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Vavilov, Nikolay Ivanovich

VAVILOV, NIKOLAY IVANOVICH

(b. Moscow, Russia, 25 November 1887; d. Saratov, U.S.S.R, 26 January 1943)

botany, agronomy, genetics, phytogeography.

Vavilov was the oldest son of Ivan Ilich Vavilov, owner of a shoe factory and at one time a member of the Moscow City Duma. His brother Sergey became a well-known physicist and president of the Soviet Academy of Sciences; his sister Lydia, a microbiologist, died of typhus during World War I; his sister Alexandra was a physician. Vavilov was married twice, the second time to his co-worker Elena Ivanovna Barulina; his sons, Oleg and Yuri, both became physicists.

Vavilov’s career was shaped by his lifelong dream of mobilizing the sciences in order to transform Russian agriculture. His meteoric rise to prominence following the Russian Revolution began twenty years of prodigious activity involving the creation of a vast research network. The picture that emerges from contemporary accounts and memoirs by some of his many co-workers suggests that his remarkable success as a scientific entrepreneur resulted from his boundless energy and winning personality. During this period Vavilov apparently took no vacations and slept no more than four or five hours a night, often on the train between Moscow and Leningrad or on a sofa in his office, where he lived much of the time. Heterodox and undogmatic, he had an excellent memory for names and faces, and seemed to dazzle almost all those with whom he came into contact. Most regarded charm as his defining characteristic,

In 1906 Vavilov graduated from a commercial high school and entered the Moscow Agricultural Institute (now the Timiryazev Academy of Agriculture). There he organized and led a student science club that conducted botanical and geographical expeditions to the Caucasus, Transcaucasia, and other regions of Russia. As a thirdyear student he presented a paper on Darwinism and experimental morphology at the institute’s Darwin centenary celebration. His student thesis on destructive garden slugs of Moscow Province was awarded the A. P. Bogdanov Prize of the Polytechnical Museum.

After graduating in 1911. Vavilov remained with D. N. Pryanishnikov in the department of special agriculture to prepare for an academic career and conducted research at the institute’s Moscow selection station, working with its director, D. L. Rudzinsky. In 1911–1912 he taught agriculture at the Golitsyn Courses for Women. He moved in 1912 to St. Petersburg, where he was a probationary employee at the Bureau of Applied Botany of the Ministry of Agriculture, headed by R. E. Regel, and at its Bureau of Mycology and Phytopathology, headed by A. A. Yachevsky.

In 1913 the department of special agriculture sent Vavilov to England. At Cambridge he attended the lectures of R. C. Punnett and worked in the laboratory of R. D. Biffen. He also worked at the John Innes Horticultural Institution at Merton (London) under William Bateson, whom he always regarded as his teacher. While in England he studied immunity in wheat, using the collection of John Percival, then the world’s largest, and conducting experiments on plant immunity simultaneously in Moscow and England.

On his return to Russia, which was hastened by the outbreak of World War I, Vavilov passed the examination for the master’s degree and completed his thesis, Immunitet rasteny k infektsionnym zabolevaniam (“Plant Immunity to Infectious Diseases”). He continued to teach in the department of special agriculture during the war (he was not drafted because of an eye problem), and in 1916 organized and conducted a botanical and geographical expedition to Iran and the Pamir.

In 1917, Vavilov was simultaneously appointed professor of genetics, selection, and special agriculture at the Voronezh Agricultural Institute and professor of agronomy and selection at Saratov University. During the next four years he organized and directed the Saratov branch of the department of applied botany and selection of the Agricultural Scientific Committee of the Commissariat of Agriculture of the Russian Soviet Federated Socialist Republic (formerly the Bureau of Applied Botany of the Ministry of Agriculture).

Vavilov came to the attention of the government in 1920–1921 and rapidly rose in prominence. Following Regel’s death in 1920, Vavilov moved to Petrograd to replace him as director of the department of applied botany. In 1924, with Lenin’s support, he reorganized the department into the All Union Institute of Applied Botany and New Cultures, which in 1930 was renamed the All Union Institute of Plant Breeding (VIR). From 1923, Vavilov was also director of the State Institute of Experimental Agronomy at Leningrad. He reorganized this institute in 1929, forming the All Union Lenin Academy of Agricultural Sciences (VASKhNIL), with the VIR at its core. He served as its first president until 1935, by which time it had more than 400 experimental stations and a staff of more than 20,000.

