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Genetic Resources of Leguminous Plants in the N.I. Vavilov Institute of Plant Industry

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Genetic Resources of Leguminous Plants in the N.I. Vavilov Institute of Plant Industry
  Genetic Resources of Leguminous Plants in the N.I. Vavilov Institute of Plant Industry     In the centre of St.-Petersburg in St. Isaac ’s Square ( B. Morskaya str. 42-44) two buildings constructed by architect Yefimov are situated. They house the N.I. Vavilov Institute of Plant Industry. There is a memorial board on one of these buildings with the following words carved: « Nikolai Ivanovich VAVILOV, a prominent biologist, worked here in 1921-1940». Nikolai I. Vavilov (1887  – 1943) is recognized as the foremost plant geographer, botanist and geneticist of contemporary times. In the early 20th century, the world did not appreciate the urgency of protecting the environment and scientists thought little about the gradual disappearance of valuable plant species. Vavilov was among the first to recognize the need for intensive plant collecting, research and preservation. He organized and took part in over 100 collecting missions in the major agricultural areas of  the world. During these expeditions, Vavilov paid special attention to leguminous crops as sources of protein and as means of increasing soil fertility (Sinskaja 1991). He considered these issues to be of the highest priority in biological and agricultural sciences for developing sustainable agricultural production. Many close colleagues of Vavilov e.g. Barulina (his second wife), Govorov, Zhukovsky, Sinskaja further developed his ideas, selecting legumes as the subject of their research. Govorov, L.I.   (1885-1941)   Prof. Leonid I. Govorov was the Russian agronomist and plant breeder; from 1915 worked at Moskow Breeding Station; from 1923 was the Head of the Steppe Station, and later was the first Head of the Department of Leguminous Crops at VIR. Vavilov and his colleagues revealed a number of important findings in plant biology through such research. However, this vast body of work has not been brought together in a single report. Vavilov’s expeditions and their outcomes   Vavilov was particularly interested in the sites of ancient agricultural civilizations and of mountainous regions. In 1916, he was sent to Iran and Pamir by the Ministry of Agriculture to determine the reasons for disease epidemics among the resident Russian garrisons. Vavilov discovered that the wheat used for bread flour was contaminated with seeds of the poisonous grass species Lolium temulentum . Vavilov continued his travels in the northern and central regions of Iran, in Pamir and other regions of Central  Asia. His main purpose was to collect early varieties of agricultural plants for testing in northern and droughty areas of Russia, and to determine the high-altitude limits of agriculture (Vavilov 1987a, 1991). He was also searching for "Persian wheat”, reported to be resistant to many diseases (Bazilevskaya and Bakhareva 1991).   Despite a careful search, Vavilov did not find "Persian wheat" in Iran. However, he did collect many other valuable accessions, among them leguminous crops including mung bean , chickpea , lentil , everlasting pea , pea , beans  and species of clover  , not known at that time in Russia. The materials collected by Vavilov in Iran and in Pamir formed the foundation of the collection of leguminous grain crops of the VIR.  As a result of his first expedition, 171 grain legumes samples were collected. Subsequent collecting missions of L.I. Govorov, N.R.Ivanov, P.M. Zykovsky, B. S. Kurlovich increased the number of legume accessions from Central Asia to 1373. The results of a detailed investigation of these were a starting-  point for developing a number of important theoretical and practical generalizations on the srcin, geography, genetics, disease resistance and evolution of cultivated plants. Vavilov’s  travels in Iran indicated that there were sources of ancient agriculture in southwest Asia. He also detected particular ecotypes in Pamir, which suggested their srcin was in those mountains. He concluded that southwestern Asia was a centre of srcin of a many legume species (Vavilov 1965b). The majority of the legumes samples from this centre were small-seeded with specific flower colours determined by recessive genes (Zhukovsky 1971). Cross-pollinated species from southwest Asia (everlasting pea, fodder beans) have a propensity for self-fertilization. In the law of homologous series (Vavilov 1920), which was largely formulated following the expedition to Iran and Pamir, Vavilov noted that variability characterized the entire Leguminosae (Vavilov 1920, 1987b). E.I. Barulina established a precise parallelism in variability of vetch and lentil samples that srcinated from Iran (Barulina 1930). Their similarity is such that even an expert finds it difficult to distinguish between seeds of these species. N.I. Vavilov  with his wife E.N. Barulina , 1926 (before their expedition to the Mediterranean). E.N. Barulina was the leading scientist in the field of genetic resources of Leguminous