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ISRAEL MICHAEL LERNER May Id, 90-June I2, 1977 BY R. W. ALLARD I. MICHAEE EERNER MADE sophisticated contributions to popu- lation, quantitative, en c! evolutionary genetics, en c! ani- mal brawling. He excellec! in teaching at all levels, from providing nonscientists with realistic concepts of science and its importance in making policy decisions regarding the future of society, to teaching acivancec! courses in ge- netics. He also hac! exceptional talent for management en c! server! with distinction in many assignments cleating with intramural affairs at the University of California en c! with scientific policy at the national ant! international levels. Despite the preclominantly scientific cast of his professional career, Lerner's primary interests throughout his life were in the humanities. It is thus remarkable that he accom- plishec! so much in science. How this came to be Lerner explainer! eloquently in the brief autobiographical state- ment he submitter! to the National Academy of Sciences upon his election in 1959. The following quotation, with minor editing, is taken from that statement: I have been a scientist, not through any overwhelming curiosity about na- ture, not because of a drive to contribute to the welfare of humanity, nor because of the promise of any aesthetic satisfaction from experimentation and generalization. Indeed my inclinations have always been in the direc- tion of the humanities (I still regard myself as an historian manque), to 167

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68 B I O G RA P H I C A L EMOIRS ward such arts as the theater or toward politics or the law. I drifted into a scientific career by following a line of least, or at best, little resistance. I was lucky in my associates, I have been fortunate in the circumstances of my personal life, and the genes I inherited interacted favorably with the environments I found myself in. That is as much as introspection can yield regarding how I came to be me. The outward facts follow. There was nothing in my family's tradition or in my home environ- ment that would have predisposed me to an academic career in science. My father, at the time of my birth, was a successful importer and exporter living in Harbin, Manchuria (then a Chinese territory under a long-term Russian government lease). The life we led was reasonably typical of middle- class prosperous Russian families with some cultural pretensions, the the- ater, lectures, and concerts occupying a fairly prominent place in our daily existence. Certain departures from the Russian norm were occasioned by Harbin's geographical position (Chinese house servants, English regarded as a more important language than French). My sister (two years older than I) and I were first taken care of by two Russian nurses and then by German governesses. By the time the Russian revolution broke out in 1917 my sister and I were being tutored at home in the regular school subjects appropriate to our ages, with English and piano lessons on the side. The revolution had a tremendous impact on Harbin and on the personal circumstances of our family. The wave of emigres passing through Harbin, among whom a high proportion belong to the intelligensia, increased cultural activities for a short period far beyond the town's proportions in size or its provincial geographical position. Former University professors (true bearded Geiheimrats rather than the American variety) were so numerous that even many sec- ondary schools were able to obtain their services for teaching. Thus, when at age 12, my home education gave way to school attendance, much of my education was by specialists in University subjects, rather than by secondary school pedagogues thus I was exposed to political economy, philosophy, literary criticism and history at a much younger age than most of my con- temporaries from elsewhere. However, although letters, humanities and social sciences were taught by former University professors in the Harbin schools, this was not the case in the natural sciences. I suppose the reason was that there were fewer scientists in Russia to start with, that fewer scien tists migrated, and that an even smaller proportion took up teaching as an occupation. Another result of the revolution was that, during my child- hood, Harbin became a center of music and theater. We had a full nine

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ISRAEL MICHAEL LERNER 169 month opera season, a symphony, at least one dramatic theater operating throughout the year, a ballet troupe, a light opera troupe, and concerts by instrumentalists and singers. I acquired from these influences a deep and lasting interest and love for the performing arts, particularly opera. My father's finances suffered such severe reverses as a result of the revolution that, instead of several tutors, my sister and I were sent to schools, private because public schools were only at the primary level. Piano les- sons, and for a very short time drawing lessons (for which I exhibited absolutely no talent), were the only extra lessons continued. During this period my parents made attempts (unsuccessful) to emigrate to Switzer- land. In the fall of 1922, I was sent to the Harbin Public Commercial School, where I spent five years, graduating in the spring of 1927. The Russian Commercial School in Harbin was a compromise between a classi- cally oriented (Gymnasium) and a technically oriented (Realschule) sec- ondary school. The direction I was to take after school was not at all clear. It was understood that I should go on to University, but whether it would be to one of the institutions in Harbin, or whether I would follow my sister to Russia (where she became a physician), or emigrate to Europe, or to America, was unclear. Of the various prospects, going to America appealed most. I knew the language and I understood that working one's way through college there was much more common than in Europe. The prospect of doing military service in Russia, the difficulties for a scion of a bourgeois entering a Russian University at that time (my sister had difficulties), the uncertain- ties as to how my further education would be financed, were factors militat- ing against going to Russia. Harbin itself, even if I were successful in com- pleting a university course in some subject, provided only dismal vistas. So, America was the choice. But, by 1927, U.S. Immigration laws had tightened and a wait of many years for a visa was likely. However, a rumor spread through our school that Canada was an equally good place to go, provided that one announced intention to engage in agriculture. Suffice it to say that I left Harbin without passport, visa, or funds and found myself in September of 1927 in Vancouver, B.C. engaged in a farm job, digging ditches and caring for chickens at $2.00 per day on the Poultry Farm of the University of British Columbia. Thus I drifted by accident into a field which interested me only casually. A factor of major importance in staying in this field was the encouragement of Vigfus F. Asmundson, then an Assistant Professor in the Department, who was engaged in Poultry Genetics re- search. I soon became his assistant and continued to work with him until I