During this period Vavilov earned his reputation as “the most widely traveled biologist of our day.” In connection with the VIR’s collecting activities, he organized more than 180 expeditions, 40 outside the Soviet Union. Collecting expeditions in which he participated include those of 1920 (Astrakhan, Tadzhikistan, and the Pamir), 1921–1922 (United States, Canada, England, France, Germany. Sweden, and Holland). 1924 (Afghanistan), 1925 (Khiva and Uzbekistan), 1926–1927 (Algeria, Tunisia, Morocco. Egypt, Syria. Palestine. Trans-Jordan, Greece, Crete, Capri, Italy, Sardinia. Sicily, Spain. Somaliland, and Ethiopia), 1927 (Germany), 1929 (western China, Taiwan, Japan, and Korea). 1930 (United Stales. Central America, and Mexico). 1931 (Denmark and Sweden), 1932– 1933 (Cuba, Peru, Bolivia, Chile, Brazil, Argentina, Uruguay, Trinidad, and Puerto Rico), 1934 (Caucasus), and 1940 (Belorussia and the western Ukraine). Vavilov was elected president of the U.S.S.R. Geographical Society in 1931 and served through 1940. His own account of his travels is Pyat kontinentov (“Five Continents”), published posthumously in 1962.

As a result of such expeditions, by 1940 the VIR had amassed a collection of more than 250,000 specimens, including 36,000 of wheat, 10,022 of maize, 23,636 of legumes, 23,200 of grasses, 17,955 of vegetables, and 12,650 of fruits. These samples were distributed throughout the system of experimental stations for cytological and genetic analysis, breeding and hybridization experiments, and testing under various growing conditions. They formed the basis for the institute’s —and Vavilov’s own —numerous publications.

Vavilov was elected a corresponding member of the Soviet Academy of Sciences in 1923 and a full member in 1929. Following the death of Y. A. Filipchenko in 1930, he became head of the genetics section of the Commission for the Exploitation of Productive Forces, transforming it into the Soviet Academy’s Laboratory of Genetics, which became in 1933 its Institute of Genetics, which he directed until 1940. There he attracted a first-rate group of researchers, including H. J. Muller (1933– 1937), and established the leading center of theoretical genetics research in the Soviet Union.

Vavilov won many awards, including the N. M. Przhevalsky Gold Medal of the Geographical Society of the U.S.S.R. (1925), the Lenin Prize (1926), and the Great Gold Medallion of the All-Union Agricultural Exhibition. He was elected a foreign member of many scientific societies, including the International Agrarian Institute of Rome (1924), the Ukrainian Academy of Sciences (1929), the Czechoslovak Academy of Sciences (1936), the Indian National Academy of Sciences (1937), the Royal Society of Edinburgh (1937), the Royal Society of London (1942), and the Linnean Society of London, the New York Geographical Society, the American Botanical Society, the Mexican Agricultural Society, and the Royal Spanish Natural History Society. He was awarded honorary doctorates by the University of Brno (1936) and the University of Sofia. In 1932 Vavilov served as vicepresident of the Sixth International Congress of Genetics (Ithaca, New York) and in 1939 was elected president of the Seventh International Congress, although he was unable to attend.

During his lifetime Vavilov published more than 350 scientific books and articles, many of which, according to his own testimony, were written or dictated in the evening, at odd moments, or while traveling. His writings manifest an encyclopedic command of agronomic and biological literature. Vavilov could read, write, and speak English, French, and German; and many of his publications in Russian include texts or lengthy abstracts in English. He served as the editor of several journals. most notably Trudy po prikladnoi botanike, genetike i selektsii (“Works on Applied Botany, Genetics. and Selection”), and oversaw the translation and publication of works by J. G. Kölreuter, Darwin, Mendel. T. H. Morgan, and H.J. Muller.

Plant Immunity . Vavilov’s earliest scientific work concerned the genetic basis of plant immunity. His first studies were undertaken during his school days in Moscow and with Biffen in England (1913– 1914). They were published in a series of studies that appeared in 1913– 1919, notably “Immunity to Fungous Diseases as a Physiological Test in Genetics and Systematics, Exemplified in Cereals” (1914). In this paper Vavilov focused on taxonomic characters in relation to susceptibility, establishing that in the one variety of Triticum vulgare (out of 580) that was immune to mildew, immunity was inherited as a single recessive character. In this and in Immunitet rasteny… Vavilov developed the concept of the degree of specialization of parasitic fungi, postulating that the more species and genera a parasitic fungus attacks, the less likely it is that resistant varieties will be found in any of the host species.

Vavilov believed that immunity had to be viewed in an evolutionary context: “Immunity is associated with the biological specialization of parasites to genera and species and is caused by divergence of the host and parasite in their evolution.” He sought to use reactions to parasites, especially to narrowly specialized parasites, as an aid to systematic botany. One product of Vavilov’s research was his discovery of a species of wheat, Triticum timopheevi, that is resistant to most of the important diseases of wheat and that has subsequently been used as a source of disease resistance by breeders in the Soviet Union and elsewhere. Like his other works. Vavilov’s monographs on plant immunity attempt to synthesize the literature in the field in order to find regularities that would aid the breeder. According to Vavilov, his studies in this field led him naturally into his studies in genetics and plant geography.

