Introduction

Lysenkoism can be defined as a system of pseudoscientific ideas and concepts supported and enforced by the governmental structures to strengthen the ideology and establish direct control over the academic community (Krementsov and DeJong-Lambert 2017). The phenomenon of Lysenkoism emerged for the first time in the history of science in the 1930s–1950s, during the period of Joseph (Iosif) Stalin’s personality cult in the USSR, and derives its name from the surname of “people’s academician” Trofim Lysenko. A large body of literature is devoted to Lysenko and Lysenkoism, and the problem of Lysenkoism has regularly been discussed at conferences and panels (Krementsov and DeJong-Lambert 2017). This academic interest is understandable: the catastrophic consequences of Lysenko’s hold on Soviet science are well known. Recent attempts to revive Lysenko’s ideas, so-called “Neolysenkoism” (Graham 2016; Kolchinsky 2014), are now also being discussed with vigor; the purpose of these attempts is to show that Lysenko was a forerunner of some of the modern academic disciplines (for instance, epigenetics). For obvious reasons, Lysenkoism has almost always been discussed in connection with genetics and agriculture. However, as demonstrated convincingly by Weiner (1988), the first victim of ideologization of science was Soviet ecology, which became essentially a test model for Stalinists’ intrusion into science. The key role in the ideological suppression (but not yet a complete denouncement) of ecology was played by the notorious Isaak (Isay) I. Prezent, who in the 1930s was one of the foremost proponents of dialectics in Soviet biology and the leader of dialectizationFootnote 1 (the imposition of dialectics) in Leningrad (Kolchinsky 2014).

It is universally known that Lysenko’s ideas were primarily concerned with heredity. In discussing heredity, Lysenko and his followers postulated the inseparable unity of an organism and its environment, and hence the system of Lysenko’s biological beliefs can be called the general theory of ecology, as was rightfully noted by Graham (1987). Lysenko’s ideas thus encompassed not only genetics but ecology as well. Nevertheless, no one has ever studied how Lysenkoism was presented in the field of aquatic ecology (hydrobiology). This seems all the more surprising given that the environmental history of Russia is currently being intensively developed (Breyfogle 2018). However, the studies of environmental historians have been focused primarily on the analysis of interactions between science and the Soviet regime in landscape transformation and natural resource management and in the development of geographical science, rather than in specific aspects of theoretical biology (Oldfield and Shaw 2016).

The whole field of the history of hydrobiology in the USSR remains largely unexplored.Footnote 2 As a result, there is a significant gap in our knowledge about this scientific community under Lysenkoism. It is clear that the models of scientists’ behavior under ideological pressure can be different for different sciences (due to their specific characteristics), and this should be taken into consideration when interpreting the phenomenon of Lysenkoism.

One of the central problems of modern-day hydrobiology is the study of production and dynamics of organic substances from the perspective of the ecosystem approach (Alimov 2003; Talling 2008). The history of the ecosystem concept is a popular topic among historians of biology (Bocking 2013; van der Valk 2014), but their studies have been concerned mostly with the general principles of ecosystem functioning (for instance, energy flows), which had been developed by scientists of the Anglo-American school (including R. Lindeman, G. E. Hutchinson, and Howard and Eugene Odum, to name only the most distinguished). Under this approach, an early stage in studying the productivity of bodies of water remains outside the focus of attention (Schwarz and Jax 2011).

In particular, there is no mention in the modern literature that the first production theory was developed by professor Vladimir I. Zhadin, who worked at the Zoological Institute of the USSR Academy of Sciences (Zoological Institute AS USSR) (Rizhinashvili 2017a). The analysis of Zhadin’s theory may present special interest because it reflects, as “in a drop of water,” the scientific content of hydrobiology of that period (Schwarz and Jax 2011): productivity, dynamics of biocoenoses (communities), typology of bodies of water, and other aspects. The theory was first published in 1940 in the monograph Fauna of Rivers and Water Reservoirs (Zhadin 1940). The monograph summarized the results of the author’s studies on patterns of changes in the riverine fauna observed in the process of building water reservoirs. The monograph was dedicated by the author to the October Revolution (“Great proletarian revolution”). The book was thus meant as a response to the calls of the time, the great communist building projects. It should be noted that at that time, there was no unified production concept anywhere in the world, only fragmented ideas on different topics.

To what extent was Zhadin’s theory a response to ideological exigencies of the time? What is the reason that it was consigned to obscurity and has never gained support in modern science? If the essence of this theory was purely ideological, did it have reasonable cognitive elements? These questions are of special interest in light of the analysis of relationships between science and the government in the USSR in the period of the personality cult. The goal of this study is to analyze the relationship between ideological and cognitive aspects in scientific theories (using Zhadin’s forgotten theory as a case study). To date, there have only been a few attempts to analyze this relationship in historical studies (see, for instance, Shaw and Oldfield 2008), and these studies have usually been conducted for the practice-oriented branches of knowledge (Hubbard 2014). The present study will allow for better understanding of the extent of ideological influence on science. The results of such a study may also be valuable for establishing the criteria to identify pseudoscience and for developing programs to fight pseudoscientific ideas (Gordin 2017; Marks 2017).

