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Biographical Memoirs: Volume 62

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Biographical Memoirs: Volume 62 MAX LUDWIG HENNING DELBRÜCK September 4, 1906-March 10, 1981 BY WILLIAM HAYES MAX DELBRÜCK, or just Max as he was called by all his associates, was one of the outstanding natural scientists of our time. A man of rare intellectual ability, and clarity of thought and perception, he excelled in theoretical physics, biology, and philosophy, and possessed a deep knowledge and appreciation of the arts. His dedication to truth, and his intolerance of half-truths and intellectual pretension, were sometimes expressed with a disturbing frankness and abruptness of manner, often construed as arrogance by those who did not know him well. His disclaimer, ''I don't believe a word of it," when told of some new experimental result or hypothesis, became famous among his colleagues. In fact, Max was very gregarious and had a rich vein of friendship and affection in his nature which he was always ready to share with others of all ages. Above all, Max was a born leader whose Socratic influence on those who worked with him was enormous, whose rare praise was something to be coveted and remembered, and whose criticism was welcomed with respect; although Reprinted from Biographical Memoirs, The Royal Society, London, England, 1982.

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Biographical Memoirs: Volume 62 he was often wrong in his scientific judgement, he was always the first to admit it. On a personal level he engendered in the minds of his friends and colleagues a deep respect and affection that they will not forget. Max was the foremost pioneer of a new approach to an understanding of fundamental biological processes, now known as molecular biology. His most significant studies concerned the multiplication in their host cells of bacterial viruses, called bacteriophages or phages for short. These tiny particles are made up of about equal parts of two chemical components, protein and nucleic acid; infection of a bacterium by a single particle is followed, about 30 minutes later, by rupture of the cell and liberation of a hundred or more progeny particles. As long ago as 1922 the American geneticist H.J. Muller had suggested phage as the simplest possible model for studying the nature and behaviour of genes. For their novel and important studies in this field, Max and his colleagues, Salvador Luria and Alfred Hershey, were awarded the Nobel Prize in Physiology or Medicine in 1969. However, no account of Max's published work can do justice to his overall influence as the leader of a formidable group of workers, many of them physicists like himself, who infused a new way of thinking, and a new life, into biological research. In addition, he was a direct source of encouragement and inspiration to young research workers of many nationalities and from many disciplines who came to work with him on bacteriophage at the California Institute of Technology, in Pasadena, California, or to attend his famous "Phage Course" at the Cold Spring Harbor Laboratory, Long Island, New York, and to whom his intellectual approach to biological problems became an inspiration for their own thinking.

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Biographical Memoirs: Volume 62 FAMILY BACKGROUND Max grew up in the Grunewald suburb of Berlin, the youngest of seven children (four girls and three boys) of an extremely prominent academic family. His father, Hans Delbrück, who was 58 years older than Max, was Professor of History at Berlin University, specializing in the history of the art of war, as well as sole editor for at least 30 years of a monthly journal, Preussische Jahrbücher, for which he wrote a column commenting on German politics. Three of his father's first cousins were, respectively, Professor of German Literature at Jena, Chief Justice of the Imperial Supreme Court, and Minister of State. His maternal great-grandfather was the famous Justus von Liebig, Professor of Chemistry at Giessen and Münich, Foreign Member of the Royal Society, and Copley Medalist. His mother's brother-in-law, Adolf von Harnack, was Professor of Theology at Berlin University and a church historian; he was also Director of the Prussian State Library and, in 1910, became co-founder and President of the Kaiser-Wilhelm-Gesellschaft. The Harnacks, the Delbrück's nearest relatives, were also a large family and lived next door, while Karl Bonhoeffer, a Professor of Psychiatry, and his family were around the corner and the Max Planck family not far away. One of the Bonhoeffer sons, Klaus, married Max Delbrück's sister Emmie. Max's family enjoyed "a modest degree of affluence and apparently the life until 1914 was pretty free and very hospitable. As war came and life became more and more of a nightmare in every respect, of course all this darkened …. I think three-quarters of the young men in the family [including his eldest brother] were killed. So that was all very sad, and in addition then there came these pretty severe food and coal shortages and then the total mess in 1918.

