JOURNAL OF NEMATOLOGY VOLUME 9 JANUARY 1977 NUMBER 1

Historical Development of Nematology in Russia~ S. G. MJUGE 2

Abstract: The development of Russian hematology is considered from the late nineteenth century to 1970. The dominant influences of I. N. Filipjev and A. A. Paramonov are discussed in the context of the persons whom they influenced and their ctmceptual approach to the problems posed hy nematodes. The advantages and disadvantages of the framework of Russian scientific administration are compared to those in the West. Key Words: Filipjev, Heterodera spp., Meloidogyne spp., Paratnonov.

Science develops through the gradual popular agricultural journals such as accumulation of information and often Khozyain (The Landlord). Sometimes these changes direction with the development of consisted of complete translations of new ideas, or new approaches to old con- foreign articles, such as one by J. Kuhn cepts. In Russian nematology, conceptual (il). landmarks and scientific leadership were Also during this period, a considerable provided by I. N. Filipjev and A. A. amount of descriptive material was pub- Paramonov. The development of nema- lished outside of Russia. Bastian (I) tried tology in Russia is discussed with respect to summarize this information in a to the contrihutions of these great men. utonograph, an objective furthered by De Period before Filip]ev: Russian nema- Man (12). However, little synthesis or tology has always lagged behind that in the analysis was attempted in these monographs. West; it began later and in many technical After Charles Darwin's ideas began to gain aspects is still behind. The earliest pub- acceptance in biology on the eve of the 20th lished descriptions of nematodes go back Century, scientific thought required more only to the last decades of the last century. than descriptions. The nematodes that were first described 1. N. Filipiev period: The first part of were those visible to the naked eye or Filipjev's work was published in 1918, and those which caused striking pathological this was followed in 1921 by a second major changes in plants. For instance, in examin- contribution tinder the modest title of Free- ing grape vine roots for the presence of living Marine Nematodes in the Vicinity Philloxera, the Crimean Philloxera Com- of SebastopoI (4). In this paper, the author mittee (10) found root-knot nematodes. described 100 new species. If he had limited Similarly, in different areas, Heterodera was himself, as was usual at that time, to their found on beets by Tolpyguin (28), by description, or had even tried to classify Veinberg (30) in the Ukraine, and by and group them upon some morphological Tarniani (27) in Georgia. criteria, it is doubtful that his work would This was an age of the collection and have met with the enormous success that description of nature attd it lacked a unify- it did. Filipjev took a great conceptual ing conceptual framework. There were step by causally relating form and func- occasional zoological studies, such as that tion. In order to analyze the marine of Golovin (8) who examined the excretory nematodes that he had collected, he drew system, and articles began to appear in some upon the available literature on free-living, soil, microbivorous, and plant-parasitic Received for publication 29 July 1976. nematodes. i Invitational review. :lnstitute of Parasitology, Macdonald College of McGill He approached morphology on the basis University, Macdonald College, Province of Quebec, H0A of the total organism and not, as previous 1C0, Canada. I am grateful to Dr. Katherine Berdnikoff for essential help in translation and to Dr. Nell A. Croll, publications had done, on the basis of Director of the Institute of Parasitology, for providing separate organs. Thus he created the first me with laboratory facilities, eneouragemetn, and help with this manuscript. logical and comprehensive system of nema-

The JOURNAL OF NEMATOLOGY for October (8: 277-358) was issued 3 November 1976. 2 Journal of Nematology, Volume 9, No. 1, .January 1977

FIG. 1-4. Early Russian biologisls. 1) Aleksei Nikolaevich Severtzev (1886-1936). 2) Ivan Nikolaevich Filipjev (1889-1940). 3) Ivan Ivanovich Schmalhausen (1884-1963). 4)Aleksander Alexsandrovich Paramonov (1891-1.~70). Russian Nematology: Mjuge 3