crops. This example has become a classic illustration of Vavilov’s l aw of homologous series in hereditary variability (Sinskaja 1969; Makasheva 1973, 1979; Vavilov 1987a, 1987b). Forms of legumes with various seed and flower colours, obeying the law of homologous series, are demonstrated by many species of lupin, peas, mung bean, string bean and fodder beans. In 1924, Vavilov organized an expedition to  Afghanistan together with V.N. Lebedev and D.D. Bukinich (to Herat, Afghan Turkestan, Gaimag, Bamian, Hindu Kush, Ba Kafiristan, Jalalabad, Kabul, Kandahar, Baquia, Helmang, Farakh and Sehistan). The findings from this expedition supported the conclusions about southwest Asia as a centre of srcin of many plants. It was established that Afghanistan was a major primary focus of formation where there existed a large diversity of many major Eurasian crops, representing an inexhaustible source of initial material for selection (especially for drought resistance). Following this hazardous and very fruitful expedition to Afghanistan, Vavilov was awarded the N.M. Przevalski Gold Medal of the Russian Geographic Society, of which Vavilov was president from 1931 to 1940. Vavilov had a special interest in the Khoresm oasis, whose proximity to Afghanistan and Iran supported the hypothesis that this territory was also a focus of formation. Vavilov inspected area around the Amu Darya River (Khiva, Urgench, Gurlen and Tashauz) in 1925. Whereas there were signs in Iran and  Afghanistan that cultivated plants had developed locally, Khoresm showed signs of connections with northeast Africa and Egypt that influenced the features of cultivated plants. White-seeded forms of peas, haricot bean and fodder bean were found in Khoresm, together with particular forms of groundnuts and  alfalfa. Khoresm oasis was characterized by an abundance of many recessive forms of cultivated plants (Vavilov 1926; Bazilevskaya and Bakhareva 1991). In 1921 Vavilov organized a trip to Canada (Ontario) and the USA. The official purpose was to search for sources of resistance to drought, necessary for restoring Russian agriculture after a severe drought in 1921. Vavilov understood from his first visit to North America that the continent was not a focus of intensive formation of plants and agriculture (Vavilov 1965b, 1987b). The main foci of plant diversity were to the south, in southern Mexico, and Central and South America. Vavilov visited a number of European countries (Britain, France, Germany, Poland, Holland and Sweden) on his return from America in 1922.  As a result of these and his subsequent travels to Germany in 1927, the USA in 1930, Canada and other  American countries in 1932-33, he added commercial varieties and a selection of legumes from many countries to the collection of VIR. Vavilov visited Mediterranean countries in 1926: northern Africa, the islands of Cyprus, Crete, Sicily and Sardinia, and southern Europe (Portugal, Spain, France and Greece). He noted the role of leguminous crops (particularly chickpea) in supplying both people and livestock with protein and as a means of increasing soil fertility (especially lupin) (Vavilov 1997; Loskutov 1999). He found that the activities of people had little effect on plant diversity in any of the coastal zones of the Mediterranean. However, a different picture emerged in inland areas, oases and southern slopes of mountains, where the influence of European civilization was apparent (Vavilov 1997). Vavilov’s special interest in early agrarian cultures led to an expedition to Syria, Palestine and Jordan, where there were traces of ancient agriculture. In Palestine in particular, he found useful forms of white lupins (going under the local name Tel Karam). These are distinguished by early maturity and are now widely used in breeding (Golovchenko et al. 1984). Plant collection in Sudan and Ethiopia yielded about 2000 accessions of local varieties. In Sudan, Vavilov found valuable forms of white lupin, and in Ethiopia found endemic forms of everlasting pea, chickpea, lentil and beans. The flora of Ethiopia is in many respects unique and Vavilov considered it to be an independent primary Abyssinian focus (Vavilov 1926), and later the Abyssinian Centre of diversity (Vavilov,1962). In 1929 Vavilov organized an expedition to China (Xinjiang  – Kashgar, Uch Turfan, Aksu, Kucha, Urumchi, Kulja and Hotan), Japan (Honshu, Kyushu and Hokkaido) and Korea. The purpose of his trip to northern and western China was to determine the role of Central Asia (within the limits of China) in the srcin of cultivated plants of eastern Asia. Returning from northern and western China, he inspected the area close to the Chinese border around Lake Issyk-Kul and the Syr Darya river basin. Vavilov concluded from the results of this trip that Central Asia does not have the features of an independent focus of ancient agriculture: all that was cultivated there had been introduced from the east. His investigations in Japan, however, confirmed it as one of the centres of cultivated plants (the East Asiatic centre). He found that, while Japan had ‘borrowed’ some cultivated plants from China, its geographical isolation, its span in latitude, and its diversity of climate and ecology generated unique features in the cultivated flora. Wild species and cultivated forms of soybean were added to the collection following these expeditions. Vavilov was interested for a long time in plant genetic resources of Latin America. In 1932, an invitation to the VI International Genetic Congress in the USA gave him an opportunity to inspect these regions. Vavilov visited Cuba, Mexico, Ecuador, Peru, Bolivia, Chile, Brazil, Argentina and Uruguay after the congress. Vavilov detected specific and varietal structures in cultivated plants, and received information about their history, srcin and wild relatives. He collected valuable forms of kidney bean, peanuts and  American species of lupin. Thi s was Vavilov’s last overseas trip.  