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Biographical Memoirs: Volume 62 So this relatively affluent residential suburb after the war became almost a ghost town" (1). World War II also brought tragedy to the Delbrück family. Two Bonhoeffer brothers, Klaus (Max's brother-in-law) and Pastor Dietrich, two Bonhoeffer sons-in-law, and two von Harnack cousins, Ernst and Arvid, together with the latter's American wife, were executed by the Nazis as leading members of the Resistance. Max's brother Justus was imprisoned by the Nazis, and liberated after the fall of Berlin but ten days later was arrested by the Russians and died in a diphtheria epidemic in a Russian camp. The husbands of two of Max's sisters also were killed by marauding soldiers in the last days of the war. EARLY INTEREST IN SCIENCE Of all the many children in the Delbrück, Harnack and Bonhoeffer families, Max was the youngest. Moreover, none of his intimates, save one, had any knowledge of, or interest in, science. The exception was Karl Friedrich Bonhoeffer, 8 years his senior, who became a distinguished physical chemist and Max's mentor and lifelong friend. Max's main boyhood interests were astronomy and mathematics. In retrospect, some 40 years later, he considered that he chose astronomy as a means of finding and establishing his own identity in an intimate society of so many able and strong personalities, all of them older than himself; but only he was an astronomer, and proclaimed himself one during his last 2-3 years at the Grunewald Gymnasium. He read popular books on the subject, was the enthusiastic possessor of a 2-inch telescope, and sometimes woke the whole household with the loudest of alarm clocks in the small hours of the morning when he had an appointment with the stars! (1) Despite Max's nuisance value, his parents proved tolerant and even helpful, while his

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Biographical Memoirs: Volume 62 knowledge of astronomy blossomed under the tutelage and friendship of Karl Friedrich Bonhoeffer. It thus became Max's intention to study astronomy at the university. In 1924, at the age of 171/2, he went first to Tübingen where Hans Rosenberg offered an introduction to astrophysics which was then in its infancy; he also took courses in mathematics and physics, but chemistry failed to attract him and he never learned this subject as a student. He spent only one semester at Tübingen and then moved for a semester to Berlin where he had free tuition because of his father's professorship there, and thence to Bonn and back to Berlin again until, in the summer of 1926, he finally settled at Göttingen for 3 years until he obtained his degree. Although Göttingen was at that time the center of excitement in theoretical physics, following Heisenberg's discovery of quantum mechanics in 1925, Max continued to be interested in astronomy and mathematics until his attempt to write a Ph.D. thesis on novae failed because, he admitted, the mathematics of astrophysical theory of the interior of stars was beyond him, while the relevant literature was in English which he did not know at the time. But in the effort he had had to learn a good deal of quantum mechanics which brought him into contact with some of the theoretical physicists, among them Max Born, Pasqual Jordan, Eugene Wigner and Walter Heitler. At this time he wrote a short paper (1929) providing formal mathematical proofs for a theorem that Wigner had used in the application of group theory to theoretical physics. Born, who was Professor of Theoretical Physics, thereupon offered Max a Teaching Assistantship, and Heitler suggested that he extend to lithium the quantum mechanical theory of the homopolar bond that had just been developed for

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Biographical Memoirs: Volume 62 hydrogen by Heitler and London. His conclusion was that the bond energy in Li2 is considerably smaller than in H2, not because of the repulsion of the K shells but because the bond electrons were two s electrons (1930a). Max recently averred that this topic turned out to be a nightmare for him because of the complexity of the mathematics involved and that he had never dared to look at his thesis again (1); but nevertheless it won him his Ph.D. Degree in 1930. EARLY CAREER IN PHYSICS (1929-32) Bristol John E. Lennard-Jones, Professor of Theoretical Physics at the new H. H. Wills Physics Laboratory, University of Bristol, England, spent some months at Göttingen in 1929 and was anxious to attract to Bristol two of Max Born's students for whom research grants had been provided. Gerhard Herzberg, then a postdoctoral Fellow, and Max Delbrüick were appointed. Max remained at Bristol for 18 months and became very friendly with Cecil F. Powell, with whom he roomed. Among other friends at that time were P. M. S. Blackett, later to become President of the Royal Society, P. A. M. Dirac and H. W. P. Skinner. Of these early associates four were later to win Nobel Prizes, three in physics (Dirac, Blackett and Powell) and one in chemistry (Herzberg). An unpublished history of the Bristol Department, written by the late Professor A. M. Tyndall, related that "M. Delbrück, Prussian by birth but cosmopolitan by nature, a theoretical physicist recommended by M. Born, brought with him intellectual stimulus, critical judgement and social entertainment which gave help and pleasure to many and sundry." Another member of the department at the time, who remembers him quite well (J. Burrow, quoted by

