LERNER, I(SADORE) MICHAEL

(b. Harbin, Manchuria, 14 May, 1910; d. Berkeley, California, 12 June 1977),

genetics, evolutionary theory, poultry husbandry.

I. Michael Lerner is best known for formulating the influential concept of genetic homeostasis, usually defined as the tendency of a Mendelian population to maintain a constant genetic composition in the face of external pressure. The concept was formally articulated in 1954 in a widely read book titled Genetic Homeostasis. Lerner’s interests in evolutionary and genetical theory largely grew out of his interests in the practical breeding of poultry. He later turned to theoretical genetics and behavior genetics and explored the social implications of biology. Over the course of his career, he made notable contributions to both practical and theoretical aspects of breeding, genetics, evolution, and understanding the relationship between biology and society.

Early Life and Education . Lerner was born in Harbin, Manchuria in 1910, of Russian Jewish parents. Manchuria at that time was a Chinese territory under Russian government lease. Lerner’s father was a businessman who ran a successful import export firm. The family was fairly prosperous and supported artistic and cultural activities popular at the time. Lerner, therefore, grew up in an environment that encouraged the love of theater, music, and the performing arts in general. He was the second of two children (his sister was two years older), who were tended to first by Russian nurses and then by German governesses. His facility with the English language owed much to this early Manchurian influence, which stressed English rather than French as a second language to Russian.

Largely tutored at home, the Lerner children began to attend formal school only after the Russian Revolution in 1917. This event had a dramatic effect on life in Harbin and on the Lerner family especially, as they suffered enormous financial reversals. Lerner and his sister were sent to private schools because the family could no longer afford tutors at home. In the fall of 1922, Lerner was sent to the Harbin Public Commercial School, which was a cross between a classically oriented gymnasium and a technically oriented Realschule. For Lerner, one of the few positive outcomes of the revolution was that many of his teachers were university-level instructors who were Russian émigrés. This was especially true of his teachers in the humanities and social sciences, who taught secondary school courses with the sophistication usually seen at the university level: Lerner took courses in philosophy, literary criticism, and history at a much earlier age and taught by individuals with advanced specialties in the area.

The other positive outcome of the revolution was that Harbin itself became a kind of refuge for Russian émigré artists as well as intellectuals who formed a lively and bustling community rich in the arts, including opera, symphony, ballet, and theater companies. Lerner’s lifelong interest in the arts and humanities owes much to this early family influence and to his educational experience in Harbin. He showed no marked interest in the sciences initially, and even well after he had earned himself a formidable reputation in genetics he continued to identify with the humanities, even going to far as to call himself a kind of “historian manqué” in one interview (Hall, 2005).

At the time of his graduation in 1927, Lerner was unsure of the direction he would take. His family had tried to emigrate to Switzerland in 1923, but had failed; Lerner leaned toward using his education as the opportunity to leave Harbin. His sister had earlier left for Russia to become a physician, but Lerner dreaded the required military service in Russia. He also feared what the new regime would make of his “bourgeois” family background, the expense of a Russian education, and the political uncertainty that clouded any future direction he might take there. Because of his comfort with the English language, he considered immigrating to the United States, but because immigration restriction laws were tightened by 1927, he decided on Canada because it was accepting immigrants, especially if they showed an interest in agricultural sciences.

Lerner’s entry into what would develop into a brilliant scientific career thus began as a result of an opportunistic decision to leave Harbin for Canada. He later described it as “the path of least resistance.” By September of 1927, Lerner landed in Vancouver, British Columbia, a preferred port of entry for people on the Pacific rim, without funds or a passport, but with the job of digging ditches and caring for the chickens on the poultry farm of the University of British Columbia. He was paid the sum of two dollars per day for this work. He eventually served as assistant to Vigfus F. Asmundson, an assistant professor in the poultry genetics department who encouraged Lerner to continue in the area by obtaining advanced degrees. Asmundson even went so far as to help pay for his education by lending him funds when needed. Lerner thus obtained his B.S. and M.S. degrees at the University of British Columbia with a specialty in poultry genetics. He also met and married a fellow classmate, Ruth Stuart, with whom he shared a number of interests.

Theodosius Dobzhansky, also a Russian émigré, would also play a critical role in Lerner’s life. They met in 1931 when Dobzhansky, who had been then working as a fellow in the laboratory of American geneticist Thomas Hunt Morgan, visited the University of British Columbia and found himself stranded there awaiting a return visa to the United States. Because both Dobzhansky and Lerner spoke Russian and because they had similar interests in genetics, the two bonded in what would become a fruitful, lifelong scientific and personal relationship. Dobzhansky’s famous charismatic influence played itself out on Lerner as his interest in genetics and its applications in evolution began to grow in the 1930s.

