. J. Exp. Med., 27: 563-68. With Jean Oliver. Experimental hemochromatosis. i. Exp. Med., 28: 629~4. Urinary siderosis. Hemosiderin granules in the urine as an aid in the diagnosis of pernicious anemia, hemochromatosis, and other diseases causing siderosis of the kidney. l. Exp. Med., 28:645-58. 1919 With G. W. Wilson.

The influence of ether anesthesia, of hemor- rhage, and of plethora from transfusion on the presser effect of minute quantities of epinephrine. J. Exp. Med., 29:173-86. With O. H. Robertson and Jean Oliver. Experiments on the pro- duction of specific antisera for infections of unknown cause. I. Type experiments with known antigens a bacterial hemotoxin (megatheriolysin), the pneumococcus, and poliomyelitis virus. II. The production of a serum effective against the agent causing a chicken sarcoma. I. Exp. Med., 29:283-304, 305-20. 1920 With G. W. Wilson and Jean Oliver. Experiments on the production of specific antisera for infections of unknown cause. III. The effects of a serum precipitin on the animals of the species furnish- ing the precipitinogen. J. Exp. Med., 31:~53-65. With L. D. Larimore. The relation of the portal blood to liver main- tenance.

A demonstration of liver atrophy conditional on com- pensation. l. Exp. Med., 31:609-32. With L. D. Larimore. The relation of the portal blood to liver maintenance. Proc. Soc. Exp. Biol. Med., 17:65-66. With P. D. McMaster. A vicious activity of the gall bladder during 296 BIOGRAPHICAL MEMOIRS biliary stasis. B. The determining {actor in the causation of white stasis bile. Proc. Soc. Exp. Biol. Med., 17:159. With L. D. Larimore. The biliary factor in liver lesions l. Exp. Med., 32:249-72. With P. D. McMaster. The concentrating activity of the gall bladder. Proc. Soc. Exp. Biol. Med., 17:215. 1921 With P. D. McMaster. The biliary obstruction required to produce jaundice. i. Exp. Med., 33:731-50. With P. D. McMaster.

The concentrating activity of the gall bladder. J. Exp. Med., 34:47-73. With P. D. McMaster. Physiological causes for the varied character of stasis bile. l. Exp. Med., 34:75-95. 1922 With O. H. Robertson. Sources of the antibodies developing after repeated transfusion. l. Exp. Med., 35:141-52. With P. D. McMaster and L. D. Larimore. Significance of the hemo- siderosis of pernicious anemia. l. Exp. Med., 35:521-31. With H. Haessler and G. O. Broun. The renal elimination of bili- rubin. l. Exp. Med., 35: 533-52. With O. H. Robertson. Lasting individual differences in the resis- tance of normal bloods to shaking. Proc. Soc. Exp. Biol. Med., 20: 37-38. With P. D. McMaster and G. O. Broun.

The experimental produc- tion of gall stones in dogs, in the absence of infection, stasis and gall bladder influence upon the bile. Proc. Soc. Exp. Biol. Med., 20: 128-38. 1923 With P. D. McMaster. A method for the permanent sterile drainage of intra-abdominal ducts, as applied to the common duct. I. Exp. Med., 37:11-19. Destruction of the red blood corpuscles in health and disease. Physiological Reviews, 3: 75-99. With P. D. McMaster and G. O. Broun. Studies on the total bile. I. FRAN CIS PEYTON ROUS 297 The effects of operation, exercise, hot weather, relief of obstruc- tion, intercurrent disease, and other normal pathological influ- ences. J. Exp. Med., 37: 395-420. With G. O. Broun and P. D. McMaster. Studies on the total bile. II. The relation of carbohydrates to the output of bile pigment. l. Exp.

Med., 37:421-29. With P. D. McMaster and G. O. Broun. Studies on the total bile. III. On the bile changes caused by a pressure obstacle to secretion; and on hydrohepatosis. i. Exp. Med., 37:685-98. With G. O. Broun and P. D. McMaster. Studies on the total bile. IV. The enterohepatic circulation of bile pigment. I. Exp. Med., 37: 699-710. With P. D. McMaster and D. R. Drury. The genesis of gall stones in the dog. Proc. Soc. Exp. Biol. Med., 20:315-18. With G. O. Broun and P. D. McMaster. Studies on the total bile. V. The relation between blood destruction and output of bile pig- ment. l. Exp. Med., 37:733-57. 1924 With P. D. McMaster and D. R. Drury.

