CC BY
17
Protistology 16 (2): 57-67 (2022) | doi:10.21685/1680-0826-2022-16-2-l Historical materials
Protistology
Protistology in the Zootomical Cabinet of the St. Petersburg University (Department of Invertebrate Zoology). People and their destinies
Sergei I. Fokin
Department of Biology, Unit of Protistology, Pisa University, Italy, and Department of Invertebrate Zoology, St. Petersburg State University, Russia
| Submitted February 12, 2022 | Accepted April 30, 2022 |
Summary
The article considers the evolution of protistological studies in the St. Petersburg University and primarily in the Zootomical Cabinet. This historical overview presents the examples of life stories and scientific achievements of the main participants of this process. Researchers belonging to different generations of zoologists, from the 1830s until the transformation of the Zootomical Cabinet into the Department of Invertebrate Zoology in 1930, are mentioned. The role of professors Schewiakoff and Dogiel in the formation of the scientific protistological school in St. Petersburg University is emphasized.
Key words: L.S. Cienkowsky, V.A. Dogiel, S.S. Kutorga, K.S. Mereschkowsky, protistological scientific school in St. Petersburg University, W.T. Schewiakoff, Zootomical Cabinet
Introduction
Protistology, which for a long time had been termed "protozoology", was considered a part of zoology until the middle of the XX century. The latter in Russian universities of the XIX century was divided into zootomy (zoology of invertebrate animals) and zoology itself, which included all studies of the vertebrate animals. The Zootomical Cabinet was separated from the Zoological one in the Imperial St. Petersburg University (ISPbU) in 1871. It is clear that speaking about protistological studies therein, it is necessary to give some retrospective overview of their evolution since the foundation of the University. Usually, quite many scientists in the
https://doi.org/10.21685/1680-0826-2022-16-2-1
© 2022 The Author(s)
Protistology © 2022 Protozoological Society Affiliated with RAS
XVIII through the first half of the XIX century were describing protists, first of all, ciliates, and other tiny organisms, unicellular or multicellular (flagellates, amoebae, rotifers, trematode larvae), under the title "infusoria" (see Ehrenberg, 1838; Kutorga, 1839). The group sometimes was considered as zoophytes. For instance, in the book of P.Ph. Gorjaninov (1796—1865) "Zoology and Zootomy used for the general benefit, especially for medicine", the author indicated that the kingdom of "animals-plants" or "zoophytes" corresponded to the common phylo-genetic root of all living organisms (Gorjaninov, 1837).
Later, E. Haeckel (1834-1913) introduced the concept of Protista or Protoctista for Protozoa to
Corresponding author: Sergei I. Fokin. Department ofBiology, Zoology-Anthropology Unit, University of Pisa, 56126 Pisa, Italy; sifokin@mail.ru
S.S. Kutorga. St. Petersburg, 1850s.
denote this group of living beings that connected plants and animals at the base ofhis first phylogene-tic tree (Haeckel, 1866). In fact, it was a conglomerate of evolutionarily different organisms, where fungi and bacteria (monera), most protists, as well as sponges were grouped together. At the same time, Haeckel initially included the infusoria in the Animalia as a part ofthe Articulata branch. Then, of course, this error, which was probably related to Ch. Ehrenberg's ideas, was corrected (Haeckel, 1879). For ciliated protists (the main part of the former infusoria group), the phylum Ciliophora Doflein, 1901 was established at the very beginning of the XX century.
In this article, I dwell primarily on the pre-Soviet period of the development of protistology at the St. Petersburg University. It is obvious that in the recent 100 years, the St. Petersburg school ofthis biological discipline has developed immensely. However, this is a rather huge field to discuss, and thus I am postponing this comprehensive discussion until future times. The previous short version of this long story was published two decades ago (Fokin, 2001a).
Pioneers of protistology at the St. Petersburg University
The study of the unicellular organisms began at ISPbU at the dawn of its existence — in the second part of the 1830s. Namely, S.S. Kutorga (1805-1861), a professor of zoology at the ISPbU, published his "Natural history of the development of Infusoria" (Kutorga, 1839). However, at that time it was just a brief episode related rather to the beginning of the broad use of a microscope in biological research and teaching. This was obviously the case partly inspired by the Ch. Ehrenberg's monograph, too (Fokin, 2001a). This remarkable monograph in two volumes was published a year earlier in Germany (Ehrenberg, 1838).