In 1933, Lerner received an offer of an assistantship to continue graduate work at the University of California, Berkeley, in the poultry husbandry department working under the direction of L. W. Taylor. The timing was propitious, because the University of British Columbia was shutting down its own poultry program. He received his PhD in 1936, and was appointed instructor in poultry husbandry in Berkeley. He received a series of accelerated promotions to professor. In 1958, he moved to the Department of Genetics as chair. Lerner became emeritus professor in 1973 and remained associated with the Berkeley campus until the end of his life.

Scientific Work . Lerner’s scientific work is generally divided into three periods. The first grew out his involvement with practical breeding, primarily with poultry. His primary interests were in improving poultry stocks and egg production: He first examined poultry growth and disease resistance, then moved on to egg production and the effects of artificial selection and inbreeding in chickens. Lerner was especially interested in the effects of selection combined with inbreeding, and devised a number of tests for the theoretical projections of gains from the selection of several different characteristics simultaneously. He formulated what is known as the “optimum selection index,” which was a way of optimizing the effects of selection that could lead to rapid increases in egg production. This latter concept was applied widely and led to the doubling of egg production in commercial flocks of domestic chicken.

For much of this research, Lerner collaborated with Berkeley colleague Everett R. Dempster, a mathematically inclined population geneticist, and Dorothy C. Lowry, his technical assistant. It was an especially productive phase of Lerner’s career, culminating with the publication of his first book, Population Genetics and Animal Improvement, in 1950. This book established Lerner as one of the leaders and original thinkers in the field of animal breeding as a whole. His reputation and influence in animal breeding were enhanced further in 1958 when he published Genetic Basis of Selection. Both books helped to take what was long thought to be folk knowledge into the realm of a quantifiable and testable science. Some of his work examining poultry growth also took him to collaborative ventures with pioneers of allometry (the science of differential growth) such as Joseph Needham and Julian Huxley.

Lerner’s second research period grew out of his direct knowledge of animal breeding and the interplay of selection, inbreeding, and their effects on the genetic composition of populations. It was also aided and abetted by a growing circle of acquaintances in the San Francisco Bay Area with an interest in exploring genetics and evolution and the integration of the two as it was emerging during the period of the evolutionary synthesis. Beginning as a graduate student, Lerner had organized a group of active younger researchers on the Berkeley campus with a keen interest in integrating knowledge of the new genetics in both practical and theoretical terms within a dynamic evolutionary framework. Berkeley genetics itself was booming at the time, with workers such as Ernest Brown Babcock, the plant geneticist who focused on the genus Crepis, and Roy Elwood Clausen and Thomas Harper Goodspeed, who were working on the genetics of Nicotiana tabacum. With Dempster, Lerner organized a fortnightly journal club, Genetics Associated. By the late 1930s the group included Donald Cameron (then a research assistant to Clausen) and James Jenkins and G. Ledyard Stebbins (junior geneticists on the Crepis project with Babcock), along with plant breeders such as Francis Smith and Alvin Clark. Lerner also interacted with German émigré physiological geneticist Richard Goldschmidt when he later joined the faculty at Berkeley, though he disagreed with Goldschmidt’s notion of sudden or abrupt evolution through macromutations. Located nearby at the California Institute of Technology, Lerner’s old friend Dobzhansky was also a frequent visitor to the Bay Area and infused colleagues with his enthusiasm for evolutionary genetics as it was emerging from his researches on varied species of Drosophila. The Berkeley campus was thus a hub for research into evolutionary genetics in the late 1930s and 1940s; Lerner’s interactions with his colleagues outside his primary department of poultry husbandry and applied genetics inspired him to think in broader and more theoretical terms.

Lerner’s increasing confidence with theoretical genetics combined with his empirical knowledge in practical animal breeding came to fruition during his second period of research into theoretical genetics. In 1954 he published the book setting forth the conceptual theory of genetic homeostasis for which he is best known. Written in 1953 over a period of six months, while on a Guggenheim Fellowship at the Institute of Genetics in Pavia and at the Istituto Italiano di Idrobiologia in Pallanza, Italy, the book was dedicated to his long-time collaborator Dempster, who helped guide him with mathematical modeling. A small book of only 134 pages, Genetic Homeostasis packed a punch and was widely discussed in the biological sciences, and not without some controversy.

The novel concept set forth in the book and captured in the title was inspired by the concept of physiological homeostasis as it had been set forth by Walter Cannon in his celebrated The Wisdom of the Body in 1932. It referred to organisms regulating their internal physiological mechanisms as a way of buffering themselves or resisting external pressures. By genetic homeostasis, Lerner had in mind the same kind of regulatory or buffering capacity in populations of organisms that enabled them to preserve optimal fitness despite external pressures. Lerner defined it in his book as the property or ability of a population of organisms “to equilibrate its genetic composition and to resist sudden changes” (p. 2). The concept bore some resemblance to the concept of “genetic inertia” formulated by C. D. Darlington and Kenneth Mather in 1949, but Lerner strongly opposed the notion that they were identical or that he should cede priority to them.