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Biographical Memoirs: Volume 62 N.T.)* describes him as a cheerful, outgoing person and one who rapidly established a reputation as a theoretician who was always ready to discuss problems of any kind with experimentalists who needed help and advice. Herzberg also recalled that he "fitted in very well with the group of younger physicists there because of his (then) gregarious ways and the ease with which he made friends." Max published two papers from Bristol in English (1930b; 1932), on topics related to the quantum mechanical theory of homopolar bonding on which he had written his thesis. Copenhagen and Zürich Following his Bristol experience, Max obtained a Rockefeller Fellowship (Physics) to study with Niels Bohr in Copenhagen where he spent the spring and summer of 1931, and then spent the last 6 months with another quantum physicist, Wolfgang Pauli, in Zürich. In Copenhagen he roomed, and collaborated on a nuclear physics project, with George Gamow (1931) with whom he established a lasting friendship. Also working with Bohr at that time was Victor Weisskopf, a very close friend since their student days together at Göttingen; they arrived in the United States almost simultaneously in 1937 (see below) and remained in personal contact until Max's death. Anyone who might infer from all this that life in Copenhagen was a staid and serious business should read Max's lighthearted and facetious account of the gaiety and practical jokes of those days, in his contribution to a George Gamow Memorial Volume (1972c). Weisskopf (pers. comm.) has commented on his wonderful sense of humor: "There was a custom in Copenhagen, at each of the early conferences *   The initials in the text that indicate the source of quotations are explained in the acknowlegements at the end of the memoir.

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Biographical Memoirs: Volume 62 organized by Niels Bohr, to have what we called a session of 'comic physics.' It was always Max who was the most spirited leader in these activities with his humour and intellectual fantasy. You must have heard of his rewriting of Goethe's Faust to make fun of the physics of that time." Max's short visit to Copenhagen became of greater importance to him than he could have imagined, for it marked the turning point in his life that changed not only his career but his philosophical outlook as well. The determining influence was Bohr's formulation of the complementarity concept as a generalized extension of Heisenberg's uncertainty principle. Thus, the propagation of light may be unambiguously defined, in a probabilistic way, either as a continuous motion of electromagnetic waves or as the exchange of individual quanta of energy related to the wavelength of the former by Planck's constant, but not by both at the same time; the two expressions of reality stand in a mutually exclusive but complementary relation to one another. According to Max, Bohr then very vigorously asked the question whether this new dialectic wouldn't be important also in other aspects of science. He talked about that a lot, especially in relation to biology, in discussing the relation between life on the one hand, and physics and chemistry on the other—whether there wasn't an experimental mutual exclusion, so that you could look at a living organism either as a living organism or as a jumble of molecules; . . . you could make observations that tell you where the molecules are, or you could make observations that tell you how the animal behaves, but there might well exist a mutually exclusive feature, analogous to the one found in atomic physics . . . in many respects Bohr wasn't sufficiently familiar with the status of the science (biology). So it was intriguing and annoying at the same time. It was sufficiently intriguing for me, though, to decide to look more deeply, specifically into the relation of atomic physics and biology—and that means learn some biology"(1).

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Biographical Memoirs: Volume 62 Much has been written of Bohr's profound influence on Max. Thus, Gunther Stent writes, "I think it is fair to say that with Max, Bohr found his most influential philosophical disciple outside the domain of physics, in that through Max, Bohr provided one of the intellectual fountainheads for the development of 20th century biology" (3). Again, Horace Judson said of Max, "His mind and style had been formed by Niels Bohr, the physicist, philosopher, poet and incessant Socratic questioner who made Copenhagen one of the capital cities of science between the wars" (4, p. 50). But Max himself saw more than this in the so-called Copenhagen Spirit, as shown by his reply to a question about the Phage Group: "Well, the phage group wasn't much of a group. I mean it was a group only in the sense that we all communicated with each other. And that the spirit was—open. This was copied straight from Copenhagen, and the circle around Bohr, so far as I was concerned. In that the first principle had to be openness. That you tell each other what you are doing and thinking. And that you don't care who—has the priority" (4, p. 61). It followed that, after a further 6 months with Lennard Jones at Bristol, Max decided to accept an appointment as assistant to Lise Meitner at the Kaiser Wilhelm Institute for Chemistry in Berlin in the autumn of 1932, because of its proximity to the Kaiser Wilhelm Institute for Biology. But before returning to Berlin he paid a short visit to Copenhagen to hear Bohr deliver his famous address, "Light and Life," to the opening meeting of the International Congress on Light Therapy in August, in which he explicitly stated his views on complementarity in biology (9). Odd though these views may seem to us now, in retrospect, this lecture confirmed Max's decision to turn to biology.