In Europe of that time, protistological views of Ch.G. Ehrenberg (1795-1876) predominated. Ehrenberg treated, for instance, ciliates as very small but perfectly organized animals with all the organ systems ofmulticellular animals. This approach was non-critically shared by S.S. Kutorga, who in his work of1839 actually gave a synopsis of Ehrenberg's famous "Die Infusionsthierchen als vollkommene organismen" (1838), stating, among other things, that there was absolutely no connection between animal and plant worlds.
The German translation of Kutorga's book — "Naturgeschichte der infusionsthiere" — was published in 1841. Thus, Prof. S.S. Kutoga can be recognized as one of the first Russian scientists to study protists (first of all, ciliates). He wrote in 1839: "This discovery [of infusoria — SF] excited all the researchers, and everybody in possession of a microscope hastened to enjoy the sight of perpetual movement of this invisible world" (Kutorga, 1839. S. 2). Therefore, for the naturalists of those times it was mainly a kind of fun and the majority of them even did not try to study these tiny organisms deeply.
An undoubted service of Prof. Kutorga was that he made Russian scientists familiar with the cellular theory ofSchleiden and Schwann. In 1841, Kutorga wrote an article where he underlined that Schwann's book had "opened a new field of microscopical investigations, provided new views and notions on living animal and plant organisms and in some way changed the direction of natural sciences" (Kutorga, 1841). Kutorga himself, however, later left zoology (even though he continued teaching it at the ISPbU) and concentrated his scientific activities mostly on geology and paleontology. It was only
L.S. Cienkowski. St. Petersburg, 1860s.
in the beginning of the 1850s that Kutorga, in his university lectures, attributed protozoa — rhizopods and ciliates — to the group of "primary animals", underlining their unicellular nature and the lack of complicated organs (Fokin, 2001a).
At the ISPbU in the middle of the XIX century, there appeared a true protistological researcher: L.S. Cienkowski (1822-1887), a student ofS.S. Kutorga, who became an outstanding Russian scientist, one of the founders of Russian protistology (Raikov, 1949; Fokin, 2001a; Kuznicki, 2003). He was a graduate of the ISPbU (1844) and later (however, unfortunately, only for a short time, 1854-1861) was a professor of botany therein. His doctoral thesis "On the low algae and ciliates" (Cienkowski, 1856) convincingly demonstrated that morphological characteristics and physiological processes in the organisms at the borderline between animals and plants coincide, and no clear characters delimitating the two kingdoms can be found. Prof. Cienkowski was the one to introduce microscope into the practice of university teaching in Russia; he discovered and described several dozens of various protists and traced life cycles of many of them. He was one of the first to investigate cyst formation in ciliates and to show experimentally the impossibility of self-birth in protists (Cienkowski, 1859). In his lectures, Cienkowski propounded the idea of the connection
between unicellular and multicellular animals and was the first to draw attention to the phenomenon of symbiosis in lower organisms.
Zootomical Cabinet of the ISPbU — the emergence and development of a protistolo-gical centre in Russia
Further, a new outbreak of interest in protists' study was already associated with the Zootomical Cabinet, and even among botanists in the ISPbU such activity also had a successful continuation: e.g., in some studies of professors A.S. Famintzin (1835-1918) and Ch.J. Gobi (1947-1919). Other researchers who contributed significantly to the progress of protistological studies at the Zootomical Cabinet of the St. Petersburg University are listed below and their brief life histories are provided.
Konstantin S. Mereschkowsky (1855-1921) -a protistologist, botanist, anthropologist, writer; the fist scientists who got his own education at the Zootomical Cabinet of the ISPbU (1880) under the supervision of Prof. N.P. Wagner (1829-1907) and became deeply interested in protistology. In the beginning of his scientific career (1877-1886), Mereschkowsky was the one to study ciliates and other protists mostly systematically (Fokin, 2008). He was the author of the first faunistic reviews on the unicellular organisms of the North and South of Russia: "Studies of the protozoans of the Russian North" (Mereschkowsky, 1878), "Materials to the ciliate fauna of the Black Sea" (Mereschkowsky, 1880), and "On some new or little-known ciliates" (Mereschkowsky, 1881). Mereschkowsky's teaching at the Zootomical Cabinet of the ISPbU (18831885) helped several then well-known researchers-zoologists. Moreover, among them was Yu.I. Andrusova (1863-1942), who studied the ciliate fauna of the Black Sea as well, and thus she could be recognized as one of the first Russian female protistologists (Fokin, 2004).