As he pointed out repeatedly, homeostasis referred to self-regulatory capacity or the ability of organisms to buffer themselves against external pressures in a way that inertia, borrowed from physics, did not. It was a component he viewed as critical to his theory. As he extended the genetic conceptualization further, Lerner stressed that the most adapted types within a population were not those that conformed to phenotypic norms, but those that would have the highest fitness by their ability to adapt to changing environments. For Lerner, heterozygosity conferred just such an advantage: Heterozygotes generated two kinds of gene products and were therefore better buffered than homozygotes against environmental changes. The argument for genetic homeostasis thus hinged on Lerner’s argument for heterozygote superiority, the mechanics of which he took great pains to explore in his book. In espousing heterozygote superiority, Lerner was echoing colleagues such as Dobzhansky, but he also drew notable critics, who pointed out that attempts to find the mechanisms for gene action had proven unsuccessful. The data simply could not support it, while other genetic phenomena such as pleiotropy (interactive gene effects) and linkage disequilibrium offered more plausible explanations.

Other elements of Lerner’s conceptual theory were well received, especially by biologists stressing integrative approaches to evolution, genetics, and development. Fundamentally, Lerner’s conceptualization of genetic homeostasis depended on developmental homeostasis, because he began with the assumption that all sexual organisms had genotypes that produced self-regulating developmental patterns and phenotypes. The relationship was explicitly stated. “Error,” he wrote in his concluding paragraph, “is minimized in successful populations by developmental homeostasis, genetic homeostasis arises as an after-effect.” The relationship between the two was so close that to some the terms homeostasis, assimilation, buffering, inertia, and canalization (terms usually associated with development) are used interchangeably. But what made Lerner’s conceptualization new to developmental biologists at the time was its linked dependency on the heterozygous condition; as Lerner himself stated, heterozygotes were better able to “stay within the norms of canalized development” (1954).

Lerner’s integration of population genetics with developmental biology was instantly appreciated by Conrad Hal Waddington, the noted developmental biologist, who had been famously frustrated by the absence of embryology or developmental biology in the synthetic theory of evolution. Population geneticists such as Dobzhansky, whose own preference was for heterozygote superiority, regarded the thesis as brilliant. Indeed, at a time when workers such as Ernst Mayr charged mathematical population genetics with being simplistic, reductionist, and “bean-bag,” Lerner offered an ambitious and satisfying alternative that integrated genetics, evolution, and developmental biology in an entirely novel way. Though critics have charged that empirical data have not supported the argument for heterozygote superiority, Lerner is still regarded as a pioneer who made a “valiant” effort in seeking to integrate “genes, organisms, and development” (Hall, 2005). In this respect, Lerner may join workers such as Waddington in being one of the early pioneers of the new field known as “evo-devo,” seeking to integrate evolutionary biology with developmental biology. Lerner is also recognized for being one of the first to appreciate the importance of the co-adapted gene pool or co-adapted gene complexes (West-Eberhard, 2003). Whatever the durability of all or parts of the concept of genetic homeostasis, the 1954 book was in its day important and influential, read by a wide circle of biologists. It was deemed an “acknowledged masterpiece” by many (Glass, “I. Michael Lerner Papers”), and deemed so important that even his critics described it as “speculative, imaginative, controversial, and influential” (Allard, 1996, p. 171).

Lerner’s final period of research began with a shift to behavior genetics and led to his entry into the field of behavioral psychology. He adopted as his model organism the common flour beetle, Tribolium spp., and designed a series of elegant and elaborate experiments to understand the interplay of genetics and behavior in the competitive behavior of this organism. This work eventually led to his appointment to the Institute of Personality Assessment and Research on the Berkeley campus, a part-time appointment that enabled him to work with graduate students in psychology. His general turn toward the interplay of genetics, evolution, and social behavior became increasingly the centerpiece of his work in the 1960s. It was also the opportunity for Lerner to integrate his lifelong passions for the humanities and social sciences. A committed teacher, Lerner designed an enormously popular course on the Berkeley campus to explore the interplay of genetics, evolution, and society, for which he developed the textbook Heredity, Evolution, and Society in 1968. The book explored contentious topics that included the extension of evolutionary and genetic theory to human affairs such as intelligence, race, and the role of selection in human social behavior broadly construed. Lerner skirted strict genetic determinism, but nonetheless believed that genetics played a powerful role in governing human affairs. The course also explored issues of world hunger, conservation, and population control, all major topics of concern to the social implications of biology. With a liberal, and indeed a markedly leftist political orientation, Lerner managed to teach this course all through the period of campus unrest and the radical politics at Berkeley through the 1960s, drawing huge crowds of students to it and becoming one of the most celebrated teachers at Berkeley. His political commitments were also apparent in his fight with Lysenkoism (an anti-genetics political campaign) in the Soviet Union; he was an outspoken critic of this regime and its effect on population genetics, writing works with provocative titles such as Genetics in the U.S.S.R.: An Obituary, and “Marxist Biology Viewed Dimly.” One of his major contributions to disseminating knowledge of the oppressive effects of Stalinist control of science was his translation and editing of Zhores A. Medvedev’s The Rise and Fall of T. D. Lysenko in 1969.

