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The Birth of the Genetic Theory of Evolution in the Soviet Union in the 1920s

Theodosius Dobzhansky

When Darwin's On the Origin of Species was published in 1859, Russia was entering a period of political reforms and a ground swell of radicalism among its intelligentsia. The coincidence was fortuitous, but it left an impress on the intellectual tradition. Evolution was accepted not only as a scientific theory but also as a part of the liberal world view. Chernyshevsky and Pisarev, standard bearers of the radical youth, proclaimed that a valid personal philosophy must rest on a solid base of natural science, and evolution was a pivotal part of that. Professor Kutorga had lectured on Darwin's theory at Moscow University in 1860 and published an account of it in 1861. A Russian translation of Darwin's classic appeared in 1864. In 1865 Timiriazev, who was as effective an exponent of Darwinism in Russia as Huxley in England or Haeckel in Germany, published a collection of essays on evolution, which went through seven editions by his death in 1920.

Politically conservative circles took a dim view of Darwin's work because of its alleged political implications rather than evolution as a biological theory. As Danilevsky, one of Darwin's critics, indicates, the issue was taken extremely seriously: "It is clear how vitally important not only for zoologists and botanists, but for any moderately intelligent person is the issue whether Darwin is or is not right. It is so important that I am firmly convinced that no equally important problem exists either in any other field of knowledge or in any realm of practical life. Indeed, this is the problem of 'to be or not to be' in the strictest and broadest sense."

The Orthodox church was never very active in its opposition to evolutionism, and nothing like laws prohibiting the teaching of evolution or a Tennessee "monkey trial" occurred in Russia. The polemics of the debate were published neither in scientific nor in religious periodicals, but mostly in general literary and sociopolitical journals intended for broad circles of educated readers. This public did not usually have much understanding of biology as a science, but its bearings on human problems were considered important. When I was in my middle teens, a conservative lady of our acquaintance asked me: "Do you really believe in this horrid theory of Darwin?" I could have replied that those who do not credit this horrid theory seem to me rather dull-witted. Critics and Defenders of Darwinism Among those who accepted evolution as a part of the new gospel, some had reservations about certain parts of Darwin's theory. The struggle for existence seemed to have particularly undesirable connotations; already Chernyshevsky held the Lamarckian "transformism" superior to Darwin's natural selection. Kropotkin's attempt to give primacy as an evolutionary force to mutual aid, instead of competition and struggle, is sufficiently well known, because his essays were published in an English periodical (1902), while Kessler's "The Law of Mutual Aid" (1880), published in Russian, was virtually ignored. The checkered career of neoLamarckism in Russia has recently been well analyzed by Gaisinovich (1968) and Bliakher (1971). The polemics about Lamarckism versus Darwinism and genetics became a caricature of scientific discussion when the problem was taken over by Marxist philosophers in the 1920s and early 1930s. The Timiriazev Institute was working on "the study and propaganda of the scientific foundations of dialectical materialism." The Communist Academy had a Section of Natural and Exact Sciences. The Faculty of Medicine of Moscow University had a Society of Materialist Physicians. The criterion of validity of theories of evolution was their congruity with dialectical materialism as construed by different authorities. I remember the frustration I felt discussing some problem of genetics or evolution with Serebrovsky, an excellent geneticist and a convinced Marxist, in 1926 or 1927. His clinching argument to me was, "Your reasoning is undialectical." The debates among the high priests of dialectics were often impassioned but inconclusive. Both Lamarckians and Darwinians claimed to be faithful dialecticians. These polemics had however a wholly unintended effect: they prepared the ground for Lysenko's simplistic brand of dialectics, which for almost a generation swept away much of biology in Russia.