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170 B I O G RA P H I C A L EMOIRS obtained my B.S. and M.S. degrees. He lent me money to pay tuition and often, when the Department budget was strained, paid me out of his own pocket for the work I was doing. It was to him that I owed my determina- tion not only to enter into an academic career, but to do so specifically in the field of Genetics (I had no inkling that he would move to Davis and I would one day become his fellow staff member in the Poultry Department at the University of California). In 1931 Theodosius Dobzhansky spent a month in Vancouver and I had nearly daily contact with him. Dobzhansky's enthusiasm for research in genetics provided very strong reinforcement for my wishes to continue graduate work but it was not until 1933 that an offer of an assistantship that I could afford to accept presented itself. It was in the Poultry Department at Berkeley with L. W. Taylor, a fact that commit- ted me to work with the chicken for the next 25 years. When Lerner receiver! his Ph.D. in genetics at Berkeley in 1936, he was appointed instructor in poultry husbandry, from which level he receiver! accelerates! promotions to pro- fessor. Thus, revolution, financial problems, periods of go- ing hungry, en c! other clire clifficulties, intersperses! with some comic relief episodes, together with much heart-warm- ing help en c! encouragement, launcher! what wouic! prove to be a remarkable career but one that took a very cliffer- ent direction from the course that might have been pre- clictec! from his early knowlecige en c! creep attraction to the humanities. The researches concluctec! by Lerner in his twenty-five years in the Department of Poultry Husbandry at Berkeley (many in collaboration with Everett R. Dempster of the Department of Genetics and Dorothy C. Lowry, his techni- cal assistant) were reported in more than 175 published papers. As a young faculty member he dealt with the inher- itance of a number of components underlying egg produc- tion, the effects of practicing selection in conjunction with inbreeding, en c! with empirical tests of theoretically pre- clictec! gains from simultaneous selection for several cliffer- ent inherited! characteristics. These studies lee! to construc

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ISRAEL MICHAEL LERNER 171 lion of selection inclices, creclitec! by commercial poultry producers as responsible for substantial increases in egg production. Two of Lerner's books (Population Genetics and Animal Improvement, 1950, en c! Genetic Basis of Selection, 1958) were highly influential in transforming animal brawling from an art to a science baser! on multifactorial Menclelian in- heritance. In another book (Genetic Homeostasis, 1954) Lerner formulates! a brilliant hypothesis relating natural selection en c! evolution that stimulates! much thought, discussion, en c! controversy (in the words of one generally unfriencITy critic, it was speculative, imaginative, controversial, en c! in- fluential) . During the late 1950s, Lerner's interests turner! increas- ingly to the ways that studies of domestic en c! laboratory animals might throw light on the genetic basis of selection en c! evolution. In 1958 he joiner! the Department of Genet- ics at Berkeley, adopting the common flour beetle as an experimental organism more suitable for his new purposes and carried out many exquisitely designed competition ex- periments. He shower! that the outcomes of his experiments were almost entirely deterministic when the experimental conditions as well as the genetic compositions of the com- peting entities were carefully controllecI. He also shower! that some of the characteristics involves! in competitive ability were behavioral. This lee! him to in-clepth studies of the technical literature in psychology, and he was invited to join, on a part-time basis, the Institute of Personality Assess- ment on the Berkeley campus. During the late 1960s en c! until his cleath in 1977, Lerner's research activities were increasingly replaced by administra- tive en c! eclitorial work en c! the summarizing of various as- pects of evolutionary genetics in numerous invites! aciciresses en c! articles. He server! as chairman of his department en c! the graduate council at Berkeley en c! on various boards of