Vavilov also traveled widely in the territory of the former Soviet Union. He paid special attention to the mountain regions of the Caucasus and Central Asia, in which there were unique cultures and wild species, whose role in the creation of cultivated plants is not doubted. The direct connection of the Caucasus with Iran, Central Asia and Turkmenistan has resulted in some similarities with the Southwest  Asiatic Centre of srcin of cultivated plants. Vavilov personally participated in expeditions to more than 50 countries. In 1938  –1940, he wrote essays on his journeys under the title "Five Continents”. The existing  pages of this manuscript were published in 1962 and 1987 in Russian (Vavilov 1962, 1987a), and later in English by IPGRI (Vavilov 1997). The characteristics of the principal legume accessions collected by Vavilov   Mung bean  ( Vigna radiata  (L.) R.Wilczek)   Vavilov collected 115 accessions of mung bean during his first expedition in 1916. Mung bean is an ancient food crop of southwest Asia. It has a high seed protein content (23.0  – 32.1%), high lysine, tryptophane and vitamin contents, and good flavour and cooks quickly (20  – 40 min). These properties have made it a favourite food of the local population. Its foliage is useful as forage for cattle and as a green manure.   Vavilov’s accessions of this crop were studied at the Central Asian branch of VIR (Popova 1937; Pavlova 1952; Ivanov 1961). From a detailed study of the assembled diversity of mung bean, G.M. Popova developed an intraspecific classification of this crop, differentiating three subspecies: Indian ( indicus ), Chinese ( chinensis ) and Iranian ( iranicus ). She also described 63 varieties of mung bean (Popova 1937). Subspecies iranicus are characterized by a twisted form of bush, a stalk height of 86-128 cm, 50-180 pods per plant (7-8 cm long), seeds of various colours (yellow, green, grey, brown) and 1000-seed weight of 32.8-49.8 g. Most accessions are early maturing, but late-maturing accessions also exist. The growing period in Central Asia is 91-122 days. The seed yield per plant ranges between 7.3 and 17.0 g. Two accessions from Afghanistan (k-2209, k-2216) are highly resistant to drought and are of particular interest. Accessions of Iranian srcin differ by having large above-ground biomass and are potentially useful for creating cultivars intended for fodder and green manure. Chickpea ( Cicer arietinum  L.)   This is an ancient crop of Iran and Central Asia and is represented by numerous local races with small seeds. Plants are small and suited only to manual harvesting. However, they are widely used in breeding. The variety ‘ Tadzhiksky 10’    was created by individual selection from Vavilov’s Tadzhikistan samples. It matures early and is res istant to fusarium wilt. This variety was used to breed the variety ‘ Zimistony  ’, which is also characterized by early maturity and high productivity in Tadzhikistan. ‘ Milutincky  ’ was developed from the Iranian sample k-327 by individual selection for Uzbekistan conditions. This cultivar is drought and disease resistant. ‘ Tashkent 511’   was created from Afghan sample k-223 through mass selection. It has enhanced protein content and disease resistance. ‘ Kubansky 16’   was developed for conditions of the Kuban region from Uzbek sample k- 16. ‘Volga - 5’ has shown itself highly productive in central Russia. It was bred using accession k-249 from Afghanistan. Lentil ( Lens esculenta  Moench)   Vavilov’s lentil accessions are mainly dwarf forms with small seeds belonging t o a large number of varieties: persica, grisea, violascens and others. There are many endemic forms of early and semi-early varieties that are drought resistant and have good cooking properties (Table 3). Pamir sample k-194 matures later than all others (growth period 86-102 days). Accessions k-5, k-196, k-434, k-435 have light-blue flowers. Material assembled by Vavilov was used as the basis for creating the commercial cultivar ‘Tadzhik 95’, which matures early, has good cooking properties and is drought resistant. ‘Azer’, now cultivated in Azerbaijan, was created using sample k-373 from northern Afghanistan. It is characteristically tall and has rhomboid beans. Its growth period is 81-95 days and it has a seed protein content of 29-30% and a 1000-seed weight of 34-37 g. Pea  ( Pisum sativum  L.)  
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