FIG. 5. Group of Nematologists at the USSR Academy of Science. Back row (left to right): K. I. Schpa- kovskay, A. A. Paramonov, E. S. Turlygina, T. V. Pioroyskay; Front row (left to right): T. A. Gluzhenko, N. I. Sumenkova. S. L. Lazarevskaya-Blinova, P. S. Krulov, S. G. Mjuge. todes which was based on scientific data. "an enemy of tile people." His published The translation of his work into foreign works were removed from many libraries languages drew considerable international and destroyed. According to official sources, attention (21). The previous unnatural I. N. Filipjev died in a concentration camp systems of Cobb (3) and Micoletzky (13) ill 1941; according to Kirjianova (9), he were overtaken. Micoletzky (14) renounced died oll October 22, 1940. Filipjev was his system of nematode classification and posthumously "rehabilitated" during the accepted that proposed by Filipjev. Khrushchev regime. In 1934, Filipjev enlarged and deep- The period between Filipjev and ened his ideas (5), and in the same year Paramonov: New ideas arose in Russian published a monumental volume--Nema- biology in the "post-Filipjev" era. A. N. todes that are Harrn[ul and Use[ul in Severtzev (24) found new methods of Agriculture (6). This publication became a relating form and function to habitat. basic text for Russian nematology and was Today, he would be called an ecologist. widely read in other countries. In 1941, Proceeding from the ideas of Darwin, he this work was reprinted in an enlarged laid a new foundation for evolutionary edition with I. H. Schuurmans-Stekhoven morphology which was a definite advance- (7), but Filipjev himself did not participate ment over the descriptive morphological in the final production. In the West, approach of Karl Gegenbaur (1826-1903) Filipjev was "reported as missing," while and tile comparative morphology of Ernst his Soviet colleagues were told that he was Haeckel. Important conceptual advances 4 Journal of Nematology, Volume 9, No. 1, January 1977 were proposed by I. I. Schmalhausen (25) years of age, he skied actively, rubbed who analyzed the mechanisms of evolution. himself with snow, and swam in ice holes! The theories of "prototypes" and "archi- He knew all that was then known about techtonics" were developed in 1944 by V. N. H. schachtii. He trained a few specialists in Beklemishev (2) who foresaw numerical pest control but left no "heir" to his area of taxonomy and phylogeny within the hematology, with the possible exception of higher taxa. B. I. Kulchitsky. Apparently, one nematode By 1936, nematology had been influ- species alone was too little to occupy two enced by the conceptual advances of generations of nematologists. evolutionary theory and by related develop- Another "species" of nematode, the ments in general biology. A new applied root-knot group, became the main object of science of helminthology had been defined study in the entire life of A. A. Ustinov of as "the study of worms and the diseases the University of Kharkov. I say "species," that they cause" (26) with its subdivision since Ustinov insisted to his dying day that "ptlytohelminthology" which concerned there was only one species of root-knot plant-parasitic nematodes and diseases nematode: Meloidogyne marioni (Cornu). caused by them. In contrast to I. I. Korab, however, Ustinov Following the demise of Filipjev in had much wider interests than just the Russian nematology, a large collection of systematics of root-knot nematodes. He also nematode specimens remained in the studied physiology and ecology. As a result Zoological Institute of the Academy of of his leadership, there remained after Sciences of the USSR. This collection was Ustinov's death a strong and versatile preserved and enlarged by Filipjev's group of plant nematologists: N. M. assistant, E. S. Kirjianova (9). She picked Ladygina (ecologist), V. G. Zinoviev up the baton from Filipjev and enriched (biochemist-physiologist), and Z. G. Volod- Russian nematology through her varied chenko (morphologist). research contributions and through training of additional nematologists. Among her In the 1930s, there was a drive to students are E. L. Krall, T. S. Ivanova, develop the rubber-bearing plant, Scorzonea A. T. Tulaganov, and many others; these iau-saghyz, because natural rubber was not are the "grandchildren" of Filipjev. available in Russia and the technology to Professor Tulaganov held the posts of synthesize rubber was not then known. Professor of Invertebrates in the Depart- The growth of S. tau-saghyz tubers was ment of Zoology at the University of often hampered by disease and nematodes. Tashkent; Rector of the University of As a result, the first specialized laboratory Samarkand; and Director, Institute of in phytonematology was opened in Moscow Parasitology, Uzbek, S.S.R. He trained in 1933 under the Directorship of N. M. many nematologists who worked on applied Sveshnikova. Sveshnikova soon became the