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Biographical Memoirs: Volume 62 THE BERLIN YEARS (1932-37) Max's appointment as assistant to Lise Meitner, who was collaborating with Otto Hahn on the results of irradiating uranium with neutrons, was, in effect, to be a consultant on theoretical physics. During this period he did write a few papers, one of which turned out to be an important contribution on the scattering of gamma rays by a Coulomb field due to polarization of the vacuum produced by that field (1933). His conclusion proved to be theoretically sound but inapplicable to the case in point, but 20 years later Hans Bethe confirmed the phenomenon and named it "Delbrück scattering." A second seminal paper with Gert Molière, which Max referred to retrospectively as "very learned" (1), attempted to apply quantum mechanics to resolve the paradox of irreversibility in statistical mechanics (1936c). Not long after the beginning of Max's Berlin period, which coincided with Hitler's rise to power, he organized a private group of five or six theoretical physicists to join in fairly regular discussions among themselves, often at his mother's house. At his suggestion some biochemists and biologists also joined the group. Among these were K. G. Zimmer whose interest was the dose effect of ionizing radiation on biological systems, and, most significantly for Max's future, N. W. Timoféeff-Ressovsky, a Russian geneticist from the Kaiser Wilhelm Institute for Brain Research who had been collaborating with Zimmer on the genetic effects of radiation for some 2 years before contact with Max was established. Timofeeff-Ressovsky's experimental organism was Drosophila, the fruit fly, which was then, and still is, very popular with geneticists because of its short generation time and the large populations that can be raised in the laboratory.

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Biographical Memoirs: Volume 62 thoughts on this theme were: "The journey of life which seems to be going outward, in the end turns out to have been going inward most of the time." On 5 March 1979 he wrote, "Im leichten Wellenschlag der Wochen treib ich dahin. Ein steuerloses Blatt bald zu verschwinden." ("I drift with the gentle undulation of the weeks. A rudderless leaf soon to disappear.") During the last few weeks of his life, Max announced one day that he had decided to live for two more years in order to complete his autobiography which he had recently started to write. Only 3 days before his death he began to dictate the chapter "Light and life" (B.C.). I WISH TO RECORD my most grateful thanks to Max's wife, Manny Delbrück, for her invaluable help in compiling this Memoir and commenting on the draft manuscript, and also to their daughter Nicola (N.D.) for her impressions of family life. Dr. Patricia Burke kindly provided me with a full bibliography, compiled with Manny's assistance, and Professor L. Hood provided an up-to-date list of Max's honours and other data. I am also indebted to many people who offered me impressions and reminiscences of Max. Personal and scientific recollections of his early career in theoretical physics were sent to me by Professor Sir Charles Frank, O.B.E., F.R.S. (C.F.) who also put me in touch with the University of Bristol, Dr. G. Herzberg, F.R.S., Professor N. Thompson (N.T.), and Professor V. F. Weisskopf (V.F.W.). Professor A. P. Eslava (A.P.E.) and Dr. E. D. Lipson provided assessments of the work of the Phycomyces Group, and Professor P. Starlinger (P.S.) enlightened me on the conception and birth of the Cologne Institute. Dr. P. M. Gresshoff, Professor G. S. Stent, and Professor M. J. D. White, F.R.S. kindly suggested appropriate amendments to the draft manuscript. Finally, I must also thank the following for permitting me to quote from their personal communications, contributions to Max's Memorial service, and other unpublished sources: Professor E. S. Anderson, F.R.S. (E.S.A.), Professor S. Benzer (S.B.), Fraulein Beate Carrière (B.C.) who recorded the last few weeks of Max's life (translated for me from the

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Biographical Memoirs: Volume 62 German by Dr. P. M. Gresshoff), Dr. D. R. Smith (D.R.S.), Professor G. S. Stent (G.S.S.), and Dr. J. D. Watson (J.D.W.). (The initials in brackets indicate the sources of quotations in the text.) FAMILY Married: 2 August 1941, Pasadena, California, to Mary Adeline Bruce (born 1917 in Butte, Montana, U.S.A.), daughter of James Latimer Bruce, mining engineer, and Leah Hills Bruce. Children: son, Jonathan, born 1947 in Nashville, Tennessee; daughter, Nicola, born 1949 in Pasadena, California; son, Tobias, born 1960 in Pasadena, California; daughter, Ludina, born 1962 in Cologne, W. Germany. HONORS Election to U.S. National Academy of Sciences—1949 American Academy of Arts and Sciences—1959 Royal Danish Academy—1960 Deutsche Akademie der Naturforscher Leopoldina—1963 Royal Society of London, Foreign Member—1967 Académie des Sciences, Paris, Associé Étranger—1979 Honorary Degrees Copenhagen University—1965: Doctor of Philosophy University of Chicago—1967: Doctor of Science Heidelberg University—1968 Harvard University—1971: Doctor of Science Gustavus Adolphus College, St. Peter, Minnesota, U.S.A.—1977: Doctor of Science University of Southern California—1981: Doctor of Science Göttingen University—1981: Doctor of Philosophy (to commemorate 50th anniversary of first degree) Awards Kimber Medal for Genetics (U.S. Academy of Science)—1964 Gregor Mendel Medal (Deutsche Akademie der Naturforscher-Leopoldina)—1967 Gross-Horwitz Prize (Columbia University)—1969 Nobel Prize for Physiology or Medicine—1969