Law of Homologous Series . Vavilov first presented his “law of homologous series in hereditary variation” at the Third All-Russian Conference on Selection and Seed-Growing at Saratov (1920). His paper was published in Russian that year, two years later in English, and in its enlarged and revised version in 1935. Punnett later suggested that Vavilov may have gotten the idea for his law in 1913, during one of Punnett’s lectures at Cambridge on parallel series of variations in species of rodents and butterflies. This is not certain, however, since the idea was common at the time. As Vavilov himself noted, the phenomenon of “parallel” or “analogous” variations in different races, breeds, species, genera, and families of animals and plants had been commented on by many earlier biologists, including Darwin, Mivart, Naudin, B. D. Walsh, de Vries, J. Duval Jouve. G. H. T. Eimer, Cope. E. Zederbauer, and P. A. Saccardo.

In his paper Vavilov sought, however, to establish a more general law. By collecting information on morphological variants or mutations in various cereals, he showed that many variants found in a given species also are found in closely related species—the more closely related the species, the greater the number of common variants. Vavilov displayed the data in tabular form and thought he had found a law or regularity (zakonomernost) in the range of variability within plant species. He himself drew the parallel between his “homologous series” and the periodic table of the elements, and stated his conclusions in symbolic form reminiscent of chemistry, thus leading the agronomist V. R. Zalensky to comment. “Biology has found its Mendeleev.” Vavilov believed that just as the periodic table had served as the basis for predicting the existence of undiscovered elements, his “homologous series” tables could be used to predict the existence of undiscovered plant forms that would fill the “gaps” in his system. He saw his law as evidence that despite the apparently random and unpredictable character of mutation and variation, the variability of species manifests an overall regularity that contributes to the orderliness of the evolutionary process.

A number of other investigators saw in Vavilov’s law a challenge to the traditional Darwinian view of evolution as governed by natural selection. L. S. Berg regarded parallel variations as evidence that evolution was an unfolding along predetermined lines. Filipchenko saw the law as evidence that the characteristics that determine species are different from those establishing higher taxonomic categories, thereby agreeing with E. D. Cope and W. L. Johannsen that microevolution and macroevolution are fundamentally different processes with fundamentally different causes. Others understood the law to suggest that mutations can be predicted, or that there are definite limits to species variability.

Responding in 1936 to criticism, Vavilov admitted that in formulating the law in 1920 and 1922, the “striking parallelisms, down to the most minute details” that he had observed in cultivated plants, had led him to be “unnecessarily categorical”:

We underestimated the variability of the genes themselves…. At that time we thought that the genes possessed by close species were identical; now we know that this is far from the case, that even very closely related species which have externally similar traits are characterized by many different genes. By concentrating our attention on the variability itself, we gave insufficient attention to the role of selection… [“Puti sovetskoy selektsii,” in Spornye voprosy genetiki i selektsii (1937)].

In the revised (1935) version of the theory, Vavilov distinguished more carefully between phenotypic and genotypic variation, and between “homologous” variation (caused by the same genes) and “analogous” variation (similar phenotypic traits caused by different genes). He also added information on parallel variations in additional species (such as those in melons, worked out in 1925) and emphasized the importance of selection. Vavilov modified his analogy between genetics and chemistry, commenting:

In the last decade genetics has moved ahead rapidly and to some degree begins to resemble chemistry, at least the chemistry of complex organic compounds…, The regularities in plant polymorphisms…, may to some measure be compared with the homologous series of organic chemistry, for example hydrocarbons…. As with chemical structures, different forms of plants and animals are characterized by a physical structure and bring to mind the systems and classes of the chemistry of crystals [Zakon gomologicheskikh ryadov v nosledstrennoy izmenchivosti, 2nd ed. (Moscow-Leningrad, 1935)].

Despite these revisions, however, he maintained the law’s validity and importance, and gave several examples of the discovery of forms that had been predicted on the basis of his theory.

The law of homologous series in hereditary variation was the basis of Vavilov’s conception of the species, most fully expressed in his 1931 monograph Linneevsky vid kak sistema (“The Linnaean Species as a System”). Vavilov regarded the “Linnaean species” (a synonym for Lotsy’s term “linneon”) as “a complex system of forms, the structure of which conforms to the law of homologous series.” For him the species was not an arbitrary division of convenience but a product of a dynamic evolutionary process operating over time and space, a process that both determined its genetic composition and gave it a real existence at any given moment.

Throughout his career Vavilov held what he regarded as a Darwinian view of evolution. As expressed in 1940, it envisioned hereditary variability as the basic material for both natural and artificial selection; the generation of new forms is increased by hybridization: and further evolution proceeds by the dispersion of species, their occupation of new territory, ecological factors, and geographical isolation. For Vavilov “the key factor in evolution, adaptation, and speciation is natural and artificial selection.”