Most evolutionists in Russia were, of course, busy doing biological research rather than philosophizing. All approaches to evolutionary problems explored in other countries had their proponents in Russia. One tendency was particularly widespread and characteristic- a preoccupation with the organic diversity and the congruity of organic structures with the ways of living in different environments. Russia has a variety of climates from arctic to subtropical, vast plains and high mountains, forests and steppes, great rivers, lakes, and seas. Exclusively laboratory workers who neither possess nor wish to have any knowledge of living beings in nature were and still are a minority. Zoological and botanical systematics, comparative anatomy, phylogenetic studies on fossil and living forms, and later genetics were pursued in universities, as well as in museums and laboratories under the aegis of the Academy of Sciences. It was this tradition that yielded such evolutionists as Kovalevsky, Pavlov, and Mechnikov in the 1800s and Berg, Chetverikov, Karpechenko, Levitsky, Philipchenko, Schmalhausen, Severtzov, Vavilov, and many others during the first third of the 1900s. Some of the numerous amateur naturalists became very competent scientists. Genetics started tardily, but developed with great elan once it did start. The first course of genetics was given in the University of St. Petersburg by Philipchenko in 1913. I never had a course of genetics at the University of Kiev, although my teacher, Kushakevich, was an excellent cytologist and adherent of the chromosome theory of heredity. At that time I was an entomologist specializing in the taxonomy of Coccinellidae (lady beetles). Mendel's laws were occasionally mentioned, though not in evolutionary context.

The theory of natural selection reached the nadir of its repute among evolutionists in the early twentieth century. Russia was no exception, despite the aging Timiriazev thundering against what he believed reactionary forces bent on discrediting Darwinism. Perhaps the chief among the panoply of arguments advanced against natural selection was the absence of a clear correspondence between the organic diversity and the diversity of the environments. In particular, the characteristics that distinguish related species, and are utilized in taxonomic keys, appear to be neither useful nor harmful to their possessors. In other words, those characteristics are neutral traits, which natural selection can neither promote nor eradicate. Identical arguments, now in vogue among the so-called non-Darwinian evolutionists, who find no sense in the amino acid sequences in the proteins of different forms of life, are to me distinctly deja vu. The eminent systematist and zoogeographer Semenov-Tian-Shansky compared nature to an artist who creates aesthetically appealing forms for no reason other than their beauty. If a viable organism can be built this way as well as that way, both kinds of organisms will exist. Natural selection can only interdict what is inviable. The concept of mutation was still rather unfamiliar to at least the older generation of biologists in the 1920s, even though the theories of de Vries and of his Russian anticipator Korzhinsky were not unknown. However, in 1919 and 1922 Philipchenko published in Priroda, a journal analogous to Science or to Nature, excellent reviews of the works of the Morgan school on the genetics of Drosophila. To me these reviews were a revelation. To most senior biologists Drosophila mutants were a collection of monstrosities, of no significance for evolution.


To criticize the theory of natural selection was easy; to put something better in its place very difficult. Some of the critics embraced neo-Lamarckism, the inheritance of acquired traits. Smirnov, a brilliant young zoologist at Moscow University, engineered in 1926 the invitation to Kammerer to come to Russia to head a laboratory in which they would have worked to establish the validity of Lamarckism, about which they had not the slightest doubt to begin with. The plan came to naught owing to Kammerer's suicide. The Lamarckists were a small minority among biologists (though not, for a time, among philosophers and politicians). (See Gaisinovich, 1968; Bliakher, 1971.)

The most widespread view was that evolutionary changes are induced by the environments in which the organisms live. But the evolutionary responses to the environment occur neither via natural selection nor via the inheritance of acquired traits. It is rather by means of direct influences of the "geographical landscape" on the germ plasm of the inhabitants of a given territory. Berg gave a most articulate exposition of this view, which he accepted as a subsidiary to his nomogenesis theory. The geographical landscape subsumes "the external environment in the widest sense, physical as well as biotic factors united in one harmonious whole." The geographical landscape "affects organisms in an imperative manner, compelling all the individuals, so far as the organization of the species permits, to vary in a determined manner. There is no place here for chance: consequences follow with the same fatal constancy as chemical reactions or physical phenomena." Just how the "harmonious whole" changes the genotype was left quite vague. Those who attempted to explicate this puzzle (Berg was not one of them) postulated direct alteration of the genetic materials by environmental factors, rather than somatic or parallel inductions favored by the Lamarckians. Evidence that could be quoted in support was scanty in the extreme: the now discredited experiments of Tower on the beetle Leptinotarsa, and the also discredited induction of inheritable eye defects by antibodies in rabbits, alleged by Guyer and Smith. Muller's discovery of gene and chromosomal mutations in Drosophila induced by ionizing radiations was, of course, a firmly established demonstration of genetic changes by an environmental agency, but hardly relevant to the geographical landscape theory.