Observations on some causes of gall stone formation. I. Experimental cholelithiasis in the absence of stasis, infection, and gall bladder influences. l. Exp. Med., 39:77-96. With I). R. Drury and P. D. McMaster. Observations on some causes of gall stone formation. II. On certain special nuclei of deposi- tion in experimental cholelithiasis. l. Exp. Med., 39:97-116. With D. R. Drury and P. D. McMaster. Observations on some causes of gall stone formation. III. The relation of the reaction of the bile to experimental cholelithiasis. l. Exp. Med., 39:403~5. With P. D. McMaster. The liver requirement of the fasting or- ganism. J. Exp. Med., 39:425-45. The relative reaction of living mammalian tissues. Science, 60:363. 1925 The relative reaction within living mammalian tissues. I. General features of vital staining with litmus. l. Exp.

Med., 41:379-97. The relative reaction within living mammalian tissues. II. On the mobilization of acid material within cells, and the reaction as influenced by the cell state. l. Exp. Med., 41 :399-411. 298 BIOGRAPHICAL MEMOIRS The relative reaction within living mammalian tissues. IlI. Indi- cated differences in the reaction of the blood and tissues on vital staining with phthaleins. T. Exp. Med., 41 :451-70. With D. R. Drury. jaundice as art expression of the physiological wastage of corpuscles. l. Exp. Med., 4 1: 60 1-9. With D. R. Drury. Suppression of bile as a result of impairment of liver function. J. Exp. Med., 41:611-22. The relative reaction within living mammalian tissues. IV. Indi- cated differences in the reaction of the organs on vital staining with phthaleins. l. Exp. Med., 41:739-59. With D.

R. Drury. Outlying acidosis. i. Am. Med. Assoc., 85:33-35. The biliary aspects of liver disease. Am. l. Med. Sci., 170:625. 1926 With D. R. Drury. The relative reaction within living mammalian tissues. V. (a) Influence of lymph-soluble tissue materials on the significance of the coloration with some phthalein indicators. V. (b) Influence of lymph-insoluble tissue materials on the signifi- cance of the coloration with some phthalein indicators. J. Exp. Med., 43:669-86, 687-701. The relative reaction within living mammalian tissues. VI. Factors determining the reaction of skin grafts; a study of the indicator method of conditions within an ischemic tissue. l. Exp. Med., 44:815-34. With W. W. Beattie. The relative reaction within living mammalian tissues. VII. The influence of changes in the reaction of the blood upon the reaction of the tissues. l. Exp.

Med., 44:835-54. 1927 With D. R. Drury and W. W. Beattie. The relative reaction within living mammalian tissues. VIII. On the course of the tissue aci- dosis secondary to blood acidosis induced with hydrochloric acid. l. Exp. Med., 45:23-39. With D. R. Drury and W. W. Beattie. The relative reaction within living mammalian tissues. IX. On the tissue reaction as influ- enced by inhalations of CO2 and by overbreathing. J. Exp. Med., 45:41-58. Pathology and the glare of the future. Reprinted from Contributions to Medical Science (Dedicated to Aldred Scott Warthin), ed. by Willard J. Stone, pp. 19-22. Ann Arbor, Mich.: George Wahr. FRANCIS PEYTON ROUS 1929 299 With D. R. Drury. Outlying acidosis due to functional ischemia. l. Exp. Med., 49:435-60.

The Modern Dance of Death. (The Linacre Lecture) London: Cambridge Univ. Press. 51 pp. With H. P. Gilding. The meaning of Bier's spots. Proc. Soc. Exp. Biol. Med., 26:497-98. With H. P. Gilding. Studies of tissue maintenance. I. The changes with diminished blood bulk. J. Exp. Med., 50:189-211. With H. P. Gilding. Studies of tissue maintenance. III. Persisting bloodlessness after functional ischemia. l. Exp. Med., 50:471-87. With H. P. Gilding. The final response of the small cutaneous ves- sels. l. Exp. Med., 50:489-512. 1930 With H. P. Gilding. Is the local vasodilatation after different tissue injuries referable to a single cause? J. Exp. Med., 51:27-39~. With H P. Gilding and F. Smith. The gradient of vascular perme- ability. J.