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Biographical Memoirs: Volume 62 REFERENCES† (1) Oral History: Max Delbrück—how it was. Engng Sri. (California Institute of Technology) March-April, pp. 21-26; May-June, pp. 21-27 (1980). (2) In: Phage and the origins of molecular biology (eds. J. Cairns, G. S. Stent & J. D. Watson). (Cold Spring Harbor Laboratory of Quantitative Biology, Long Island, New York) (1968). (3) Tributes to Max Delbrück delivered at a Memorial Service in celebration of his memory, held at the California Institute of Technology on 19 April 1981; abstracted in Engng Sri. (California Institute of Technology) June, pp. 18-20. (1981). (4) Judson, H. F. The eighth day of creation. New York: Simon & Schuster. (1979). (5) Mullins, N. C. The development of a scientific speciality: the Phage Group and the origins of molecular biology. Minerva X, 1,51-82 (1972). (6) Stent, G. S. Obituary: Max Delbrück. Trends in biochemical sciences, 6(5), III-IV (1981). (7) The life of a Nobel wife. Engng Sci. (California Institute of Technology) March-April, pp. 14-24 (1977). (8) Benzer, S. Fine structure of a genetic region of bacteriophage. Proc. natl. Acad. Sci. U.S.A. 41, 344-354 (1955); see also, On the topography of the genetic fine structure, ibid. 47, 403-415 (1961). (9) Bohr, N. 1933 Light and life. Nature, Lond. 131, 421-423; 457459. (10) Bohr, N. 1965 Light and life revisited. In: Essays 1958-62 in atomic physics and human knowledge. New York: John Wiley. (11) Delbrück, M. 1978 The arrow of time—beginning and end. Engng Sci. (California Institute of Technology) 42, October. (12) Doermann, A. H. 1952 The intracellular growth of bacteriophages. I. The liberation of intracellular bacteriophage T4 by premature lysis with another phage or with cyanide. J. gen. Physiol. 35, 646-656. t   References 1-7 are to books and general articles quoted in discussion of Max's career and personality and are not in alphabetical order. The remaining references, 8-20 inclusive, are to scientific papers, listed alphabetically in the usual way.

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Biographical Memoirs: Volume 62 (13) Hershey, A. D. 1946 Spontaneous mutations in bacterial viruses, Cold Spring Harb. Symp. quant. Biol. 11, 67-77. (14) Hershey, A. D. & Chase, M. 1952 Independent functions of viral protein and nucleic acid in growth of bacteriophages. J. gen. Physiol. 36, 39-56. (15) Lipson, E. D. 1980 Sensory transduction in Phycomyces photoresponses. In: The blue light syndrome (ed. H. Senger). Berlin, Heidelberg, New York: Springer. (16) Schrödinger, E. 1944 What is life? (Cambridge University Press, Cambridge, England, New York, 1945: republished Doubleday, Inc., New York: Anchor A88, 1956). (17) Stent, G. S. 1968 That was the molecular biology that was. Science 160, 390-395. (18) Stent, G. S. & Calender, R. 1978 Molecular genetics: an introductory narrative, 2nd ed. San Francisco: W. H. Freeman & Co. (19) Watson, J. D. 1968 The double helix p. 217; facsimile of letter on 6 pp. following p. 226. London: Weidenfield & Nicolson. (20) Watson, J. D. & Crick, F. H. C. 1953 Genetic implications of the structure of deoxyribonucleic acid. Nature, Lond. 171, 964-967.