Centers of Origin of Cultivated Plants . Vavilov’s work on the origin of cultivated plants was largely responsible for his worldwide reputation. His interest in the problem dates from his 1917 article proposing that rye originated as a weed in the southern wheat fields and gradually supplanted wheat by climatic selection in northern and mountainous regions, to which it was better adapted. His first general theoretical statement on the subject was begun in 1924 and published in 1926 as Tsentry proiskhozhdenia kulturnykh rasteny (“Centers of Origin of Cultivated Plants”), which in the same year brought Vavilov one of the first Lenin Prizes. His views were elaborated and modified in subsequent monographs (1927, 1931, 1935, 1940).

Vavilov dedicated his book to Alphonse de Candolle, whose Origine cies plantes cultivées (1882) was the major work in the field prior to Vavilov’s. Whereas Candolle had sought to establish centers of origin by using archaeological, historical, and linguistic—as well as botanical—data, Vavilov used these only as “correctives” to his “differential systematic geographical method,” which relied heavily on genetic and cytological analysis. As outlined in 1926, this method involved the division of plants into linneons and genetic groups by means of systematic morphology and cytological, genetic, and immunological analysis; the establishment of their geographical ranges; the “detailed determination of the varieties and races of each species (more precisely, the heritable varying traits and the general system of hereditary variability)”; and the geographical mapping of the variant forms of a plant group. as well as those of closely related varieties and wild species, in order to determine “geographical centers of concentrated diversity” (areas that contain the greatest “diversity of endemic varieties”), which Vavilov believed to be “as a rule, the primary center of origin.” In 1927 he added, “The primary centers often include a large number of genetically dominant traits.”

Using this method, Vavilov found that the centers of origin of many plants coincide with one another and with areas known to be sites of early civilization. In 1926 he identified five such primary centers: southwestern Asia (from India through Central Asia to the Black Sea); southeastern Asia (China. Korea, and Japan); the Mediterranean coast; Abyssinia and Eritrea; and the mountainous regions of South America and Mexico. Following his expeditions to the Americas and Asia, he increased the number of primary centers to eight (1935), dividing the American center into two (Mexico and Central America, and the Andes) and the southwest Asiatic into three: the Indian (which now included Indochina and the East Indies); the Central Asiatic (northwest India through Uzbekistan); and an area running from eastern Turkey through the Caucasus to Afghanistan. In addition, within the Andean or South American center he acknowledged distinct regions in which different plants had originated (Peru. Bolivia, Ecuador; Chile: Brazil and Paraguay). By 1940 his classification listed thirteen such regions, organized into seven basic centers.

According to Vavilov’s final version (1940), these seven centers are the southwest Asiatic (greatest number of species of wheat, barley, rye, and many fruits); the south Asian tropical (sugarcane, rice, many tropical fruits and vegetables); the east Asiatic (millet, soy, many vegetables and fruits); the Mediterranean (the olive, many vegetables, fodder, and certain fruits); the Abyssinian (teff, the Abyssinian banana [ensete], several species of sorghum, and some species of hard wheat); the Central American (corn, upland cotton, many species of beans, pumpkins, tobacco, cocoa, sweet potato, peppers, and tropical fruits [guava, soursop and sweetsop, avocado]); and the Andean (potatoes, okra, the cinchona tree, and the coca shrub). In all, by 1935 Vavilov believed that he and his co-workers had located the center of origin of more than 600 species of cultivated plants.

For Vavilov these primarily mountainous regions, with their wide variety of environmental conditions and natural isolating barriers, provided ideal conditions both for speciation and for human cultivation of new breeds. He never stated why he regarded “centers of diversity” as “centers of origin”; but by 1931 he apparently had recognized the importance of isolation in producing the diversity that he observed in these regions: “Schematically… as a result of inbreeding and mutation, advantageously recessive forms separate out and are formed toward the periphery of the ancient range of a species of cultivated plant under isolation (on islands, in mountains)” (“Rol Tsentralnoy Azii…,” in Trudy po prikladnoi botanike, genetike i selektsii, 26 , no. 3 [1931]). Vavilov saw these centers of origin as containing the full variability and “hidden potential” of the various species,

In December 1936 Vavilov acknowledged that in the preceding decade his thinking on the subject had changed.

At the beginning, we assumed that the majority of genes distinguishing current cultivated plants were really located at these centers, assuming that races and varieties dispersed from definite regions exactly as had whole species. Like geneticists in the first decade of this century, we conceived of genes as being more stable than they later turned out to be. In fact, in these centers we found an enormous number of genes, even more than we supposed at the outset, but at the same time newly formed genes, often remarkably valuable, were found in secondary regions on the periphery, and in the area between the periphery and the centers [“Puti sovetskoy selektsii,” in Spontye voprosy genetikii seiektsii(1937)].

For Vavilov his studies not only served to clarify the evolutionary process by demonstrating how species are formed but also had a direct “utilitarian goal”: by making the full range of hereditary diversity of a species available lo the breeder, the theory provided the “key to the mobilization of the plant resources of the globe.”