Exp. Med., 51: 807-30. 1931 With F. Smith. The gradient of vascular permeability. II. The con- ditions in frog and chicken muscle, and in the mammalian dia- phragm. J. Exp. Med., 53:195-217. With F. Smith. The gradient of vascular permeability. III. The gradient along the capillaries and venules of frog skin. l. Exp. Med., 53:219-42. With F. Smith. The gradient of vascular permeability. IV. The per- meability of the cutaneous venules and its functional significance. J. Exp. Med., 54:499 514. 1932 With P. D. McMaster and S. S. Hudack. The relation of hydrostatic pressure to the gradient of capillary permeability. l. Exp. Med., 55:203-21. With E. Botsford. The incidence of cancer in tarred and sheltered mice. J. Exp.

Med., 55:247-66. 300 BIOGRAPHICAL MEMOIRS 1933 With P. D. McMaster and S. S. Hudack. The fixation of certain viruses on the cells of susceptible animals and the protection afforded by such cells. Proc. Soc. Exp. Biol. Med., 31:90-91. 1934 Withy. W. Beard. The neoplastic traits of a mammalian growth due to a filterable virus the Shope rabbit papilloma. Science, 79: 437-38. With i. W. Beard. Selection with the magnet and cultivation of reticuloendothelial cells (Kupffer cells). J. Exp. Med., 59:577-91. With i. W. Beard. The characters of Kupffer cells living in vitro. J. Exp. Med., 59~:593-607. With l. W. Beard. A virus-induced mammalian growth with the characters of a tumor (the Shope rabbit papilloma). I. The growth on implantation within favorable hosts.

l. Exp. Med., 60:701-22. With I. W. Beard. A virus-induced mammalian growth with the characters of a tumor (the Shope rabbit papilloma). II. Experi- mental alterations of the growth on the skin: morphological considerations: the phenomena of retrogression. J. Exp. Med., 60:723-37. With i. W. Beard. A virus-induced mammalian growth with the characters of a tumor (the Shope rabbit papilloma). III. Further characters of the growth: general discussion. l. Exp. Med., 60: 741-66. 1935 With l. W. Beard. Carcinomatous changes in virus-induced papillo- mas of the skin of the rabbit. Proc. Soc. Exp. Biol. Med., 32: 578-80. With P. D. McMaster and S. S. Hudack. The fixation and protection of viruses by the cells of susceptible animals. i. Exp. Med., 61: 657-88. With i. W. Beard.

The progression to carcinoma of virus-induced rabbit papillomas (Shope). J. Exp. Med., 62:523~8. With l. W. Beard. Effectiveness of the Shope papilloma virus in vari- ous American rabbits. Proc. Soc. Exp. Biol. Med., 33:191-93. - FRANCIS PEYTON ROUS 301 With John G. Kidd and l. W. Beard. Certain factors determining the course of virus-induced tumors. Proc. Soc. Exp. Biol. Med., 33: 193-95. With J. W. Beard. A comparison of the tar tumors of rabbits and the virus-induced tumors. Proc. Soc. Exp. Biol. Med., 33:358-60. 1936 With l. G. Kidd. The carcinogenic effect of a virus upon tarred skin. Science, 83:468-69. With l. G. Kidd and l. W. Beard. Serological reactions with a virus causing rabbit papillomas which become cancerous. I.

Tests of the blood of animals carrying the papilloma. l. Exp. Med., 64: 63-77. Aldred Scott Warthin. In: Dictionary of American Biography, vol. 19, pp. 493-94. New York: Charles Scribner & Sons. With l. G. Kidd and i. W. Beard. Serological reactions with a virus causing rabbit papillomas which become cancerous. II. Tests of the blood of animals carrying various epithelial tumors. J. Exp. Med., 64:79-96. With J. G. Kidd and l. W. Beard. Observations on the relation of the virus causing rabbit papillomas to the cancers deriving therefrom. I. The influence of the host species and of the pathogenic activity and concentration of the virus. J. Exp. Med., 64:385-400. With l. W. Beard and I. G. Kidd. Observations on the relation of the virus causing rabbit papillomas to the cancers deriving therefrom. II. The evidence provided by the tumors; general considerations. J. Exp.