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Biographical Memoirs: Volume 62 BIBLIOGRAPHY 1929 Ergänzung zur Gruppentheorie der Terme. Z. Phys. 51, 181-187. 1930a Quantitatives zur Theorie der homöopolaren Bindung. Ann. Physik. 5, 36-58. b The interaction of inert gases. Proc. R. Soc. Lond. A 129, 686-698. 1931 (With G. GAMOW) Uebergangswahrscheinlichkeiten von angeregten Kerncn. Z. Phys. 72, 492-499. 1932 Possible existence of multiply charged particles of mass one. Nature Lond. 130, 626-627 (Erratum on p. 660). 1933 Zusatz bei der Korrektur; appendix to L. Meitner and H. Kösters, Ueberstreuung kurzwelliger g- strahlen. Z. Phys. 84, 137-144. 1935a (With L. MEITNER) Der Aufbau des Atomkern, natürlichc und künstliche Kernumwandlungen. (Book, 62 pages.) Berlin: J. Springer. b (With N. W. TIMOFÉEFF-RESSOVSKY and K. G. ZIMMER) Ueber die Natur der Genmutation und der Genstruktur. Nachr. Ges. Wiss. Göttingen 6 N.F. Nr. 13, 190-245. 1936a (With N. W. TIMOFÉEFF-RESSOVSKY) Strahlengenetische Versuche über sichtbare Mutationen und die Mutabilität, einzelner Gene bei Drosophila melanogaster. Z. indukt. Abstamm.—u. VerebLehre 71, 322-334. b (With N. W. TIMOFÉEFF-RESSOVSKY) Cosmic rays and the origin of species. Nature, Lond. 137, 358-359. c (With G. MOLIÈRE) Statistische Quanten-mechanik und Thermodynamik. Abh. preuss. Akad. Wiss. Nr. 1, 1-46. 1939 (With E. L. ELLIS) The growth of bacteriophage. J. gen. Physiol. 22, 365-384. 1940a Statistical fluctuations in autocatalytic reactions. J. chem. Phys. 8, 120-124. b Radiation and the hereditary mechanism. Am. Nat. 74, 350-362. c The growth of bacteriophage and lysis of the host. J. gen. Physiol. 23, 643-660. d Adsorption of bacteriophage under various physiological conditions of the host. J. gen. Physiol. 23, 631-642.

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Biographical Memoirs: Volume 62 e (With L. PAULING) The nature of the intermolecular forces operative in biological processes. Science 92, 77-79. 1941 A theory of autocatalytic synthesis of polypeptides and its application to the problem of chromosome reproduction. Cold Spring Harb. Symp. quant. Biol. 9, 122-124. 1942 (With S. E. LURIA) Interference between bacterial viruses. I. Interference between two bacterial viruses acting upon the same host and the mechanism of virus growth. Archs Biochem. 1, 111-141. a (With S. E. LURIA) Interference between bacterial viruses. II. Interference between inactivated bacterial virus and active virus of the same strain and of a different strain. Archs Biochem. 1, 207-218. b Bacterial viruses (bacteriophages). Adv. Enzymol 2, 1-32. 1943a (With S. E. LURIA) Mutations of bacteria from virus sensitivity to virus resistance. Genetics 28, 491-511. b (With S. E. LURIA & T. F. ANDERSON) Electron microscope studies of bacterial viruses. J. Bact. 46, 57-76. 1944a A statistical problem. J. Tenn. Acad. Sci. 19, 177-178. b (With S. E. LURIA) A comparison of the action of sulpha-drugs on the growth of a bacterial virus and its host. Proc. Indiana Acad. Sci. 53, 28-29 (Abstract). 1945a Spontaneous mutations of bacteria. Ann. Mo. bot. Gdn. 32, 223-233. b The burst size distribution in the growth of bacterial viruses. J. Bact. 50, 131-35. c Effects of specific antisera on the growth of bacterial viruses. J. Bact. 50, 137-150. d Interference between bacterial viruses. III. The mutual exclusion and the depressor effect. J. Bact. 50, 151-170. 1946a Bacterial viruses or bacteriophages. Biol. Rev. 21, 30-40. b Experiments with bacterial viruses (bacteriophages). Harvey Lect. Series 41, 161-187. c (With W. T. BAILEY, Jnr.) Induced mutation in bacterial viruses. Cold Spring Harb: Symp. quant. Biol. 11, 33-37. 1947 Über Bacteriophagen. Naturwissenschaften 34, 301-306. 1948a Biochemical mutants of bacterial viruses. J. Bact. 56, 1-10. b (With MARY BRUCE DELBRÜCK) Bacterial viruses and sex. Scient.