problems but published little. Possibly leading authority on the chemical control even great-great grandchildren of Filipjev of soil nematodes in the Soviet Union. She can be derived from this tree of nema- trained L. A. Guskova, widely known in tological development. The main direction U.S.A., who is now Head of the Department of the Tulaganov school is systematics anti of Phytohelminthology of the All Union faunistic studies. Institute for the Protection of Plants in Soviet nematology has not been derived Leningrad. T. S. Skarbilovitch succeeded entirely from Filipjev and his school. In Sveshnikova in the tau-saghyz laboratory the Ukraine, in the early 1920s, I. I. Korab and later devoted much time to the study began working in the Agricultural Institute of the biology of Heterodera, especially of Bielaya Tserkov (White Church). He was H. schachtii. concerned with Heterodera schachtii since In the early 1920s, the zoologist N. M. this was the center of the sugar beet Ktdagin developed an interest in nema- industry in Russia. As a novice nematol- todes. It was a small part of his professional ogist, I made a pilgrimage to see him. As career and he produced only two articles in the oldest of our Soviet colleagues, he 1927 and 1928. His real contribution was astonished me with his youthfulness. At 70 to acquaint one of his students, A. A. Russian Nematology: Mjuge 5 Paramonov, with nematodes and nema- of this work, lie conducted research into tology. the specificity of plant parasites with A. A. Paramonov: Paramonov became respect to agricultural practice in Russia Professor of Darwinism in 1937 and (18). On the basis of earlier studies by Professor of Zoology in 1941 at the Agri- Severtzev, Schmalhausen, and Beklemis- cultural Academy of K. I. Timiriazeff in chev, he produced a cohesive theory of the Moscow. He held both these appointments phylogeny of nematodes (19, 20). The until 1948. Paramonov received a sound breadth of his vision led to the idea that basic background in biology under Pro- nematodes were to be considered as com- fessor O. Butschli at the University of ponents in the natural "ecosystem" and Heidelberg. He was also influenced by the that their presence was to be viewed as a leading professors of that period in Moscow consequence of the environmental condi- and St. Petersburg. Paramonov was first a tions. This view enabled correlations to be theoretician, but because of his vocation drawn between nematode populations and and position within an agricultural acad- the habitats of their plant hosts. At its best, emy, he made contributions to applied this approach could be used to predict agriculture. outbreaks of disease which related to agri- Although Paramonov began to be cultural practices and climatic conditions. interested in nematodes in 1925, his main Together with his co-workers, Paramonov interests until 1948 had been in general moved towards a centralized theory for the biology. He was the author or co-author of "therapy of phytohelminthoses." almost all of the more popular textbooks He presented his theoretical principles and books published in the U.S.S.R. on the in a three-volume work: The Basic Prin- subjects of Darwinism and zoology. Thes~ ciples of PhytoheIminthology (21, 22, 23). included encyclopedias and the Soviet All of these volumes were translated edition of Bremm's Life of the Animals. into English. Unfortunately, Paramonov Because of the politicizing of Soviet biology did not complete the last of his planned in 1948, Paramonov was forced to abandon volumes in which he intended to describe the university chairs that he occupied and the theory of controlling "phytohelminth- he turned exclusively to helminthology. oses." He died on June ll, 1970, and left Paramonov applied for his retirement the most radiant memory in all who were pension in 1970 but did not expect to have privileged to know him. his application accepted. He wept when, as Despite his important position and the an economy measure, his retirement was fact that he was a man of great principle, agreed upon, and he died one month later. he was a remarkably kind, gentle, and From 1952 until his death in 1970, he was sympathetic colleague. He lived in a com- Head of the Section of Phytohelminthology munal apartment and occupied one room of the Hehninthological Laboratory of the which was divided into three "cubby-holes" Academy of Sciences of the USSR. by plywood walls. In addition to normal Paramonov viewed phytohelminthology as inhabitants, there were always some a complex discipline in which the compre- wounded pigeons, stray kittens, or puppies. hensive