Who knows what could have been the scientific future of Mereschkowsky in the ISPbU, but in the autumn of 1886, he left the University due to complicated family circumstances and the decline ofhis health. In any case, as it was re-described very recently (Fokin, 2021a), finally he came back to protistology again (mainly in the Kazan University) where he succeeded to publish a seminal article "About Nature and Origin of the Chromatophores in the Plant Kingdom" (Mereschkowsky, 1905)
K.S. Mereschkowsky. St. Petersburg, 1884.
and then a book entitled "A Theory of Two Plasms as the Basis of Symbiogenesis, a New Theory of the Origin of Organisms" (Mereschkowsky, 1910). Prof. Mereschkowsky developed his novel view on evolution, the theory of symbiogenesis, using a classical protistological object — diatoms. These studies nowadays rank Mereschkowsky among the founders of the symbiogenetic theory of eukaryotic cell origin (Khahina, 1979; Fokin, 2005; Kutschera and Niklas, 2005). Unfortunately, Mereschkowsky did not have any successors in the ISPbU who would support the protistological direction of studies. However, it is definitely not the case of the famous protistologist belonging to the next university generation — W.T. Schewiakoff, who was a student in the ISPbU at that time. It is more than likely that they might have met at the University, but Schewiakoff never mentioned Mereschkowsky as his university teacher. However, a delusive viewpoint that Schewiakoff was a student of K.S. Mereschkowsky can still be found in the Internet publications.
Wladimir T. Schewiakoff (1859—1930) — the most well-known Russian protistologist of the late
XIX — the first third of the XX century; Doctor of Philosophy at Heidelberg University, Doctor of Zoology, Professor at the ISPbU and the Imperial Women's Pedagogical Institute (in the Soviet times — Irkutsk State University); corresponding member of the Imperial St. Petersburg Academy of Sciences (later — the USSR Academy of Sciences and the Russian Academy of Sciences). With his fundamental research, Prof. Schewiakoff made a significant contribution to the study of unicellular organisms belonging to ciliated protists and acan-harian radiolarians (Fokin, 2019, 2021b). He can be indicated as a founder ofprotistological scientific school at the ISPbU.
It is important to briefly remind the readers about some pages of his biography related to the University. Wladimir Schewiakoff graduated from the Reformated Church School (1877) in St. Petersburg — his hometown, and entered the Mining Institute where he studied until 1881. Then Wladimir decided to transfer to the Department of Natural Sciences at the Physical-Mathematical Faculty of the ISPbU. From 1881 until 1884, he was specializing there at the Zootomical Cabinet under the supervision of Prof. N.P. Wagner as an external student. Schewiakoff had not completed the University course when he decided to continue his education abroad. He chose Heidelberg University. The director of the Zoological Institute in Heidelberg was Prof. Otto Butschli (1845—1920), a zoologist and protistologist of the worldwide fame. Butschli found Schewiakoff to be not only a bright student (1885—1889) but also a reliable assistant and later (1891—1894) — a colleague.
In October 1893, the Imperial Academy of Sciences in St. Petersburg elected Dr. Schewiakoff a laboratory assistant at the newly formed Special Zoological Laboratory of the Academy, headed by Prof. A.O. Kowalevsky (1840—1901). In spring 1894, Shewiakoff defended the Magister's (PhD) thesis "On the Biology of Protozoa" (Schewiakoff, 1894) in St. Petersburg and in the autumn of the same year he finally settled in Russia. In February of 1896, he defended in the ISPbU the Doctoral (Dr. Sc.) thesis entitled "The Organization and Systematics of Infusoria Aspirotricha (Holotricha Auctorum)", which simultaneously was published as a book (Schewiakoff, 1896) and the scientist became the Head of the Zootomical Cabinet.
One of the best known publications of Sche-wiakoff is the monograph "Acantharia". The scientist worked on it at the Naples Zoological Station
(the Gulf of Naples) in 1899, 1902, 1905 and 1925/ 1926. This means that the scientist conducted a significant part of the research dedicated to acantharians (at that time a class of Radiolaria) being a member of the ISPbU (Schewiakoff, 1902).