More interesting was Berg's theory of nomogenesis (1922). A man of splendid intellect and great personal charm. Berg developed his views in a book marshaling an abundance of evidence comparable to Darwin's On the Origin of Species. For a year or two after reading Berg's work I was close to becoming a partisan of nomogenesis. To Berg the phylogeny is basically similar to the ontogeny - the development follows paths predetermined by causes internal to the organism. The environment and natural selection permit or prohibit the survival of a new form but are not competent to bring it into existence. Evolutionary changes are subject to laws analogous to, and perhaps consubstantial with, those of embryonic development. With impeccable honesty. Berg admits that his putative "laws" leave unexplained the adaptedness of organisms to their environments. He postulates that adaptedness is an immanent property of living matter. Nomogenesis was attacked from all sides. Dialectical Marxists rightly saw that nomogenesis borders on vitalism. Lamarckians were displeased that Berg not so much denied as ignored acquired traits and somatic induction. His book has no reference to Kammerer's works being hotly debated at that time. Genetics also receives inadequate attention, which is hardly surprising because Berg wrote his book mostly before biological literature published abroad since 1914 had reached the blockaded Russia.

The Rise of Genetics

Scientists in Russia had an extraordinary experience of being almost wholly isolated from world science for about seven years, and then obtaining access at once to what was accomplished during this period by their colleagues abroad. To those interested in genetics and evolution this amounted to a sudden revelation. The review of the work of the Morgan school on Drosophila published by Philipchenko in 1922 was an eyeopener to many. Muller visited Russia in 1922 and left a fair number of cultures of Drosophila mutants at Koltsov's Institute of Experimental Biology in Moscow the seed from which grew the Russian school of Drosophila genetics. Moscow geneticists were generous with the gift; I was enabled to start working with Drosophila mutants in Kiev early in 1923, as were other converts to Drosophila studies elsewhere. I myself left Russia on the morning of December 4, 1927, having received a postgraduate fellowship to work one or two years in Morgan's laboratory. I did not suspect at all, when I left, that I might stay on after the expiration of the fellowship.

There were three nuclei of genetics research in the 1920s. By far the largest were the Institute of Applied Botany headed by Vavilov, with headquarters in Petrograd (later Leningrad), Koltsov's Institute of Experimental Biology in Moscow, and Philipchenko's department of genetics at the University of Leningrad. Vavilov, soon joined by Karpechenko, Levitsky, and numerous other botanical and agricultural geneticists, maintained his familiarity with current ideas and research in genetics and evolution, despite what to almost anybody else would have been a crushing load of administrative responsibilities. Koltsov's personal interests were in cell biology, in what later became molecular genetics, and finally in eugenics. Two of the several sections in his Institute of Experimental Biology were headed by Chetverikov and Serebrovsky. They attracted a number of talented young researchers, some of whom became eventually leading figures in evolutionary biology in Russia and abroad. Among them were Timofeeff-Ressovsky, Astaurov, Romashov, and Dubinin.

Sergei Chetverikov

Chetverikov's classic paper (1926) showed that evolution is brought about by natural selection acting on a store of genetic variability generated by mutation. Between 1930 and 1932, the same theory was expounded with greater mathematical refinements by Fisher, Wright, and Haldane. The four "founding fathers" thus provided the cornerstone of what later became the biological, or synthetic, theory of evolution. Chetverikov's work would have received much greater renown had it not been published in Russian in a journal with limited distribution abroad. As it was. Fisher, Wright, and Haldane were simply unaware of Chetverikov's contribution. It did, however, stimulate a great deal of research activity in Russia and eventually abroad. As a background against which to see the crosscurrents in evolutionary biology in Russia in the 1920s and 1930s, brief sketches of some of the personalities involved in the apparently sudden blossoming of evolutionary studies may be useful.