study of nematodes must merge One could always come to him with any-- harmoniously with questions of plant even the most trivial-questions, or without pathology. He soon assembled a group of any question at all. Any time of the day or phytohelminthologists interested in all night, one could count on a warm welcome aspects of nematology. and we, his students, all too often took The first ecological classification of advantage of this. nematodes was published by Paramonov The study of nematodes in Russia: The (17) in 1952. In it he analyzed the relation- works of Filipjev and Paramonov have left ships that existed between plants and a lasting impression on Russian nematology. animals. This system was an outgrowth of The first instructions for collecting nema- the system of Filipjev and, as an existing todes and examining diseased plants were natural system, it has stood the test of those of Filipjev (1932). Filipjev (4) became time. In 1962, Paramonov (21) enlarged so convinced of the ubiquity and ecological the system further. As a peripheral aspect specialization of nematodes that, describing 6 Journal of Nematology, Volume 9, No. 1, January 1977 nematodes of the Petrograd area in 1921, proteolytic enzymes was found in all the he described species which, in his opinion, species investigated. The activity of this should have existed there. The species latter enzyme was linked to the presence were, as it happens, discovered th_r~" at a of specific hydrogen donors that activated later date! it and played an important role in the Since every faunist strives to discover respiration of the host. By influencing the new species, most Russian researchers did respiratory metabolism of hosts, it became the same and concentrated on the "exotic possible to inhibit the proteolytic activity areas" such as Armenia, Azerbaijian, White of the nematodes. This reaction reduced Russia, and Central Asia. Meanwhile, their nutrition and their sexual reproduc- nematodes in the central part of Russia tion (16). remained completely unknown (20). The goal orientation of the research In contrast, Paramonov felt that the clone by all members of Paramonov's group study of playtohelminthoses should be gradually became side-tracked into indi- comprehensive. He wished to proceed by vidual projects. Up to a point, the qtmlities developing a total description of the fauna, of Paramonov played a negative role, for not only by a one-time, widespread he understood the individual creative sampling but by regular sampling of the aspirations of tile members of his group same habitat. Then, the various factors and permitted them to develop. The real promoting the increase or decrease of disintegration of the central theme came pathogenic species should be elucidated; about by the destructive nature of Soviet and ecologists should study the inter- science. relationships of nematodes amongst each Contemporary factors in Russian nema- other and with fungi, insects, bacteria, and tology: It may be seen that Russian other organisms. Therapy, Paramonov nematology is more classical and concep- believed, should follow the studies of tual, even in its approach to control, than physiologists and biochemists who would the more pragmatic approach of Western establish the factors that could decrease science or nematology. Recent develop- populations. This approach was the essence ments have not favored entrepreneurial of applied Russian nematology. approaches. This situation is related both The development of therapy for to the security and confidentiality of some phytohelminthoses: Paramonov organized a government laboratories and to the par- group of physiologists whose role it was to ticipation of the USSR in the International find "nematostatic" substances (E. S. Patent System. Turlygina) and to study the physiology of The Academy of Science is officially nematode nutrition (S. G. Mjuge). Turly- totally independent in its scientific affairs. gina (29) showed in vitro that the sexual It is there that theoretical problems are reproduction of Rhabditis spp. was in- tackled and the results are ultimately hibited by potassium rhodanite, ammonium transmitted to the appropriate government nitrate, and sodium salicilic acid. The same ministries. From a practical point of view, substances inhibit the laying of eggs by control of science in USSR is in the hands Meloidogyne incognita if plant hosts are of the Central Committee of the Com- treated. munist Party of the Soviet Union. Within S. G. Mjuge determined that some plant the committee, there is a "Department of nematodes possess extra-corporeal enzymes