The Imperial Academy of Sciences elected Prof. W.T. Schewiakoff its corresponding member in 1908. In 1911, Schewiakoffwas appointed Vice-minister of the Ministry of People's Education of Russia; and he held this position until 1917. With this appointment, unfortunately, he left the University forever.
Thus, Prof. Schewiakoff gave almost 20 years of his life to the ISPbU. As a teacher-zoologist and the Head of the Zootomical Cabinet he radically reorganized the institution (now — the Department of Invertebrate Zoology at the St. Petersburg State University) and the entire system of teaching this biological discipline at the University, raising the methodology of teaching students to a new quality level. W.T. Schewiakoff is one of the founders of the Russian protozoological scientific school and the educator of a whole galaxy ofwell-known native zoologists and general biologists (Fokin, 2011b).
Here it is necessary to emphasize the obvious connection of the Russian protistological scientific school with the German school of Prof. O. Butschli. At the Zoological Institute of the Heidelberg University, more than 50 Russian students and scientists studied and worked in the last quarter of the XIX — early XX centuries; some of them (first of all, W.T. Schewiakoff) were engaged in protistological topics (Fokin, 2012).
From a scientific point of view, it should be especially noted that Prof. Schewiakoff was a pioneer in several research areas. He was the first who described the mitosis in rhizopods and apicomplexians; detected the mitochondria in ciliates; described after Prof. O. Butschli the nucleoid in bacteria, and gave the first scientific explanation of the gliding movement type in gregarins. The scientist made the first attempt to apply Darwin's views to the zoogeography ofprotists and was one of the first who attempted to justify the saltational way of evolution (Fokin, 2021b).
It may be especially important to stress within this article that during his work at the University, several his students became true protistologists or carried out important protistological investigations. Some of them made a significant contribution to the further development ofthe scientific school laid
W.T. Schewiakoff. St. Petersburg, 1906.
down at the ISPbU at the border of the XIX and XX centuries, largely due to the efforts made by Prof. Schewiakoff. Below, I briefly mention some ofthose names and fates, although not all of them became protistologists.
Sergei I. Metalnikoff (1870-1946) — a protis-tologist, zoologist, physiologist and immunologist. He specialized in invertebrate zoology at the ISPbU (1894-1896) under the supervision of Prof. W.T. Schewiakoff. Metalnikoff earned his PhD in zoology (1908) and became the Head of the Biological Laboratory in St. Petersburg, as well as a deputy director of the Higher Lesgaft Courses (1909). In a short time, he was also selected for a professorship in zoology at the St. Petersburg Higher Women's Courses (1911). Before 1917, Prof. Metalnikoff was already a well-known authority within the Russian scientific community (Fokin, 2011a). As a protistologist, he made seminal studies dedicated to the feeding process in ciliates and their ability to select food (Metalnikoff, 1907), physiology of intracellular ingestion in Protozoa (Metalnikoff, 1911), and on the question of immortality of unicellular Protozoa (Metalnikoff and Galadjiev,
S.I. Metalnikoff. St. Petersburg, 1907.
S.V. Averinzev. Heidelberg, Germany, 1900.
1916).
After the Bolsheviks' coup in 1917, the scientist emigrated from Russia to France, where he headed one of the laboratories at the Pasteur Institute (Paris). There, the main directions of Prof. Metal-nikofFs research were immunology and the control of agricultural pests using bacteria that are pathogenic to pests (Fokin, 2022).
Sergei V. Averinzev (1875—1957) — a protis-tologist, zoologist, ichthyologist, teacher. Averinzev entered the Natural Sciences Department of the Physical-Mathematical Faculty of the ISPbU in 1896. He was a student of Prof. W.T. Schewiakoff. In his fourth year at the University, Averinzev was awarded the University gold medal for the research work "Morphology and Systematics of the Family Halteria Clap. et Lachm" (Ciliophora, Oligotrichia). He graduated from the University with the 1st degree diploma and stayed there for a two-year period of training for a professorship. Averinzev published his first work in protistology, "On the Protozoa Fauna of Bologoye and its Environs", in 1900. In 1903— 1904, Averinzev passed the Magister's examinations and in 1904—1909 was the Head (working-manager) of the Murman Biological Station of the St. Petersburg Naturalists' Society. Having defended the Magister's thesis (PhD) "Freshwater Rhizopo-
da" (1906), he was appointed Privatdocent in the Zootomical Cabinet of the ISPbU. He worked there until 1909. At the same time, he taught at the Higher Women's (Bestuzhevskye) Courses at the Psychoneurological Institute and at the Agricultural Courses named after I.A. Stebut, where he was elected a professor. From 1901 until 1913, S.V. Averinzev often worked in the field ofprotistology in Western European countries. For instance, he stayed at the Zoological Institute of Heidelberg University under the supervision of Prof. O. Butschli (1901).