Chetverikov started as a naturalist working on systematics and ecology of Lepidoptera. He had a contemplative turn of mind and, although he was a steady worker, he published little. His interest in the expansions and contractions of the populations of various species in time led to the publication in 1905 of "Waves of Life," a rather short but allusive article. His great 1926 essay is only 52 pages long, although it contains enough material to make an at least medium-sized book. Chetverikov was at his best in face-to-face discussions with colleagues and students. His sharp and rigorous analysis of evolutionary problems was accompanied by constructive ideas and suggestions for research. He shared his ideas with his interlocutors with complete selflessness. Trying to establish his priority seemed to him unworthy of a scientist.

I had the privilege of several visits with him in his laboratory in Moscow. Though at that time I was only a bit more than half as old as he was, Chetverikov was unstinting with his time and inspiring with his advice. He disliked writing letters, however, which proved to be a blessing in disguise for me. Otherwise I might have shared the fate of Timofeeff-Ressovsky in Germany, instead of coming to work with Morgan in the United States. Chetverikov was an experimentalist as well as a theoretician. Even before the publication of his 1926 theoretical paper, he initiated a study of what later became known as genetic loads, in natural populations of Drosophila tnelanogaster. The results were reported briefly in a paper read in 1927 at the International Genetics Congress in Berlin, and in a very short paper published in Russian in 1928. In 1929 Chetverikov was exiled, because, it was rumored, he was denounced to the political police by one of his students. This incident occurred before Stalin's terror reached its peaks of brutality. Although Chetverikov was never able to develop the work begun so auspiciously with his many students in Moscow, he remained alive until 1959. Recognition, in the form of the Darwin medal award by the German Academy Leopoldina, reached him shortly before he died at the age of seventy-nine.

Alexander Serebrovsky

Serebrovsky accepted Chetverikov's theory of evolution. His own work developed quite independently and in somewhat different directions, however. He was an adherent of the now almost forgotten presenceabsence theory, originated chiefly by Bateson in England, which for reasons beyond my comprehension seemed to Serebrovsky particularly congenial with Marxist dialectics. This theory postulated that most mutations are minute deficiencies, little holes in the chromosomes. Serebrovsky attempted to measure the size of the assumed "hole," by estimation of the map distance in centimorgans between two marker genes on both sides of a mutant in Drosophila melanogaster. Together with his student Dubinin, he interpreted the so-called step allelism at the gene locus of scute in Drosophila in terms of overlapping deficiencies in a series of linearly arranged subgenes.

Of more lasting interest and significance are Serebrovsky's studies on gene geography, first published in Russian in 1927. Chicken populations in the Caucasus (Dagestan) are highly polymorphic, and the gene frequencies differ considerably in nearby mountain valleys, as well as elsewhere in the Soviet Union. This led Serebrovsky to formulate the concept of the gene pool, which he called the gene fund ("genofond"). His interest in the human gene pool landed him in trouble. Independently from Muller, Serebrovsky advanced a plan of improvement of human populations by means of artificial insemination of women with the semen of eugenically desirable donors. This plan was too much for the then poet laureate Demian Bedny (eventually himself destroyed in Stalin's terror). Serebrovsky was blasted and ridiculed in Pravda and elsewhere by totally ignorant hacks. When Lysenko became the ruler of biology, Serebrovsky turned to applied genetics and put forward the idea of pest control by means of release of masses of individuals that would produce sterility or lethality in the recipient populations. This idea was surely ahead of his time, but he had no opportunity to make even small-scale tests of its applicability. When genetics, particularly Drosophila genetics, was banned, Serebrovsky, the ardent believer in communism and dialectical materialism, died a forlorn and embittered man, just when Lysenko reached the peak of his glory.