Selection as a Science . Throughout his career Vavilov sought to transform Soviet agriculture by creating new breeds of plants suited to its various environmental conditions. In 1912 he wrote that genetics “gives the foundations for the planned intervention of man in the creation of nature, the rules that govern the creation of forms.” In 1925 he held forth “the possibility, in Timiryazev’s words, of ’sculpting organic forms at will,’” suggesting that “in the near future man will be able, by means of crossing, to synthesize forms such as are absolutely unknown in nature.” At other times he saw his task as “the planned and rational utilization of the plant resources of the terrestrial globe” (1917), or finding “the way to seize the bastions of the fortress” of the plant kingdom (1926). He regarded “the complete control of the organism” as “the final purpose of modern biology” (1934).

Vavilov’s approach to this task was synthetic, in 1931 he wrote:

Biological phenomena connected with speciation are too complex to be reduced to simple physicochemical processes. But even complex biological processes manifest definite rules and laws which the biologist must study and which lead the investigator to the mastery of the formative process [Linneevsky via kuk sistema (1931)].

He looked upon his law of homologous series and his theory of the centers of formation of cultivated plants as precisely this kind of biological law or rule.

In the mid-1930’s Vavilov sought to characterize selection as an independent scientific discipline. In his pamphlet Selektsia kak nauka (“Selection as a Science,” 1934), he presented what he considered to be its seven basic divisions: phytogeography; studies of hereditary variability (the laws of heredity and the study of mutations); studies of the influence of the environment in creating varieties and affecting their development; hybridization theory; selection theory (centering on different forms of plant reproduction); studies of selection for particular properties (such as immunity, resistance to cold or drought, photoperiodism, and chemical composition); and the study of selection in particular plants.

As conceived by Vavilov, selection was a highly complex science. For controlling heredity it drew on genetics, cytology, and embryology; and in the actual process of selection it drew on physiology, chemistry, technology, phytopathology, and entomology, “transforming and differentiating them in accordance with the final task of creating a variety.” He regarded selection as a natural extension of Darwin’s theory of evolution and defined selection succinctly as “evolution directed by the will of man.”

As director of the VIR, Vavilov put his concept of selection into practice. For him the ultimate purpose of the expeditions he sent or led was to provide Soviet breeders with the full potential of the species, the raw material for sculpting new breeds. Beginning in 1923, he conducted experiments in “geographical sowing” that involved growing varieties at more than 100 sites throughout the Soviet Union in order to ascertain their suitability for planting under various conditions. He set up cyclical interbreeding of geographically close and distant forms of cereal grains, green beans, and flax in both fields and hothouses.

On the basis of these studies, Vavilov developed an agroecological classification of cultivated plants. The data for such a classification were published in brief form in 1940 in The New Systematics and fully in 1957 as Mirovye resursy… (“World Resources…”). In this work Vavilov distinguished ninety-five types of ecological areas in relation to climate, soil, and geography, and the varieties of cereal grains, legumes, and flax best suited to them, as well as the role of selection in creating especially adapted varieties.

Vavilov’s undertaking was possible only because of the manpower and resources at his disposal, and the latter were made available to him because of his promises to transform Soviet agriculture. In a 1931 newspaper article he wrote that “in practice, agronomy knows no bounds” and suggested that “the cultivation of vegetables, root and tuberous plants, and fodder grasses can go right up to the northernmost boundaries of the Eurasian landmass, not only to the northern boundaries of the taiga… but into the tundra.” Vavilov apparently believed that the task could be accomplished within a few decades and that the prospects were limitless.

Science and Politics . Throughout most of his career, Vavilov felt that the Soviet Union gave scientists and their work the greatest respect of any country in the world and offered the widest opportunities for serving mankind; therefore the cruelties of the regime had to be overlooked. In 1920 his presentation of his law of homologous series led conference delegates to send telegrams of praise to the commissars of agriculture and education. Sometime during this period, perhaps in connection with the 1921 famine, he apparently came to the attention of Lenin, who had been impressed by K. A. Timiryazev’s translation of William Sumner Harwood’s The New Earth: A Recital of the Triumphs of Modern Agriculture in America (New York. 1906). During the subsequent fifteen years Vavilov enjoyed the strong support of the government, which financed his rapidly expanding activities. He quickly grew in prominence and influence, serving as one of the few non-Communist members of the Central Executive Committee of the government (1926–1935) and, for a time, on the Central Executive Committee of the Russian Soviet Federated Socialist Republic (1927– 1929).

From 1928 to 1932, when the Academy of Sciences was “Bolshevized” and greater ideological conformity began to be demanded of scientific specialists, Vavilov started to introduce the language of dialectical materialism into his writings, offering “The Linnaean Species as a System” (1931) as a model for fellow scientists.