In the summer of 1902, he investigated the chemical composition of the shells of marine Rhizopods at the Naples Zoological Station. He visited the zoological stations and museums of Trondheim, Bergen, Hamburg and Helgoland in order to study their operation (1904). In 1910, he was awarded the scholarship of the Academy of Sciences for conducting research at the Buitenzorg Botanical Garden in Java. He also visited a number of places in the Eastern Africa. In 1914, he defended his Doctoral thesis (Dr. Sc.), "Materials on Morphology and History of Parasitic Protozoa" (microsporidia, gregarines, and ciliates).
Since 1917, Averinzev mainly concentrated on the problems of commercial ichthyology and fisheries at the Barents Sea. In this field, he worked then in Moscow, Tver and Yakutsk. After his return
from Siberia, the scientist was invited to lecture on zoology in the Fergana Pedagogical Institute (Uzbekistan). With several breaks, he taught at the Pedagogical Institute until 1940. Then S.V. Averinzev resumed teaching in Moscow, in a regional Pedagogical Institute, where he was the Head of the Department of Zoology until May of 1953. Averinzev was the author of more than 250 works on protistology, ichthyology and fishery, including several manuals and textbooks on zoology.
Valentine A. Dogiel (1882-1955) — an outstanding Russian zoologist, well known in the world by his numerous publications in the international scientific journals (1906-1927). Dogiel graduated from the Zootomical Cabinet of the ISPbU in 1904 under the supervision of Prof. Schewiakoff who mentioned Dogiel as his favorite student. Dogiel got his PhD in 1910 and Dr. Sc. degree in 1913. During a couple of first decades, the scientific activity of V.A. Dogiel was mainly within protistology, but using parasitic representatives from different groups of protists. Finally, Prof. Dogiel got a very wide scope of scientific interests. He became a parasitologist (Dogiel, 1941) and the founder of ecological parasitology — a special, still rapidly developing branch ofparasitology. His achievements in the fields of comparative anatomy and evolution were summed up in the fundamental monographs: "Comparative Anatomy of Invertebrates" (Dogiel, 1938, 1940) and "Oligomerization of Homologous Organs as One of the Major Ways of Animal Evolution" (Dogiel, 1954). His "first love", which lasted, however, the whole lifetime, was protistology. He studied protists from various systematic groups: gregarines, flagellates, ciliates, microsporidia, radiolaria (Mazurmovich and Poljansky, 1980). A fundamental monograph "General protozoology" (Dogiel, 1951) was the result of 40 years of investigations carried out by Prof. Dogiel and his disciples, and of the university course "General Protistology", which Dogiel had taught for many years. The latter was the first Russian handbook in this field of biology. Fourteen years later, it was published in English, after the text had been refined by Dogiel's disciples (Dogiel, 1965). As a teacher ofinvertebrate zoology, he succeeded in publishing the textbook (Dogiel, 1934), which had a number of editions and is still partly in use by students.
Prof. Dogiel had educated many protistologists. It is worth mentioning briefly here the almost forgotten name of G.N. Gassovsky (1893-1959),
V.A. Dogiel. Petrograd, early 1920s.
a graduate of the Zootomical Cabinet of 1916. Despite the repeated changes in the type of scientific and teaching activities as well as places of residence (from Vladivostok to Kishenev and Cherkasy), he had retained the interest to study protists for the whole his life (Gassovsky, 1960). Some of Prof. Dogiel's former students became well-known scientists abroad. Among such his disciples were C.A. Hoare (1892-1984), later on known as an English protistologist; the founder of the Polish Academy of Sciences - a protozoologist J. Dembowski (1889-1963), and some others.