Science" which directs scientific affairs, for digestion of host tissues. These secre- including applied problems. For instance tions were considered a target for destroying in 1965, the Central Committee was nematodes. In order to provide a firm basis persuaded that the spread of cotton wilt for understanding extra-corporeal secre- was caused by nematodes. As a result, the tions, and in the traditions of Filipjev and Academy of Sciences received the suggestion Paramonov, it was decided to examine the that all helminthologists should become evolutionary pattern of secretory function involved in the solution of this problem. (15). Mjuge demonstrated tlmt the catalytic I. I. Skrijabin had to display enormous enzymes for carbohydrates varied among political talents to avoid total commitment species, but that a cathepsin-type of of himself and other members of his group Russian Nematology: Mjuge 7 to this assignment. It later transpired that nematologists have been conceptual ad- nematodes did not play an important role vances built around the framework of I. N. in the spread of Fusarium and Verticillium Filipjev and A. A. Paramonov. Their ideas cotton wilt. There are other parallel have continued into the contemporary era situations in most Communist cotmtries. where theoretical advances, not practical Because of these restrictions, the Central discoveries, are the politically safe ones. Committee usually develops "safe" themes If I have succeeded in showing the pre- of broad agricultural significance. Practical eminent position of my old friend and goals are avoided because this is the only teacher A. A. Paramonov, then the aim of mechanism for scientific advancement. By this article will have been achieved. This chance, I have preserved a copy of my review does not discuss developments after "Examiner's Report" from my doctoral 1972, but nothing I have heard makes me dissertation. The first words are: "There is feel that there have been significant changes no data or information that could be the in political or scientific direction since that subject of an invention or a discovery in date. this work." This clause appeared in the examiner's reports after the USSR joined LITERATURE CITED the International Patent Association. Prior !. BASTIAN, H. C. 1866. On the anatomy and to that date, proven foreign nematicides physiology of the nematodes, parasitic and were analyzed and similar componnds re- free; with observations on their zoological synthesized in the Institute of Y. V. position and alfinities to the Echinoderms. Samoilov in Moscow. In this way, the Phil trans. Roy. Soc. Lond., 156:545-638. 2. BEKLEMISHEV, V. N. 1944. Osnovy sravnitelnoi Soviet preparations No. 23 (analogue of anatomii bespozvonochnykh (Fundamentals farbiate) and No. 93 (analogue of DD) of comparative anatomy of invertebrates). were created. But in the 1950s, the situation 420 p. changed. The authorities began to fear 3. COBB, N. A. 1919. The orders and classes of nemas. Contributions to the Science of that Soviet discoveries would be exploited Nematology 8:214-216. abroad, and it was forbidden to publish 4. FILIPJEV. I. N. 1921. Free-living marine nonpatented works that were of any nematodes in the vicinity of Sebastopol. Part practical value. Since patents can take 2 I. Trudy Osoh. Zool. Lab. Sevastopol, Biol. years, and authors advance partly through Stantsii Ross. Akad. Nauk. 2:1-350. 5. FILIPJEV, I. N. 1934a. The classification of the publications, they often argue that their free-living nematodes and their relation to work is of no value. Should an author the parasitic nematodes. Smithson. Misc. deceive the Commission, he will be cruelly Colin. 89:1-63. punished. The reader must be aware of this 6. FILIPJEV, 1. N. 1934b. Nematody vrednye i problem in assessing the nature of nema- poleznye v. selskom -- knozyaistve (Nematodes that are harmful and useful in agriculture). tological work currently being developed in Moscow-I,enit~grad. Sel. hoz Giz, 1-440. Russia. 7. FILIPJ EV, I. N., and SCHUURMANS- There are "closed" phytohelmintholog- STEKHOVEN, J. H. 1941. A manual of ical laboratories in the USSR, perhaps agricultural helminthology. E. J. Brill, Leiden. 888 p. because Karl Marx connected the first 8. GOLOVIN, E. P. 1902. Nabludeniya had nema- revolutionary movement in Europe with a todami. Exskretornyi apparat. (Observations potato famine. Selected nematologists, on Nematodes. Excretory apparatus.) Kazan. earning higher salaries, are actively working 120 p. on secret subjects in the Ministry of War. 9. KIRJIANOVA, E. S. 1959. K. 70-Letiju so dnia rozhdenia vydaiushchegosia sovetskogo It is difficult to find out what they are oucbienogo professora. Ivan Nikolaevich doing, but secrets do leak out. Heterodera Filipjev. The 70th anniversary of the birthday rostochiensis and .4phelenchoides besseyi of the outstanding Soviet scientist, Professor are most certainly considered as tools of Ivan Nikolaevich Filipjev. Izv. Akad. Nauk biological warfare! tadzhik. SSSR. 2:51-155. 