Beginning in 1931, Vavilov attracted criticism for failing to produce desired agricultural results. In response, he pointed to what had already been achieved and gave increasing emphasis in his public presentations to techniques that could yield immediate benefit. In so doing, he helped bring to prominence T. D. Lysenko, who had become known in the Soviet Union in the late 1920’s and early 1930’s for his work on “vernalization” (yarovizatsia), the treatment of seeds before planting to alter their development in such a way as to permit additional crop yields. In an article in Izvestia (6 November 1933), he hailed Lysenko’s method as a “revolutionary discovery of Soviet science” that had given “brilliant results” in tests conducted during the preceding year on “tens of thousands” of varieties, and that had made possible the cultivation of wheat from Afghanistan, India, Australia, and South Africa within the Arctic Circle. In 1935 Vavilov termed Lysenko’s method a “powerful tool” in the selection of many plants and foresaw a “total revision” of the classification of the world’s plant resources on the basis of their reaction to vernalization. He suggested that Lysenko’s theory of the “stage development” of plants “also opens exceptional possibilities… for the utilization of the world’s plant resources.”

In 1935 Vavilov was attacked for Soviet agricultural failures by Lysenko and his follower I. I. Prezent, among others. In particular, he was accused of pandering to foreign science, of wasting government money on useless collecting expeditions abroad, and of espousing “idealist” theories of homologous series and centers of origin, which one writer characterized as “fiascoes.” The attacks on Vavilov were related to more general attacks on genetics as being a formalistic, “idealistic” science contrary to dialectical materialism and Darwinism, and linked to foreign influences, notably fascism.

Vavilov lost his posts as member of the government’s Central Executive Committee and as president of VASKhNIL in 1935. His replacement in the latter post was A. I. Muralov; following Muralov’s arrest in July 1937, it was G. K. Meister; and after Meister’s arrest in February 1938, Lysenko assumed the post. Vavilov may have been under arrest briefly in 1936. The Seventh International Congress of Genetics, scheduled to be held at Moscow in August 1937, was postponed and was finally held two years later at Edinburgh. As president of VASKhNIL, Lysenko interfered in Vavilov’s work at the VIR by appointing one of his followers as Vavilov’s assistant, firing members of Vavilov’s staff, and generally undermining his direction of the institute.

Until 1939, Vavilov defended genetics and his own work but did not strongly criticize Lysenko, calling instead for cooperation and urging more objective discussions. In that year, however, Vavilov became much more critical of Lysenko. Zhores Medvedev quotes the following line from a speech Vavilov made to a session of the regional bureau of the VIR in March 1939: “We shall go to the pyre, we shall burn, but we shall not retreat from our convictions.” The statement proved to be prophetic. On 6 August 1940, while on a collecting expedition in the western Ukraine near Chernovtsy, Vavilov was arrested by agents of the NKVD; shortly thereafter his associate G. D. Karpechenko and others were arrested.

On 9 July 1941 the Military Collegium of the Supreme Court found Vavilov guilty, under Article 58 of the Soviet constitution, of belonging to a rightist organization, spying for England, leading the “Peasant Labor Party,” conducting sabotage in agriculture, and maintaining links with émigrés, for which crimes he was sentenced to death. Through the efforts of his brother Sergey and his former teacher Pryanishnikov, his sentence was commuted to ten years’ imprisonment. Following his sentencing he was moved to the inner prison of the NKVD in Moscow. In October 1941, during the evacuation of prisoners to the interior of the country, Vavilov was moved to the Saratov prison and was placed for several months in an underground death cell, where he suffered from malnutrition. He died in prison on 26 January 1943 and probably was buried in the Voskresensky Cemetery in Saratov.

After Stalin’s death Vavilov’s case was reopened on the initiative of his family. On 2 September 1955 he was posthumously rehabilitated by the Soviet Supreme Court, and a week later the Presidium of the Academy of Sciences placed him on its list of deceased members. Since 1957 most of his works have been published, many for the first time. In December 1967 the new All-Union Society of Geneticists and Selectionists was named for Vavilov.

In the West and the Soviet Union, Vavilov has come to be regarded as one of the outstanding geneticists of the twentieth century, a symbol of the best aspects of Soviet science, and a martyr for scientific truth.

BIBLIOGRAPHY

I. Original Works. Vavilov’s more than 350 publications include the following early works: Golye slizni (ulitki), povrezhdayushchie polya i ogorody v Moskovskoy gubernii (“Destructive Slugs of the Fields and Gardens of Moscow Province”;s Moscow. 1910): “Genetika i ee otnoshenie k agronomii” (“Genetics and Its Relation to Agronomy”), in Otchet Golitsynskikh zhenskikh kursov za 1911 god po khozyaystvennoi i za 1911–1912 uchebnyi god po uchebnoi chasti (“Report of the Golitsyn Courses for Women for the 1911 Economic Year and the 1911–1912 Academic Year”; Moscow, 1912). 77–87; “Immunity to Fungous Diseases as a Physiological Test in Genetics and Systematics. Exemplified in Cereals,” in Journal of Genetics, 4 , no. 1 (1914), 49–65; “O proiskhozhdenii kulturnoy rzhi” (“On the Origin of Cultivated Rye”), in Trudy Byuro po prikladnoi botanike, 10 , nos. 7–10 (1917), 561–590; “Sovremennye zadachi selskokhozyaystvennogo rastenievodstva” (“Current Tasks of Agricultural Plant Breeding”), in Selsko-khozyaistvennyi vestnik Yugo-Vostoka, nos. 19–21 (1917), 3–10; and Immunitet rasteny k infektsionnym zabolevaniam (“Plant Immunity to Infectious Diseases”; Moscow, 1919).