Boris Ph. Sokolov (1889-1979) — a proto-zoologist, medical doctor, biochemist, politician, journalist, and writer. Sokolov was a graduate of the Zootomical Cabinet at the ISPbU (1912) partly under supervision of W.T. Schewiakoff, partly in the times of V.A. Dogiel, and the Medical faculty of the Psychoneurological Institute (1916). He participated as a military doctor on the medical train at the South-Western front of the World War I (1917). Sokolov was a member ofthe All-Russian Constituent Assembly (1918) and of the provisional government of the Northern region, Arkhangelsk
B.Ph. Sokolov. Lakeland, USA, end of 1950s.
(1920). Scientifically, he worked as an assistant at the Zoological Department of the Petrograd Scientific Institute (1914-1920), mainly on protists. In 1920, he emigrated from Russia. He lived a long life, two thirds ofwhich had passed outside of Russia. Having left Russia in 1920, he certainly could not expect to find a place in the West and continue his scientific career. However, unlike most of his colleagues, he was seriously involved in the political processes that took place in Russia during the years of the Civil War and the revolution of 1917. After emigration to Estonia, Dr. Sokolov soon received a British visa and briefly stayed in London, where he apparently failed to find a job. For some time, also of no avail, he tried to settle in Prague and Paris and, from the spring of 1922, temporarily settled in Belgium. In the West, he mainly studied the mechanisms of cancer as well as the biological properties of antibiotics and vitamins. His PhD dissertation (defended in Prague, in 1924), however, was dedicated to regeneration phenomenon in ciliates, and part of this research was carried out in Russia (partly at the ISPbU). Dr. Sokolov was also a journalist and a writer. He was the author of more than 30 popular scientific books
published in the United States. In America, his scientific career was quite successful: after several changes, he had found a permanent scientific position. During the last decades of his life, Prof.
B.Ph. Sokolov worked as the Head of the Cancer Research Laboratory at Lakeland Southern College (Florida, USA).
Cecil Arthur Hoar (1892—1984) — zoologist-protistologist. Hoar's life story differed from that of other zoologists who had left Russia. Cecil's father, the journalist Arthur Stowell Hoare, was an Englishman, and his mother, a Russian citizen named Amalia Chalet (Aimee Shalet), was a singer who had studied her craft at the Kiev Conservatory.
C.A. Hoar was born in Holland and had a dual (English-Russian) citizenship, but he had lived with his mother in St. Petersburg from the age of six.
He graduated from the Petrograd State University in 1917 with the first-class degree in zoology. There, Hoar had studied invertebrate zoology, as well as protistology, under the supervision of Prof. V.A. Dogiel. He was then appointed an assistant at the Army Medical Academy. A promising start for a scientific career, but, regrettably, it was interrupted. The Red terror of late 1918—1919 also affected foreigners living in Petrograd. For Hoar, fortunately, it only resulted in two months in prison in 1919. A year later, in June 1920, he and his wife left Russia. The departure was formally authorized by the Academy of Sciences of Russia as a mission to study blood parasites and pathogenic protozoa in the scientific centres of France and England. Hoar was to return to Russia in April 1921. Given that Cecil Arthur's father was English, and the scientist had dual citizenship, going to England was rather akin to repatriation.
Hoar worked for about 40 years in the Department of protozoology in the Welcome Bureau, London, UK. His curators at the initial stages of his career in the UK were scientists who were known in the field, such as professors C. Dobell and Ch. Wenyon. Hoar studied some representatives from different groups of protists that included the parasitic species — Amoebozoa, Ciliophora, Apicomplexa — but most of all parasitic flagellates — Trypanosomatina (Kinetoplastidae). The fundamental monograph by Prof. C. A. Hoar, "The Trypanosomes of Mammals — A Zoological Monograph" (1972), is dedicated to one of the most dangerous parasites for humans, namely, parasitic protozoa — trypanosomes. In total, Hoar published
C.A. Hoar. London, United Kingdom, 1930.
more than 180 works; in 1950, he was elected a Fellow of the Royal Society (Goodwin and Bruce-Chwatt, 1985). It is symbolic that academician C.A. Hoar, soon after his visits to the USSR (1959, 1960) took part, together with professors Yu.I. Poljansky and E.M. Cheissin, in editing and publishing the English-language translation of "General Protistology" — the wonderful book by their teacher, Prof. V.A. Dogiel, published in Oxford in 1965 (Fokin, 2022).