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Biographical Memoirs: Volume 62 Am. 179, 46-51; (Nov.); German translation in Naturwiss. Rundsch. 7, 301-306 (1949). 1949a Génétique du bactériophage. Colloques int. Cent. natr. Rerh. scient. 8, 91-103. b A physicist looks at biology. Trans. Conn. Acad. Arts Sci. 38, 173-190. 1951a (With J. J. WEIGLE) Mutual exclusion between an infecting phage and a carried phage. J. Bact. 62, 301-318. b (With R. K. CLAYTON) Purple bacteria. Scient. Am. 62, 68-72. 1953 (With N. VISCONTI) The mechanism of genetic recombination in phage. Genetics 38, 5-33. 1954a Wie vermehrt sich ein Bakteriophage? Angew. Chem. 66, 391-395. b On the replication of DNA. Proc. natl. Acad. Sci. U.S.A. 40, 783-788. 1956a Current views on the reproduction of bacteriophage. Scientia 91, 118-126. b (With W. REICHARDT) System analysis of the light growth reactions of Phycomyces. In: Cellular mechanisms in differentiation and growth (ed. D. Rudnick). Princeton University Press. 1957 (With G. S. STENT) On the mechanism of DNA replication. In The chemical basis of heredity (ea. W. D. McElroy & B. Glass). Baltimore: Johns Hopkins Press. 1958a Bacteriophage genetics. Proc. IV. int. Poliomyelitis Congr. New York: Lippincott. b (With S. W. GOLOMB & L. R. WELCH) Construction and properties of comma-free codes. Biol. Meddr. 23, 1-3. c (With R. COHEN) Distribution of stretch and twist along the growing zone of the sporangiophore of Phycomyces and the distribution of response to a periodic illumination program. J. Cell. comp. Physiol. 52, 361-388. 1959 (With R. COHEN) Photoreactions in Phycomyces, Growth and the tropic responses to the stimulation of narrow test areas. J. gen. Physiol. 42, 677-695. 1960 (With W. SHROPSHIRE, Jnr) Action and transmission spectra of Phycomyces. Pl. Physiol. 35, 194-204. 1961a (With D. VARJU) Photoreactions in Phycomyces. Responses

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Biographical Memoirs: Volume 62 to the stimulation of narrow test areas with ultraviolet light . J. gen. Physiol. 44, 1177-1188. b (With D. VARJU & Lois EDGAR) Interplay between the reactions to light and to gravity in Phycomyces. J. gen. Physiol. 45, 47-58. 1962a Knotting problems in biology. In: Mathematical problems in the biological sciences; Proc. of Symposia in Applied Mathematics 14, 55-68. b (With H. E. JOHNS & S. A. RAPAPORT) Photochemistry of thymine dimers. J. molec. Biol. 4, 104-114. c Ein Hinweis auf einige neue Gedanken in der Biologie. Phys. Bl. 18, 559-562. d Genetik und die Synthese ''lebender Substance". Der Mathematische und Naturwissenschaftliche Unterricht, Band 15, 241-244. 1963a Der Lichtsinn von Phycomyces. Ber. dt. bot. Ges. 75, 411-430. b (With L. GROSSMAN) The effects of monochromatic UV light on the structure and function of polyU. Fedn Proc. Fedn Am. Socs exp. Biol. 22, 538 (Abstract). c Inwiefern ist die Biologie zu schwierig für die Biologen? In: Physikertagung Stuttgart 1963. (Mosbach/Baden: Physik Verlag). d Das Begriffsschema der Molekular-genetik. Nova Acta Leopoldina NF 26, 9-16. e Über Verebungeschemie. Arbeitsgemeinschaflf. Forschung des Landes Nordhein-Westfalen 125, 1-39. f Die Verebungschemie. Naturwiss. Rundsch. 16, 85-89. 1965 Primary transduction mechanisms in sensory physiology and the search for suitable experimental systems. Isr. J. Med. Sci. 1:1363-1365. 1966 (With others) General discussion. Radiat. Res. Suppl 6, 227-234. 1967a Molecular aspects of genetics. In: Heritage from Mendel (ed. R. A. Brink) Madison, Wisconsin: University of Wisconsin Press. b (With K. L. ZANKEL & PATRICIA V. BURKE) Absorption and screening in Phycomyces. J. gen. Physiol. 50, 1893-1906. 1968a (With G. ADAM) Reduction of dimensionality in biological diffusion processes. In: Structural chemistry and molecular