Works from the 1920’s are Zakon gomologicheskikh ryadov v nasledstvennoy izmenchivosti (“The Law of Homologous Series in Hereditary Variation”; Saratov, 1920; 2nd ed., rev. and enl., Moscow-Leningrad, 1935), also in English as “The Law of Homologous Series in Variation,” in Journal of Genetics, 12 , no. 1 (1922). 47–89; Polevye kultury yugo-vostoka (“Field Cultures of the Southeast”; Petrograd, 1922); “O geneticheskoy prirode ozimykh i yarovykh rasteny” (“On the Genetic Nature of Winter and Spring Plants”), in Izvestiya Saratovskogo selsko-khozyaistvennogo instituta, 1 no. 1 (1923), 17–41, written with E. S. Kuznetsova (with English abstract); “K poznaniyu myagkikh pshenits (sistematiko-geografichesky ocherk)” (“Toward an Understanding of Soft Wheats [a Systematic Geographical Essay]”), in Trudy po prikladnoi botanike, genetike i selektsii, 13 . no. 1 (1923). 140–257; “K filogenezu pshenits: Gibridologichesky analiz vida Triticum persicum Vav. i mezhduvidovaya gibridizatsia u pshenits” (“Toward a Phylogeny of Wheat: An Analysis of the Hybrids of Triticum persicum Vav. and Interspecific Hybridization in Wheat”), ibid., 15 , no. 1 (1925), 3–159, written with S. V. Yakushkina (with English abstract); “Ocherednye zadachi selskokhozyaystvennogo rastenie-vodstva (rastitelnye bogatstva zemli i ikh ispolzovanie)” (“The Next Tasks of Agricultural Plant Breeding [the Plant Riches of the Earth and Their Utilization]”), in Pravda (2 Aug. 1925), 5–6; Tsentry proiskhozhdenia kulturnykh rasteny (“Centers of Origin of Cultivated Plants”; Leningrad, 1926), with Russian and English texts; “Geograficheskie zakonomernosti v raspredelenii genov kulturnykh rasteny” (“Geographical Regularities in the Distribution of Genes in Cultivated Plants”), in Trudy po prikladnoi botanike, genetike i selektsii, 17 . no, 3 (1927), 411–428; “Geographische Genzentren unserer Kulturpflanzen,” in Verhandlungen des V. internationalen Kongresses für Vererbungswissenschaft, Berlin, 1927, which is supp. 1 of Zeitschrift für induktive Abstammungs- und Vererbungslehre (1928), 342–369; Zemledelchesky Afganistan (“Agricultural Afghanistan”; Leningrad, 1929), which is supp. 33 of Trudy poprikladnoi botanike, genetike i selektsii, written with D. D. Bukinich; and “Genetika” (“Genetics”), in Bolshaya sovetskaya entsiklopedia. XV (1929). 191–201.

Vavilov subsequently published “Gomologicheskie ryady” (“Homologous Series”), in Bolshaya sovetskaya entsiklopedia, XVII (1930), 586–587; Linneevsky vid kak sistema (“The Linnaean Species as a System”; Moscow-Leningrad, 1931): “Rol Tsentralnoy Azii v proiskhozhdenii kulturnykh rasteny (predvaritelnoe soobshchenie o rezultatakh ekspeditsii v Tsentralnuyu Aziyu v 1929 g.)” (“The Role of Central Asia in the Origin of Cultivated Plants [a Preliminary Communication on the Results of the Expedition to Central Asia in 1929]”), in Trudy po prikladnoi botanike, genetike i selektsii, 26 , no. 3 (1931), 3–44; “Meksika i Tsentralnaya Amerika kak osnovnoy tsentr proiskhozhdenia kulturnykh rasteny Novogo Sveta (predvaritelnoe soobshchenie o rezultatakh ekspeditsii v Severenuyu Ameriku v 1930 g.)” (“Mexico and Central America as a Fundamental Center of the Origin of Cultivated Plants of the New World [a Preliminary Communication on the Results of an Expedition to North America in 1930]”), ibid., 135– 199, with English and Russian texts; “Sovetskaya agronomia k XVI godovshchine Oktyabrya” (“Soviet Agronomy on the Sixteenth Anniversary of October”), in Izvestia (6 Nov. 1933), 3; Selektsia kak nauka (“Selection as a Science”; Moscow-Leningrad, 1934); Nauchnye osnovy selektsii pshenitsy (“The Scientific Basis of Wheat Selection”; Moscow-Leningrad, 1935); Uchenie ob immunitete rasteny k infektsionnym zabolevaniam (“Studies of the immunity of Plants to Infectious Diseases”; Moscow-Leningrad, 1935); Botaniko-geograficheskie osnovy selektsii (“The Phytogeographical Basis of Selection”; Moscow- Leningrad, 1935); and “Puti sovetskoy selektsii” (“The Courses Followed by Soviet Selection”), in Spornye voprosy genetiki i selektsii (“Disputed Questions of Genetics and Selection”; Moscow-Leningrad, 1937), 11–38, 462–473; and “The New Systematics of Cultivated Plants.” in Julian Huxley, ed.. The New Systematics (Oxford, 19401,549–566.