Med., 64:401-24. The virus tumors and the tumor problem. (Harvey Lecture) Ameri- can Journal of Cancer, 28:233-71. 1937 With J. G. Kidd. Effect of the papilloma virus (Shope) upon the tar warts of rabbits. Proc. Soc. Exp. Biol. Med., 37: ~ 18-20. 1938 With l. G. Kidd. The carcinogenic effect of a papilloma virus on the tarred skin of rabbits. I. Description of the phenomenon. l. Exp. Med., 67:399~28. 302 BIOGRAPHICAL MEMOIRS With l. W. Beard. The fate of vaccinia virus on cultivation in vitro with Kupffer cells (reticulo-endothelial cells). J. Exp. Med., 67: 883-910. With l. G. Kidd. The carcinogenic effect of a papilloma virus on the tarred skin of rabbits. II. Major factors determining the phenomenon; the manifold effects of tarring.

l. Exp. Med., 68: 529-61. 1939 With l. G. Kidd. A comparison of virus-induced rabbit tumors with the tumors of unknown cause elicited by tarring. i. Exp. Med., 69: 399-424. 1940 With J. G. Kidd. Cancers deriving from the virus papillomas of wild rabbits under natural conditions. l. Exp. Med., 71:469-93. With i. G. Kidd. The activating, transforming, and carcinogenic effects of the rabbit papilloma virus (Shope) upon implanted tar tumors. l. Exp. Med., 7 1: 787-8 1 1 . With J. G. Kidd. A transplantable rabbit carcinoma originating in a virus-induced papilloma and containing the virus in masked or altered form. l. Exp. Med., 71 :813-37. 1941 With J. G. Kidd. Conditional neoplasms and subthreshold neo- plastic states. A study of the tar tumors of rabbits. J. Exp.

Med., 73:365-89. With Ian MacKenzie. The experimental disclosure of latent neo- plastic changes in tarred skin. J. Exp. Med., 73:391~15. The conditions determining cancer. (The William Henry Welch Lecture, I) J. Mt. Sinai Hosp., 8:184 - 85. The known causes of cancer. (The William Henry Welch Lecture, II) J. Mt. Sinai Hosp., 8: 186. With W. F. Friedewald. The carginogenic effect of methylcholan threne and of tar on rabbit papillomas due to a virus. Science 94:495-96. FRAN CIS PEYTON ROUS 1943 303 Viruses and tumors. In: Virus Diseases, p. 147. Ithaca, N.Y.: Cornell Univ. Press. The nearer causes of cancer. i. Am. Med. Assoc., 122:573-84. 1944 With W. F. Friedewald.

The effect of chemical carcinogens on virus- inducedrabbit papillomas. l. Exp. Med., 79:511-37. With W. F. Friedewald. The initiating and promoting elements in tumor production. An analysis of the effects of tar, benzpyrene, and methylcholanthrene on rabbit skin. J. Exp. Med., 80: 101-25. With W. F. Friedewald. The determining influence of tar, benzpy- rene, and methylcholanthrene on the character of the benign tumors induced therewith in rabbit skin. l. Exp. Med., 80: 127~4. lg45 With W. E. Smith. The neoplastic potentialities of mouse embryo tissues. I. The findings with skin of C strain embryos transplanted to adult animals. l. Exp. Med., 81:597-619. The neoplastic potentialities of mouse embryo tissues. II. Contribu- tory experiments; results with skin of C3H and Webster-Swiss embryos; general considerations. J. Exp.

Med., 81:621~5. 1946 The activation of skin grafts. l. Exp. Med., 83:383-99. Concerning the cancer problem. American Scientist, 34 (July>:329. Simon Flexner and the Journal of Experimental Medicine. J. Exp. Med., 84(1):i. 1947 Recent advances in cancer research. Bulletin of the New York Academy of Medicine, 23:65-78. The lamentable decline in self-satisfaction. Proceedings of the Cha- raka Club, 11 :7. Karl Landsteiner, 1868-1943. Obit. Not. Fell. R. Soc. Lond., 5:295- 312. 304 BIOGRAPHICAL MEMOIRS 1948 Simon Flexner and medical discovery. Science, 107:611-13. With W. E. Smith. The neoplastic potentialities of mouse embryo tissues. IV. Lung adenomas in baby mice as result of prenatal exposure to urethane. i.