Juri Philipchenko

Philipchenko built a school perhaps smaller than those of Chetverikov and Serebrovsky, but his impact on the evolutionary and genetical thought in Russia was, if anything, greater. He wrote six textbooks and numerous review articles on various aspects of genetics and evolution that were used in institutions of higher learning until Lysenko's pogrom. The first course of genetics in Russia was given by Philipchenko from 1913 until he was forced to resign his professorship at the University of Leningrad a year before his death in 1930. His books and lectures were didactically flawless: the depth and breadth of his scholarship were as admired by his students and colleagues as they were feared by his opponents. And yet this man of superb intellect fell short of fulfillment of his capabilities, in part because of his untimely death, and in part because of the skeptical and overcritical turn of his mind. He was always able to see a host of alternative explanations for any phenomenon and to poke holes in any theory. Yet while a deficiency of critical ability is fatal to a scientist, its excess may be frustrating; life is too short to test every alternative, and one has to accept the most probable one, subject, of course, to possible falsification.

Philipchenko welcomed Chetverikov's mutation-natural selection theory, but stressed that it might be only a part, and a rather minor part of the story. The genes and the Mendelian inheritance may be pertinent only to superficial traits, distinguishing individuals within a species or species within a genus; fundamental features, those of orders, classes, and phyla, may be inherited and changed by quite different and as yet unknown mechanisms. This view was a minority position but scientifically quite respectable in the 1920s. Similar opinions were held, for example, by the Danish geneticist Johannsen. Though a zoologist by training, in the last decade of his life Philipchenko worked on the genetics and ontogeny of certain characteristics in species of wheat and other grasses. The results were neither in accord with his expectations nor flatly contradictory. Had he lived longer he might well have changed his evolutionary credo. As it was, he still played a most useful role not only as a teacher but also as a stimulant of critical thought and research in evolutionary biology.

Between 1924 and 1927 I was the counterpart of an assistant professor in Philipchenko's department in Leningrad. Moscow is just a night railway journey away, so I visited there rather frequently. At that time at least, the second-class tickets were reasonably cheap. I did not, however, participate in the actual work done at the Koltsov Institute. When I visited there, I looked at what they were doing but I cannot claim the honor of having been one of Chetverikov's students. The subject of my own research in Philipchenko's laboratory in the years 1924-1927 was the effect of pleiotropic genes in Drosophila. However, I never lost my interest in Coccinellidae.

As an entomologist I had read On the Origin of Species, I believe in 1915, when I was all of fifteen years old. I also read German entomological literature because my main foreign language then was German. One of my beloved books was that of Standfuss, describing experiments on butterflies and moths. The transition from that to Drosophila was very easy. Berg's nomogenesis was a temporary infatuation: it sounded very remarkable and it took me some time to recover my independent judgment. Berg was a very impressive and certainly brilliant man.

The Great Institutes

Vavilov headed the Institute of Applied Botany (later the Institute of Plant Industry, still later the Lenin Academy of Agricultural Sciences). Koltsov was the director of the Institute of Experimental Biology. In its heyday Vavilov's institute had local sections and experiment stations in all parts of the Soviet Union and included among its research workers, if only on a part-time basis, nearly every geneticist and cytologist working with plants as materials. Koltsov's institute was not quite so large, but it had a powerful group of both young and old research workers. Research and thinking of the then avant-garde evolutionary biology were active in both institutions.

Vavilov and Koltsov were eminent scientists in their own right. As administrators, they necessarily sacrificed much of their personal research potentials for the sake of furthering the work of others. In a sense, their greatest discoveries were the outstanding creative scientists whom they chose as their colleagues and leaders of various research projects. In 1922 Vavilov established the law of homologous series in genetic variabilities. Notably in related species of cultivated plants, but also in wild species, the mutation process generates alleles of homologous genes that produce phenotypically similar variants. Artificial and natural selections then choose from this assortment the genotypes suitable for survival and exploitation. Some overenthusiastic reporters went so far as to compare this law with the periodic system of chemical elements. Vavilov's other major contribution, based on his extensive familiarity with cultivated plants of the world, was locating the geographic "centers of origin" of the domesticated forms. He was inclined to equate these centers with regions of maximum genetic variability in the respective species. For a few years preceding his arrest and death in prison, Vavilov was stripped of administrative work and devoted his time to studies of the evolution of cultivated plants; his publications of that period seem to lack much of his former ardor.