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172 B I O G RA P H I C A L EMOIRS the statewide University of California system en c! the U.S. Department of Agriculture. He also servec! as editor of the journal Evolution en c! as secretary of the International Com- mission on Genetic Congresses. Lerner consiclerec! teaching to be of primary importance. Among his major contributions to teaching were a book titles! Heredity, Evolution and Society en c! an associates! course clesignec! to provide nonscientists with unclerstancling of the role of science in formulating sounc! public policy. Both the course en c! the book were highly popular at Berkeley, en c! both have been wiclely imitated. Lerner receiver! many honors en c! recognitions. Those he values! most, in aciclition to membership in the National Academy of Sciences, were election to the American AcacI- emy of Arts en c! Sciences, to the vice-presiclency of the Ameri- can Society of Naturalists, as a foreign member of the Florentine Academia clef Georgofili, en c! as editor of Evolu- tion. He also values! receiving the Borden Awarc! en c! GoIc! Mecial, the Belling Prize in Genetics, the Poultry Science Research Award, en c! honorary degrees conferred by the University of British Columbia en c! the University of Edinburgh. Throughout his life Lerner followed a demanding ethi- cal imperative. He was meticulously honest en c! straightfor . ... ware! in expressing his opinions while at the same time man- a~in~ to avoic! offense. During the last years of his life his health was poor. His death on June 12, 1977, at age sixty- seven, follower! a series of major abclominal operations as well as operations for cataracts en c! a cletachec! retina, with complications from emphysema. His courage cluring these tribulations was remarkable. Lerner greatly enjoyed many aspects of life en c! conveyor! his pleasure to others. He is greatly missed by his many friends throughout the world. It is appropriate to close this memoir with an appreciation of

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ISRAEL MICHAEL LERNER 173 Ruth Stewart! Lerner, his classmate at the University of Brit- ish Columbia. She proviclec! him with acivice, encourage- ment, en c! support, all greatly appreciated, throughout the forty years of their marriage. I first met Michael Lerner in Berkeley, probably in 1939 or 1940, when I was an unclergracluate student on the Davis campus of the University of California. Starting in 1946, when I joiner! the faculty at Davis, many opportunities arose to talk to Lerner at Davis, Berkeley, en c! at scientific meet- ings at various places in North America en c! Europe. I acI- mirec! his breadth of knowlecige in biology en c! the hu- manities en c! treasurec! his friendship en c! counsel over the more than three clecacles I was privilegec! to continue my association with him.

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174 B I O G RA P H I C A L S E L E C T E D EMOIRS B I B L I O G RAP H Y 1932 With V. S. Asmundson. Inheritance of growth rate in the domestic fowl. Sci. Agric. 2:652-64. 1933 With J. V. Bierly and V. E. Palmer. Fowl paralysis (Neurolymphomatosis gallinarum) in chicks under three months of age. Can. f. Res. 8:30. 1936 Heterogony in the axial skeleton of the creeper fowl. Am. Nat. 70:595- 98. 1937 With L. W. Taylor. The spurious nature of linkage between the length of laying year and sexual maturity in the fowl. Am. Nat. 71 :617-22. 1939 The shape of the chick embryo growth curve. Science 89:16-17. Allometric studies in poultry. In Proc. 7th World Poultry Congress. Cleveland, pp. 85-88. 1940 With L. W. Taylor. The effect of controlled culling on the efficiency of progeny tests. 7. Agric. Res. 61: 755-64. With J. Needham. The terminology of relative growth rates. Nature 146:618. 1941 With J. S. Huxley and J. Needham. Terminology of relative growth rates. Nature 148:225. 1943 The failure of selection to modify shank-growth ratios of the do- mestic fowl. Genetics 28:119-32.

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ISRAEL MICHAEL LERNER 1944 175 Lethal and sublethal characters in farm animals. 7. Hered. 35:219-24. 1947 With L. N. Hazel. Population genetics of a poultry flock under se- lection. Genetics 32:325-39. With E. R. Dempster. The optimum structure of breeding flocks. 1. Rate of genetic improvement under different plans. Genetics 32:555- 66. With E. R. Dempster. Heritability of threshold characters. Genetics 35:212-34. 1948 With E. R. Dempster. Some aspects of evolutionary theory in the light of recent work in animal breeding. Evolution 2:19-28. With D. Lowry. The heritability of accumulative monthly and an- nual egg production. Poult. Sci. 27:67-78. 1949 With A. Robertson. The heritability of all-or-none traits: viability in poultry. Genetics 34: 395-411. 1950 Population Genetics and Animal Improvement. Cambridge: Cambridge University Press. Genetics in the U.S.S.R.: An Obituary. University of British Columbia Publ. Lecture Series 8. 1954 Genetic Homeostasis. Edinburgh: Oliver and Boyd. 1955 Concluding survey. Cold Spring Harbor Symp. Quant. Biol. 20:334-40. 1958 Genetic Basis of Selection. New York: J ohn Wiley. The concept of natural selection: a centennial view. Proc. Am. Philos. Soc. 103:173-82. 1960 Marxist biology viewed dimly. Am. Nat. 91:45-55.