Vavilov’s posthumously published writings include The Origin, Variation, Immunity and Breeding of Cultivated Plants, which is Chronica botanica, 13, nos. 1–6 (1951); Mirovye resursy sortov khlebnykh zlakov, zernovykh bobovykh, Ina i ikh ispolzovanie v selektsii (“World Resources of Varieties of Cereal Grains, Legumes, Flax, and Their Use in Selection”; Moscow-Leningrad. 1957); izbrannye tmdy (“Selected Works”), 5 vols. (Moscow-Leningrad, 1959– 1965); “Zakony estestvennogo immuniteta rasteny k infektsionnym zabolevaniam” (“Laws of the Natural Immunity of Plants to Infectious Diseases”), in Izvestiia Akademii nauk SSSR. biol. ser. (1961). no. 1. 117–157; Pyat kontinentov (“Five Continents”; Moscow, 1962); Izbrannye sochinenia: Genetika i selektsia (“Selected Works: Genetics and Selection”; Moscow, 1966); and Izbrannye proizvedenia (“Selected Works”), edited and compiled by F. K. Bakhteev, 2 vols. (Leningrad, 1967), with a biography by P. M. Zhukovsky and a complete bibliography.

II. Secondary Literature. See B. L. Astaurov, “N. I. Vavilov i obshchestvo genetikov i seiektsionerov SSSR” (“Vavilov and the Soviet Society of Geneticists and Selectionists”), in N. I. Vavilov i selskokhozy aystvennaya nauka (“Vavilov and Agricultural Science”; Moscow, 1969). 84–89; F. K. Bakhteev. “Nikolaj Ivanovič Vavilov. Zu seinem 70. Geburtstag,” in TAG; internationale Zeitschrift für theoretische und angewandte Genetik, 28 , no. 4 (1958), 161–166, with portrait; T. Dobzhansky, “N. I. Vavilov, a Martyr of Genetics,” in Journal of Heredity, 38 , no. 8 (1947), 226–232, with portrait; N. P. Duhtnin. “N. I. Vavilov kak genetik” (“Vavilov as a Geneticist”), in Genetika, 4 . no. 3 (1968), 18–27; A. E. Gaissinovitch, “N. I. Vavilov, in Commemoration of the 25th Anniversary of His Death,” in Folia Mendeliana, no. 3 (1968), 55–58, with portrait; R. Ruggles Gates, “Vavilov and the Soviets,” in Science and Culture, 12 . no. 3 (1947), 423–427; S. C. Harland and C. D. Darlington, “Prof. N. I. Vavilov, For. Mem. R. S.,” in Nature, 156 (24 Nov, 1945), 621–622; David Joravsky, “The Vavilov Brothers,” in Stark Review, 24, no. 3 (1965), 381–394; and N. A. Maisurian, “Zhizn i deyatelnost N. I. Vavilova” (“Vavilov’s Life and Activity”), in Genetika, 4 , no. 3 (1968), 7– 17.

See also Paul C. Mangelsdorf, “Nikolai Ivanovich Vavilov,” in Genetics, 38 , no. 1 (1953), 1–4, with portrait; Zhores A. Medvedev, The Rise and Fall of T. D. Lysenko, 1. Michael Lerner. Irans. (New York- London, 1969), 17–19, 37–41. 52–77; Nikolay Ivanovah Vavilov (Moscow, 1967), with intro. by A. A. Fedorov and complete bibliography compiled by R. I. Goryacheva and L. M. Zhukova; M. M. Novikov, “N. I. Vavilov i genetika” (“Vavilov and Genetics”), in Velikani Rossyskogo estestvoznania (“Giants of Russian Natural Science”; Frankfürt, 1960), 141–150; A. I. Revenkova. Nikolay Ivanovich Vavilov (Moscow, 1962); Ryadom s N. I. Vavilovym: Shornik vospominany (“Side by Side With Vavilov: A Collection of Reminiscences”; Moscow, 1963), compiled by his son, Y. N. Vavilov; and “Uchen-ye-genetiki o Nikolae Ivanoviche Vavilove” (“Genetic Scientists on Vavilov”), in Genetika, 4. no, 3 (1968), 49–57.

Mark B. Adams

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