The most famous of Dogiel's disciples in pro-tistology during the Soviet time were professors Yu.I. Poljansky (1904-1993), A.A. Strelkov (19031977), E.M. Cheissin (1907-1968), I.B. Raikov (1932-1998) and L.N. Seravin (1931-2010), who had worked in Leningrad - St. Petersburg and were, in their turn, the teachers of almost all contemporary Russian protozoologists (Skarlato, 1999; Muller, 1999; Fokin, 2007, 2018) as well as of many researchers of protists in other countries. Using their joint efforts and now also the efforts of their students and followers, the St. Petersburg School of Protistology has gained the worldwide
fame and continues its successful development in the XXI century.
References
Cienkowski L.S. 1856. On the low algae and ciliates. SPb (in Russian).
Cienkowski L.S. 1859. On the selfbirth. SPb (in Russian).
Dogiel V.A. 1934. Zoology of Invertebrates. BioMedPress, Leningrad (in Russian).
Dogiel V.A. 1938. Comparative anatomy of invertebrates. Vol. 1. Uchpedgiz, Leningrad (in Russian).
Dogiel V.A. 1940. Comparative anatomy of invertebrates. Vol. 2. Narkompros, Leningrad (in Russian).
Dogiel V.A. 1941. General parasitology. Uchpedgiz, Leningrad (in Russian).
Dogiel V.A. 1951. General protistology. Sov. Sc. Press, Moscow (in Russian).
Dogiel V.A. 1954. Oligomerization of homologous organs as one of the major ways of animal evolution. Leningrad University Press, Leningrad (in Russian).
Dogiel V.A. (revised by J. I. Poljanskij and E. M. Cheissin). 1965. General protozoology. Oxford University Press, England.
Ehrenberg C.G. 1838. Die Infusionsthierchen als vollkommene organismen. Leipzig.
Fokin S.I. 2000. Professor W.T. Schewiakoff: life and science. Protist. 151: 182-189.
Fokin S.I. 2001a.The rise and development of protistology in St. Petersburg, Russia. Protistology. 2 (1): 68-72.
Fokin S.I. 2001b. A love 50 years long. Professor V.A. Dogiel as a Protistologist. Protist. 152: 379387.
Fokin S.I. 2005. The 100th anniversary of Russian Mereschkowsky's symbiogenesis paper. ISS Symbiosis International, 10, 4.
Fokin S.I. 2007. Phylum Ciliophora Doflein, 1901. General description. In: Protista Part 2. Handbook on zoology. "Nauka", St. Petersburg, pp. 371-414.
Fokin S.I. 2008. Konstantin Sergeevich Me-rezhkovskiy (1855-1921). In: Russian scientists at the Naples zoological station 1874-1934. (Eds: Fokin S.I. and Groeben Ch.). Giannini Editore, pp. 125-129.
Fokin S.I. 2011a. Documents do not burn -
they simply are not read. Biologist S.I. Metalnikov (1870-1946). In: Right to the name, Biography of 20th century. SPb, pp. 181-189 (in Russian).
Fokin S.I. 2011b. People and collections of the Zootomical Cabinet of the Imperial Peters-burg University. In: The Zootomical Cabinet (Department of Invertebrate Zoology) of St. Petersburg University. On the 140th anniversary of its founding (Ed. Fokin S.I.). Scientific PressKMK, Moscow, pp. 12-42 (In Russian).
Fokin S.I. 2012. Otto Bütschli and his Russian students. Science in Russia. 188 (2): 89-94.
Fokin S.I. 2019a. Protistologist Wladimir Timo-pheyevich Schewiakoff(1859-1930). On the shores ofthe Gulf of Naples. Protistology. 13 (4): 246-255.
Fokin S.I. 2019b. Relentless flight of time. Yury Ivanovich Poljansky (1904-1993) by eyes of the pupil, colleague, historian of science. Studies in the history of biology. 11 (4): 9-45.
Fokin S.I. 2021a. Konstantin Sergeyevitch Merezhkovsky (1855-1921) - the protistologist who discovered the origin of eukaryotic cells. Protistology. 15 (1): 41-45.
Fokin S.I. 2021b. Unforgotten life. Wladimir Timopheyevitch Schewiakoff (1859-1930). KMK Scientific Press, Moscow (in Russian with English summary).
Fokin S.I. 2022. The roads of Russian emigrant zoologists. In: In foreign lands: the migration of scientists for political or economic reasons (Eds.: Borgato M.T. and Phili Ch.). Trends in the History of Science. 5. Chapter 12, pp. 263-290.