Levitsky and Karpechenko were two outstanding evolutionary cytogeneticists among Vavilov's cohorts. Levitsky, originally professor of botany at the Faculty of Agriculture in Kiev, published The Material Basis of Heredity (1924) and Cytological Basis of Evolution (1939). Karpechenko obtained fertile allopolyploid hybrids of radish and cabbage, a classic example of emergence of a full-fledged new species, by doubling the chromosomal complements in otherwise sterile interspecific hybrids. Both Levitsky and Karpechenko were articulate speakers and writers and stimulated a great deal of genetic and evolutionary research among botanists, not only in Leningrad but elsewhere in the far-flung system of Vavilov's experiment stations. Inevitably they ran afoul of Lysenkoism, were arrested in 1941, and died in prisons or concentration camps. Koltsov was a man of multifarious interests and knowledge, of imposing presence, and with the eloquence of a spellbinding orator. His public lectures were events memorable to his peers and to beginners alike. His main field of research was cell biology; some Russian writers claimed that Koltsov discovered the main features of the genetic code [see Ref.1], although he believed (as did biologists in general in his time) that the genetic information is stored in proteins rather than in the nucleic acids. He realized clearly the basic importance of Mendelian genetics for the elucidation of evolutionary processes and gave full support to Chetverikov, Serebrovsky, and other geneticists working in his institute and elsewhere. One of the Koltsovians, Timofeeff-Ressovsky, had a remarkable life; for about two decades after leaving Koltsov's institute he worked with great success in Germany, then spent some years in Soviet prisons, and finally emerged the most eminent evolutionary geneticist in the Soviet Union.

Eugenics seemed to be an important application of genetics to Koltsov. He was the organizer and the moving spirit of the Russian Eugenics Society and of its periodical, a perilous undertaking in the Soviet Union, especially when eugenics was among the slogans of the waxing Nazism in Germany. No matter how much effort Koltsov and Philipchenko made to separate their eugenics from the racist teachings of the Nazis, they were easy marks for the accusations of the Lysenkoists that all eugenics, and in fact all genetics, are pernicious inventions of the capitalists. The last years of Koltsov's life, though he avoided arrest and imprisonment, were poisoned by unceasing harassments. He was lucky to die of a sudden heart attack; his wife committed suicide within a few hours after his death.

The Flourishing State of Russian Genetics

One of the most impressive aspects of Russian genetics in the 1920s was the size of the genetics establishment. In England, few were interested in evolutionary genetics besides Bateson, Fisher, Haldane, Huxley, and Ford. In Germany, there was some interest in physiological genetics, but aside from Baur there was no population geneticist until Timofeeff-Ressovsky arrived from Russia.

In contrast, three schools flourished in Russia, each with numerous investigators. Numerically the largest school, of course, was that of Vavilov, although much of its research was applied. Chetverikov's group consisted in 1927 of twelve people doing Drosophila research, as well as five visitors to the group (Zhivago, a cytologist, Frolova, Serebrovsky, Sacharov, and Koltsov). Serebrovsky had his own group that included Dubinin among others. Frolova has the distinction of having discovered the difference between European Drosophila obscura and U.S. "obscura." When she was unable to cross the two stocks, she examined the chromosomes and found them to be quite different. She and Astaurov described the American sibling species as D. pseudoobscur a.

It is sometimes remarked that Russian geneticists seemed to be much more of the naturalist type than pure experimentalists, perhaps because of the great size and environmental diversity of the country. New and unusual animals and plants were steadily collected and brought for study in university laboratories, zoological and botanical museums, and marine institutes. Most biologists had experience working in the field and observing living beings in their habitats. Most young biologists traditionally spent a year or more abroad in "postdoctoral" work (although there were no doctorates in our sense), usually including visits to Naples' marine zoological station or similar institutions. It was taken for granted that a biologist must know animals or plants, or both.