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Biographical Memoirs: Volume 62 biology (ed. A. Rich & N. Davidson) San Francisco: W. H. Freeman & Co. b Biologie moléculaire: la prochaine étape. Science, Paris 56, 7-13. c (With G. MEISSNER) Carotenes and retinal in Phycomyces mutants. Pl. Physiol. 43, 1279-1283. 1969 (With K. BERGMAN, PATRICIA V. BURKE, E. CERDÁ-OLMEDO et al. Phycomyces. Bact. Rev. 33, 99-157. 1970a (With M. PETZUCH) Effects of cold periods on the stimulus response system of Phycomyces. J. gen. Physiol. 56, 297-308. b A physicist's renewed look at biology—twenty years later. In: Les Prix Nobel en 1969. (The Nobel Foundation, Stockholm); Science 168, 1312-1315. c Lipid bilayers as models of biological membranes. In: The neurosciences: second study program (ed. F. O. Schmitt). New York: Rockefeller University Press. 1971 Aristotle-totle-totle. In: Of Microbes and Life (André Lwoff Festschrift) (ed. J. Monod & E. Borek). New York: Columbia University Press. 1972a Homo scientificus according to Beckett. In Science, Scientists and Society (ed. W. Beranek, Jnr). Tarrytown-on-Hudson, New York: Bogden & Quigley Inc. German translation in Neue Sammlung 12, 528-542. b Signal transducers: Terra incognita of molecular biology. Angew. Chem. Int. Ed. Engl. 11, 1-7. c Out of this world. In Cosmology, Fusion and Other Matters (George Gamow Memorial Volume) (ed. F. Reines). Boulder, Colorado: Colorado Associated University Press. 1973a (With T. OOTAKI & ANITA C. LIGHTY. Complementation between mutants of Phycomyces deficient with respect to carotenogensis. Molec. gen. Genet. 121, 57-70. b Anfänge der Wahrnehmung. Karl-August-Forster Lectures 10 (1973) (Akad: Wissenschaften und der Literatur, Mainz). c (With W.-J. HSU & D. C. AILION) Carotenogensis in Phycomyces. Phytochemistry 13, 1463-1468. 1975a (With R. J. COHEN, Y. N. JAN, & J. MATRICON) Avoidance response, house response, and wind responses of the sporangiophore of Phycomyces. J. gen. Physiol. 66, 67-95. b (With A. P. ESLAVA, M. I. ALVAREZ, & PATRICIA V. BURKE)

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Biographical Memoirs: Volume 62 Genetic recombination in sexual crosses of Phycomyces. Genetics 80, 445-462. c (With P. G. SAFFMAN) Brownian motion in biological membranes. Proc. natl. Acad. Sci. U.S.A. 72, 3111-3113. d (With A. P. ESLAVA & M. I. ALVAREZ) Meiosis in Phycomyces. Proc. natn. Acad. Sci. U.S.A. 72, 4076-4080. 1976a (With A. KATZIR & D. PRESTI) Responses of Phycomyces indicating optical excitation of the lowest triplet state of riboflavin. Proc. natn. Acad. Sci. U.S.A. 73, 1969-1973. b Light and life III. Carlsberg Res. Commun. 41, 299-309. c How Aristotle discovered DNA. In: Physics and Our World; A Symposium in Honor of Victor F. Weisskopf. (Massachussetts Institute of Technology) 1974 (American Institute of Physics, New York). 1977a (With D. PRESTI & W.-J. HSU) Phototropism in Phycomyces mutants lacking ß-carotene. Photochem. Photobiol. 26, 403-405. b (With E. CERDÁ-OLMEDO) El comportamiento de Phycomyces. In: Genetica Microbiana. (Editorial Alhambra, Madrid, Spain). 1978a Erinnerung an Max Born. In: Max-Born-Gymnasium Germering: Jahresbericht 1977/78 (ed. H. Baumann). Max-Born-Gymnasium, Germering). b (With D. PRESTI) Photoreceptors of biosynthesis, energy storage, and vision. Pl. Cell Environ. 1, 81-100. c Virology revisted. In: Proceedings of International Symposium on Molecular Basis of Host Virus Interaction. Benares Hindu University, October 1976 (ed. M. Chakravorty). Princeton, New Jersey: Science Press. d Mind from Matter?? In: The Nature of Life XIII Nobel Conference (ed. N. H. Heidcamp). Baltimore, Maryland: University Park Press. (Shorter version in The American Scholar 47, 339-353.) 1979a (With T. OOKTAKI) An unstable gene in Phycomyces. Genetics 92, 27-48. b (With M. JAYARAM & D. PRESTI) Light-induced carotene synthesis in Phycomyces. Exp. Mycol. 3, 42-52.

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Biographical Memoirs: Volume 62 1980a Was Bose-Einstein statistics arrived at serendipity? J. chem. Educ. 57, 467-474. b (With M. JAYARAM & LESLIE LEUTWILER) Light-induced carotene synthesis in mutants of Phycomyces with abnormal phototropism. Photochem. Photobiol. 32, 241-245. 1981 (With M. K. OTTO, M. JAYARAM, & R. M. HAMILTON) Replacement of riboflavin by an analogue in the blue-light photoreceptor of Phycomyces. Proc. natn. Acad. Sci. U.S.A. 78, 266-269.