Gassovsky G.N. 1960. The practical significance of ciliated protists and the history of their study in Ukraine. Collection of works of Zoological Museum. Kiev. 29, pp. 58-90 (in Ukrainian).
Goodwin L.G. and Bruce-Chwatt L.J. 1985. Cecil Arthur Hoare. 6 March 1892-23 August 1984. Biographical Memoirs of Fellows ofthe Royal Society. 31: 295-323.
Gorjaninov P.Ph. 1837. Zoology and Zootomy used for the general benefit, especially for medicine. SPb (in Russian).
Haeckel E. 1866. Generalle Morphologie der Organismen: allgemeine Grundzüge der organischen Formen-Wissenschaft, mechanisch begründet durch die von C. Darwin reformirte Decendenz-Theorie. Berlin.
Haeckel E. 1879. The evolution ofman: a popular expositions. Vol. 1. New York.
Khahina L.N. 1979. The problems of symbio-genesis. Nauka, Leningrad (in Russian).
Kutorga S.S.1839. Natural history of infusoria. SPb (in Russian).
Kutorga S.S. 1841. Review of science in the Western Europe in 1839. Min. Peop. Educ. Journal. 29, part V: 1—46 (in Russian).
Kutschera U. and Niklas K.J. 2005. Edno-symbiosis, cell evolution, and speciation. Theory in Biosciences. 124: 1—24.
Kuznicki L. 2003. Protozoologia w Polsce 1861—2001. Warszawa: Polska Akademia Nauk (in Polish).
Mazurmovich B.N. and Poljansky G.I. 1980. Valentin Alexandrovich Dogiel. Science Press, Moscow (in Russian).
Mereschkowsky K. 1878. Studies on protozoans of the North of Russia. Travaux de la Société des naturalistes de St. Petersbourg. 8 (1): 203—288 (in Russian).
Mereschkowsky C.S. 1880. Materials for ciliates fauna ofthe Black Sea. Proc. SPb Soc. Nat.11, 3-13 (in Russian).
Mereschkowsky C. 1881. On some new or little-known infusoria. Annals and Magazine of Natural History. 5th series. 7: 209-219.
Mereschkowsky C. 1905. Über Natur und Ursprung der Chromatophoren im Pflanzenreiche. Biol. Centralbl. 25: 593-604.
Mereschkowsky C. 1910. Theorie der zwei Plasmaarten als Grundlage der Symbiogenese, einer neuen Lehre von der Entstehung der Organismen. Biol. Centralbl. 30: 278-303; 321-347; 353-367.
Metalnikoff S.I. 1907. On feeding process of ciliates and its ability to select a food. Proc. SPb Nat. Soc. 38: 175-181 (in Russian).
Metalnikoff S.I. 1911. On physiology of intracellular ingestion in Protozoa. News SPb Biol. lab. 11: 3-121.
Metalnikoff S.I. and Galadjiev M.A. 1916. On the question of immortality of unicellular Protozoa. News Imp. Acad. Sci. SPb, pp. 1809-1816 (in Russian).
Mûller M. 1999. Per aspera ad Protozoa et astra. Yuri I. Polyansky and Russian protistology. Protist. 150: 431-437.
Raikov B.E. 1949. L.S. Cienkowski - the founder of microbiology in Russia. Microbiology. 18, 562-570 (in Russian).
SchewiakoffW.T. 1894. On biology ofprotozoa. Memoires de l'Academie des Sciences St. Petersburg. 75 (1): 1-96 (in Russian).
Schewiakoff W.T. 1896. Organization and systematics of Infusoria Aspirotricha (Holotricha
Auctorum). Memoires de l'Academie des Sciences St. Petersburg, VIII ser., 8 (1): 1-420 (in Russian).
Schewiakoff W. 1902. Beitrage zur Kenntniss der Radiolaria-Acanthometrea. Memoires de l'Academie Imperiale des Sciences de St. Petersbourg. VIII ser., 12 (10): 1-40.
SchewiakoffW. 1926. Acantharia. Fauna e flora del Golfo di Napoli. Pubblicazioni della Stazione zoologica di Napoli. 37: 1-755.
Skarlato S.O. 1999. Protistology in Sankt-Petersburg. In: Petersburg Academy of Science among the other academies of the world. SPb scientific center of RAS Press, SPb (in Russian).