Severtsev and Schmalhausen

Severtsev and Schmalhausen were evolutionary morphologists, working on comparative anatomy and embryology of vertebrates. Severtsev's interests centered on the relationships between phylogeny and ontogeny, and led him to conclusions in many ways resembling those of Rensch and Simpson published two decades later. Severtsev viewed the mechanisms that bring about evolution in the light of classical Darwinism; he regarded natural selection the leading agent and considered the inheritance of acquired traits neither proved nor completely ruled out. He was certainly not opposed to the genetical theories of evolution being developed by Chetverikov and others, but rather felt that geneticists should do their business while he and his collaborators were doing theirs. To enthusiastic young geneticists in the mid-1920s this attitude seemed narrow-minded if not outright retrograde. Philipchenko was among those who pointed out that the narrow-mindedness was more likely the foible of the young enthusiasts because evolutionary biology is multidimensional, not only a specialized branch of genetics.

Schmalhausen was one of Severtsev's pupils, but by the late twenties while a professor of zoology in Kiev, he became interested in the genetic factors bringing about evolutionary changes. As Severtsev's successor as director of the Institute of Evolutionary Morphology in Moscow, Schmalhausen published a series of books on evolution that belong to the period of the emergence of the synthetic theory rather than to its incipience. Schmalhausen's analysis of the varieties of natural selection is fully modern and has in no way lost its significance today. As could have been expected, Schmalhausen was attacked by Lysenkoists and expelled from the directorship of his institute, but fortunately avoided the arrest and imprisonment that then commonly befell biologists who refused to compromise their scientific and personal integrity. In his last years Schmalhausen became interested in cybernetics and its applications to evolutionary biology.

The Emergence of a Paradigm

In his essay on the Russian school of evolutionary genetics, Adams (1968) remarks: "The Russian school is important both because of what it ended and what it began. Many authors have alluded to the estrangement between two traditions in biology which characterized its history in the early decades of this century: the experimentalist' and the naturalist' traditions. It is significant, then, that the Russian school is one of the earliest to draw from both traditions in order to clarify the evolutionary process." Russian biologists habitually viewed the evolutionism in two contexts: that of its philosophical implications, and in the light of life as it exists in pristine nature. To be acceptable, a paradigm had to pass scrutiny from these points of view, in addition to being valid on purely factual grounds.

Why had such a paradigm emerged when it did and not earlier or later? Interest in evolutionary problems was traditional among Russian biologists since the nineteenth century. Except for the years during World War One and the subsequent revolutionary upheaval, Russia was not a scientific backwater. Current scientific literature in major European languages was not only available but it was read rather more systematically than is the foreign-language literature now in the United States. Acquaintance with the experimental work of the Morgan school, and with the findings of other geneticists in Europe and in the United States, became possible only in about 1921. However, this exposure would have been insufficient without Chetverikov's insight and without the experiments on the genetics of natural populations that he promptly undertook to test the validity of his theoretical construct.

It now seems almost unbelievable that quite authoritative biologists in the 1920s and 1930s in Russia as well as in Europe and the United States contended that the phenomenon of mutation has no bearing whatever on evolution. Drosophila mutants described by the Morgan school were monstrosities, and they were all found in laboratory bottles, not in natural populations, products of some kind of disruption of hereditary materials in highly artificial environments. The claim of de Vries and his followers that new species arise by single mutational leaps seemed incompatible with everything that systematists and paleontologists had learned about species in living and fossil forms. It was therefore quite logical and necessary that the experimental work of Chetverikov and his collaborators and students was directed toward testing for the presence of mutants in natural populations of Drosophila. They explicitly considered their work to be the decisive test of the validity of the theory, and the results of the test turned out to be positive. After Chetverikov ran afoul of the political police and was banished from Moscow, the work went ahead under the direction of Timofeeff-Ressovsky in Germany, and by Dubinin, Romashov, and others in the Soviet Union.