16. KOMITET, K. F. 1890. Otchet za 1889 god. Conclusion: We see that nematology (Crimean Philloxera Association. Report for began to develop in Russia shortly after it 1889.) Yalta, 1-19. did in the West, and that it has gained 11. KUHN, J. 1889. Sposob unichtozheniia sveklo- vichnych nematodov. (Method of destroying much from its counterparts overseas. The the beet nematodes. From foreign literature.) international contributions of the Russian Sel Khoz. kazakhst. 161:58.68. 8 Journal of Nematology, Volume 9, No. 1, January 1977

12. MAN, DE J. G. 1884. Die frei in der reinen Geological and morphological characteristics Erde und im sussen Wasser lebenden Nema- of Phytonematodes. General principles of toden der niederlandischen Fauna. Leiden, taxonomy. Izx. Akad. Nauk. SSSR. 480 p. 206 p. 22. I'ARAMONOV, A. A. 1964. Osnovy phyto- 13. MICOLETZKY, H. 1922. Die freilebenden Erd- gelmintologii. II. (Basic Principles of Phyto- Nematoden mit besonderer Berucksichtigung nematology.) Taxonomy of Phytonematodes. der Steiermark und der Bukowina, zugleich Izv. Akad. Nauk. SSSR. 466 p. mit einer Revision samtlicher nicht mariner, 23. I'ARAMONOV, A. A. 1970. Osnovy phyto- freilebender Nematoden in Form yon Genus- gehnintologii. III. (Basic Principles of Phyto- Beschreibungen nnd Bestimmungs-schlusseln. hematology. Taxonomy of Phytonematodes.) Arch. Natnrgesch. (1921) 87 Abt(*): 1-320; (9): Izv. Akad. Nauk. SSSR. 254 p. 321-6.50. 24. SEVERTZEV, A. N. 1939. Morphologicheskie 14. MICOLETZKY, H. 1925. Die freilebenden, zakonomernosti Etolyutsii. (Morphological subwasser und Moor-Nematoden Danemarks. patterns of evolution.) Moscow-lzv. Akad. Mere. Acad. Rny. Soc. et lettres 1)anemark Nauk. SSSR. 591 p. Sect. sci. 8-me set, X, 2:1-256. 25. SCHMALHAUSEN, 1. I. 1939. Puti i zako- 15. MJUGE, S. G. 1956. Sravnitelno-physiologiches- nomernosti evolutsionogo protsessa. (Paths kaya kharakteristika phytonematode. (Com- and patterns of evolution.) Moscow-Leningrad parative physiological characteristics of Izv. Akad. Nauk SSSR. 231 p. phytonematodes.) Trudy nauchnoj, konj. 26. SKRIJABIN, K. 1. 1946. Stroitelstvo sovietskni parazitologov USSR, Kiev. gelminthologii. /The development of Soviet 16. MJUGE, S. G. 1965. Physiologia pitaniya helminthology.) Moscow-l.eningrad. Izv. Akad. phytonematod i problemy terapii gelmintozov Nauk SSSR. 221 p. rastenii. (Physiology of phytonematode feed- 27. TARNIANI, I. 1897. Sveklovichmaia nematoda ing and problems of therapy of plant (Heterodera schachtii Schmidt). Nasekc~mye helminthoses.) Akad. Nauk. SSSR. Moscow. i drugii zhivotnye nanosiaschchie vred v 17. PARAMONOV, A. A. 1952. Opyt ekologicheskoi selskom khoziaistve. (The beet nematode, klassifikatsii phytonematod. (On the ecological Heterodera schachtii Schmidt, in "Insects classification of plant nematodes.) Trndy and other animals that are harmful to gel'mint. Lab. 6:338-369. agriculture"). St. Petersburg. 62 p. 18. PARAMONOV, A. A. 1954. Spetsifischnost 28. TOLPYGUIN, M. 1880. Istriblenie sveklovich- phytogelmiutov i. ee znachenie v selskokhoz- nykh-vskhodov mikroskopicheskimi -zhivtnymi. yaistvennoi praktike. (Specificity of phyto- (The destruction of beet seedlings by micro- helminths and its importance in agriculture.) scopic animals.) Zap. Kievkogo, otd. Imper. Zool. Zh, 33:1002-1024. russk, techn, obsch, po sveklosakharnoi 19. PARAMONOV, A. A. 1956. K. revizii sistemy promysh. 10:238-240. Rhabditat rastenii. (Revision of the classifica- 29. TURLYG1NA, E. S. 1958. Voprosy teorii terapii tion of Rhabditids of plants.) 8:85-111. rastenii zakrytogo grounta pri gallovom 20. PARAMONOV, A. A. 1958. Glawlye hapravlenija nematodoze. (Questions of the theory of the evolutsii phytonematod otryada rhabditid i therapy of root-knot infestations of plants tylankhid. (Main trends in the evolution of in pot-like systems). Sborn. rabot molodykh phytonematodes of the orders Rhabditida and phytogelmintologov, lzv. Akad. Nauk SSSR. Tylenchida.) Zool. Zh. 37:736-749. 82-98. 21. PARAMONOV, A. A. 1962. Osnovy phyto- 30. VEINBERG, G. 1891. Odin iz parazitov sakharnoi gelmintologii. I. (Basic Principles of Phyto- svekly. (One of the parasites of sugar beet.) helminthology.) 1. The origin of nematodes. Sel'. Khoz. kazakhst. 27:28.