Exp. Med., 88:529-54. 1949 Robert Gladding Green~41895-1947~. Science, 109:93. Simon Flexner, 1863-1946. Obit. Not. Fell. R. Soc. Lond., 6:409-30. 1950 With W. F. Friedewald. The pathogenesis of deferred cancer. A study of the after-effects of methylcholanthrene upon rabbit skin. J. Exp. Med., 91:459 83. 1951 With S. Rogers. Joint action of a chemical carcinogen and a neoplas- tic virus to induce cancer in rabbits. Results of exposing epi- dermal cells to a carcinogenic hydrocarbon at time of infection with the Shope papilloma virus. i. Exp. Med., 93:459-88. 1952 With S. Rogers. The occurrence in tarred rabbit skin of minor, almost imperceptible, neoplastic changes.

Cancer Res., 12:292. (A) With W. E. Smith and J. G. Kidd. Experiments on the cause of the rabbit carcinomas derived from virus-induced papillomas. I. Propagation of several of the cancers in sucklings, with etiologi- cal tests. J. Exp. Med., 9~5:299-317. What cancer research has done. The Listener, 48: 138-42 (issue dated 24 July). With J. G. Kidd and W. E. Smith. Experiments on the cause of the rabbit carcinomas derived from virus-induced papillomas. II. Loss by the V X 2 carcinoma of the power to immunize hosts against the papilloma virus. J. Exp. Med., 96:159-74. Cancer research as viewed along the years. Acta Physiologica Latino- americana, 3: 18~87. FRANCIS PEYTON ROUS 1955 305 With K. Dumbell.

Are carcinogens responsible for the superimposed neoplastic changes occurring in mouse tumor cells? The effect of methylcholanthrene and urethane on pulmonary adenomas and of methylcholanthrene on mammary carcinomas. l. Exp. Med., 102:517-43. 1958 With R. A. Allen. Fatal keratomas due to deep homografts of the benign papillomas of tarred mouse skin. Normal proclivities and neoplastic disabilities as determinants of tumor course. l. Exp. Med., 107:63-85. 1959 Surmise and fact on the nature of cancer. Nature, Land., 183:1357- 61. 1960 Leo Loeb, 1869-1959. Cancer, 13: 3-4. The possible role of viruses in cancer. Opening remarks and sum- mary of informal discussions (first session). Cancer Res., 20:672- 76, 707-11. Charles Oberling, research worker on the nature of cancer.

Science, 132: 1534-35. 1961 The scope of carcinogenesis. Extrait de Acta Union Internationale Contre le Cancer, 17: 262-65. 1962 With l. S. Henderson. The plating of tumor components on the subcutaneous expanses of young mice: findings with benign and malignant epidermal growths and with mammary carcinomas. J. Exp. Med., 115: 1221-29. The disagreement amongst doctors. In: On Cancer and Hormones, p. 49. Chicago: Univ. of Chicago Press. Reprinted in Midway, no. 12:72-79. 306 BIOGRAPHICAL MEMOIRS 1964 Henry James and the mouse. Perspect. Biol. Med., 7:433-34. With l. S. Henderson. Further experiments on the cause of sequen- tial neoplastic changes. The effects of 20-methylcholanthrene on transplanted epidermal mouse papillomas and the derivative carcinomas. J.

Exp. Med., 120: 197-221. 1965 Viruses and tumour causation: an appraisal of Nature, Land., 207:457-63. present knowledge. The viruses and us. In: Perspectives in Virology, ed. by Morris Pol- lard, vol. 4, pp. 305-8. New York: Hoeber Medical Division, Harper & Row Pubs., Inc. A great friend: George de Hevesy. International Journal of Applied Radiation and Isotopes, 16:516-18. 1966 The lamentable decline in self-satisfaction. Perspect. Biol. Med., 9: 439-49. Biography and speeches at the Nobel Banquet. In: Les Prix Nobel en 1966. Stockholm: Kungl. Boktryckeriet P. A. Norstedt 8c Soner. - 1967 The fly in the social ointment. Perspect. Biol. Med., 11: 1-8. The challenge to man of the neoplastic cell.

Cancer Res., 27:1919, 1960; also in Science, 157: 24-28.

Question: What is the more familiar name of Samuel Peyton Jones, a character in an animated series for children?
Answer: Fireman Sam