It is tempting to represent a scientific discovery of a given person or of a group of persons at a given time too neatly as an automatic consequence of a certain scientific tradition. After all, the basic features of the mutation-natural selection theory of evolution were discovered almost simultaneously and certainly quite independently from Chetverikov by Fisher, Wright, and Haldane. The personal backgrounds and the scientific traditions of this Anglo-American trinity were quite different from Chetverikov's, as well as from each other. It is well known that Mendelism and genetics in general were, by a singular miscomprehension, regarded as contradictory to Darwin's theory of natural selection in the early decades of the twentieth century. The short but important theoretical paper by Hardy in 1908 should have dispelled this miscomprehension, and in fact it served as a point of departure for Chetverikov as well as for the other founding fathers (the parallel work of Weinberg was not then considered). By the 1920s the mutation-natural selection theory was definitely in the air, and four persons independently seized it. Analogously, in the thirties and forties, what we now call the synthetic theory was in the air, a logical complement to the Chetverikov-Fisher-Wright and Haldane theoretical base, and at least eight biologists of different backgrounds and traditions have brought it down to earth. Some of them, fortunately, are still alive in the 1970s and can tell their stories.


Because of Dobzhansky's death before his selection had been fully prepared for publication, there is no list of references such as appears after the other selections. Many of the books and articles cited by Dobzhansky appear, however, in other chapters of this book [Ernst Mayr etc. The Evolutionary Synthesis Perspectives on the Unification of Biology, 1980; The second edition 1998], and complete information is given there. Eds.


Ref1. (Yury Neretin) ... some Russian writers claimed that Koltsov discovered the main features of the genetic code. Simon Shnoll claims that the idea that genetics is based on duplication of long moleculas was firstly formulated by Koltsov . Later it became well-known in Europe due to Timofeeff-Resovsky.

A quotation

Этот парадокс — малое число молекул и множество признаков — впечатался а сознание Кольцова. Он думал о нем многие годы, пытаясь разрешить разными способами. Он опубликовал свое решение в 1927 году в докладе на 3-м Всесоюзном съезде зоологов, анатомов и гистологов, т.е. через 34 года. Ответ этот звучит торжественно. Сущность его такова: признаки, передаваемые по наследству, определяются линейным расположением мономеров в полимерных молекулах. (Кольцов думал, что это последовательность аминокислот в полипептидах.) Эти гигантские полимерные молекулы (их в самом деле мало в клетке) размножаются по принципу матриц: свободные мономеры из раствора сначала выстраиваются вдоль «родительской» молекулы — матрицы, выстраиваются в соответствии с последовательностью мономеров в этой матрице. А затем образуются химические связи, сшивающие прилипшие к матрице мономеры с образованием точной «дочерней» копии. Эти полимерные молекулы размножаются как кристаллы. Читателю ясно, что это — формулировка самой главной идеи биологии XX века.

Идея матричного синтеза была забыта не понявшими ее современниками. Забыта не всеми. В 1925 году учеников Н.К.Кольцова и С.С.Четверикова — супругов Тимофеевых-Ресовских по рекомендации Н.К.Кольцова и Н.А.Семашко советское правительство командировало для работы в Германию, в институт профессора Фогта. Громогласный, энциклопедически образованный Н.В.Тимофеев-Ресовский рассказывал об этой идее на своих ставших знаменитыми семинарах в Берлин-Бухе, в своих многочисленных докладах на конференциях, семинарах, съездах в разных странах, в том числе на семинарах Нильса Бора в Копенгагене. Идея матричного синтеза особенно легко усваивалась физиками. Это можно понять. «Нет пророка в своем (биологическом) отечестве». Пророк — биолог в отечестве физиков — тут ему внимают с почтением и доверием. Да и по существу — главные доводы в пользу матричной концепции из области физики. Совершенно невероятно выстраивание в правильной последовательности мономеров в полимерной цепи в обычных химических реакциях...

Когда великий физик Эрвин Шредингер читал свои лекции (по теоретической биологии!) в университете, его предметом и был взгляд на биологию с точки зрения физика. Он основывал свои взгляды на представлениях об одномерном кристалле, на термодинамике и матричной концепции, полагая, что она, эта концепция, общепринята у биологов. Когда его лекции были опубликованы в виде знаменитой книги «What is Life? The Physical Aspect of the Living Cell», Дж.Б. Холден откликнулся на нее статьей в «Nature»: концепция не общепринята в биологии, а принадлежит великому российскому биологу Кольцову!