Even though Lerner had as technical and exacting a scientific career as one could imagine, he managed to interweave his extensive love of the humanities, including history, literature, and philosophy with his science throughout much of his career. He was renowned as a skilled and indeed an erudite writer, and a master of the synthetic and original thought-piece. He also brought some of his insights into the human condition not only to his teaching, but also to his administrative work, which began to occupy his time beginning in the late 1960s and 1970s. Lerner had an amiable personality and was much liked and respected at Berkeley. He held a number of prominent positions there, along with roles on committees and boards in the University of California system. He was also an active organizer of the International Congresses of Genetics and was elected to prominent offices, including membership in the National Academy of Sciences in 1959. In the last decade of his life, he was plagued with poor health that included cataracts, a detached retina, and emphysema, and he required a number of abdominal operations. He worked through many of these illnesses but finally succumbed in 1977.

BIBLIOGRAPHY

WORKS BY LERNER

Genetics in the U.S.S.R.: An Obituary. Vancouver: University of British Columbia, 1950.

Population Genetics and Animal Improvement as Illustrated by the Inheritance of Egg Production. Cambridge, U.K.: Cambridge University Press, 1950.

Genetic Homeostasis. New York: Wiley, 1954. The Genetic Basis of Selection. New York: Wiley, 1958.

“Marxist Biology Viewed Dimly.” American Naturalist 93

(1960): 45–55.

Heredity, Evolution, and Society. San Francisco: Freeman, 1968.

As editor and translator. Zhores A. Medvedev. The Rise and Fall of T. D. Lysenko. New York: Columbia University Press, 1969.

OTHER SOURCES

Allard, Robert W. “Israel Michael Lerner.” Biographical Memoirs National Academy of Sciences 69 (1996): 167–173. Name is erroneously presented. Includes excerpts from an oral history memoir with a description of Lerner’s early life.

Cannon, Walter B. The Wisdom of the Body. New York: Norton, 1932.

Darlington, Cyril D., and Kenneth Mather. The Elements of Genetics. London: Allen and Unwin, 1949.

Dempster, Everett R., M. M. Green, and S. Washburn. “I Michael Lerner, 1910–1977.” Genetics 88 (1978): 139–140.

Futuyma, Douglas J. Evolutionary Biology, 3rd ed. Sunderland, MA: Sinauer Associates, 1998.

Glass, Bentley. “Israel Michael Lerner.” Yearbook of the American Philosophical Society (1984–1985): 130–135. Name is erroneously presented.

———. “I. Michael Lerner Papers.” Guide to the Genetics Collections at the American Philosophical Society Library.

Hall, Brian K. “Fifty Years Later: I. Michael Lerner’s Genetic Homeostasis (1954): A Valiant Attempt to Integrate Genes, Organisms and Environment.” Journal of Experimental Zoology 304B (2005): 187–197. Includes a detailed explanation of genetic homeostasis and offers a historical assessment of its applicability as well as its scientific rigor.

Mayr, Ernst, and William B. Provine, eds. The Evolutionary

Synthesis: Perspectives on the Unification of Biology. Cambridge, MA: Harvard University Press, 1980.

Patrikeeff, Felix. Politics in Exile: Manchuria and the Balance of Power in Northeastern Asia, 1924–1931. New York: Palgrave Press, 2002.

Siegel, Paul B. “I. Michael Lerner (1910–1977): Specialist and Generalist.” Behavior Genetics 8, no. 3 (1978): 225–226.

Smocovitis, Vassiliki Betty. “Keeping Up with Dobzhansky: G. Ledyard Stebbins, Plant Evolution, and the Evolutionary Synthesis.” History and Philosophy of the Life Sciences 8, no. 8 (2006): 9–48.

West-Eberhard, Mary Jane. Developmental Plasticity and Evolution. Oxford, U.K.: Oxford University Press, 2003.

Wolff, David. To the Harbin Station: The Liberal Alternative in Russian Manchuria, 1898–1914. Stanford, CA: Stanford University Press, 1999.

Woolf, Charles M., and Therese A. Markow. “Genetic Models for Developmental Homeostasis: Historical Perspective.” Tucson: University of Arizona, 1994. Available from http://eebweb.arizona.edu.

Vassiliki Betty Smocovitis