To achieve this goal, I will give an overview of the situation in Soviet ecology and hydrobiology in the 1930s. The focal point of this review will be the problem of productivity. Then I will examine Zhadin’s scientific activity during that period and present and analyze his productivity theory for bodies of water. Finally, I will consider the prospect of further development of Zhadin’s theory and its place in hydrobiology and discuss possible reasons for why this theory was forgotten.

An Overview of Ecology and Natural Resource Management in the USSR in the 1930s

By the beginning of the 1930s, the idea of transformation and reconstruction of nature for the benefit of the developing socialist society had begun to be firmly implemented in the Soviet Union (Vorontsov 1992; Weiner 1988; Weiner 2002). The exploration, inventory, and exploitation of resources were also instigated by famine that spread across several regions of the USSR in 1932–1933 (Tauger 1991).

Nature was viewed by the Soviet government as something hostile to humanity, which needed not only to be exploited but to be conquered. The spontaneity of natural forces was thought to be detrimental to the economy. The idea was that nature lacks order: all species are randomly mixed, and therefore intrusion into biological communities could not have any negative impact. This meant that flora and fauna needed planned and targeted changes and improvement. The ongoing collectivization and industrialization were supplemented by much praised acclimatization (Lysenko’s vernalization would very soon be added to this triad). Acclimatization was seen as essential for the reconstruction of fauna, to increase, for instance, its economic value. Prezent justified acclimatization by stating that nature always has untapped resources that can be utilized by an introduced species. Lysenko later used this thesis to put forward an idea about the lack of overpopulation in the biological communities and, as a consequence, the absence of intraspecific competition (Lysenko 1949).

In this ideological climate, the studies that were done as pure science—for the sake of science, without any practical application—were declared not only useless but even harmful. The slogan of that period was “practice as the criterion of truth” (Roll-Hansen 2008). In November 1935 at the Congress of Stakhanovites, Stalin spoke his famous words:

Science which has severed contact with practice, with experience —what sort of science is that? If science was the thing it is represented to be by certain of our conservative comrades, it would have perished for humanity long ago.Footnote 3

All research had to be directed toward finding a way to tame nature. Stalin wished to exclude any uncertainty from both society and nature (Graham 1987).

The mid-1930s was a time in Soviet science and biology that saw heated and wide-ranging discussions on a number of theoretical problems, often held with active participation of Party ideologists (Kolchinsky 2014). Unlike later discussions [particularly, the infamous 1948 VASKhNILFootnote 4 session (Rossianov 1993)], these debates were not only driven by ideology, but were also incited by unresolved objective problems of science. At the same time, the Soviet Union was hit by the first wave of mass repressions, and “conservative comrades” (scientists with suspected heterodox views) began to perish. The pioneer of the studies on the productivity of biological communities in the Soviet Union was the ornithologist and ecologist Vladimir V. Stanchinskiy (Weiner 1988). Stanchinskiy was the first in the world to establish the quantitative relationships between autotrophic and heterotrophic components of the community. He is rightfully viewed as a forerunner of R. Lindeman’s trophodynamic ideas. Stanchinskiy was arrested in 1933. Among the hydrobiologists who fell victim to repressions before 1932 was, for instance, Arvid L. Behning, the director of the Volga Biological Station. Behning was the pioneer of hydrobiological investigation of rivers in the USSR (Talling 2008). In 1933, all personnel of the Murmansk Biological Station were arrested and the station ceased to exist. Scientific discussions and meetings, therefore, could have far-reaching ramifications.

In ecology, this kind of discussion took place in 1934 at the Botanical Institute of the USSR Academy of Sciences (Anonymous 1934). Prezent openly attacked the use of mathematical methods in ecological studies (Weiner 1988). The targets of his critique were the works of Stanchinskiy. He thought the “usurpation” of biological phenomena by mathematical methods was unacceptable. This attack on mathematics was likely engendered by Prezent’s desire to impose a system of ideas in biology that would oppose mechanistic and reductionist views (Weiner 1988). He advocated the idea that “specific biological content” should not be reduced to “algebraic symbols” (Anonymous 1934, p. 61) and that quantitative conclusions should be tested by experimental methods. In the examples that he provided to illustrate the absurdity of using mathematical methods for the analysis of the study’s results, he simplified to the extreme and distorted the data and conclusions presented by Stanchinskiy and his colleagues. Prezent was well aware of the implications that his words would have in the ideological climate of the mid-1930s and was deliberately preparing the ground for this to happen. Consequently, a seemingly reasonable and logical demand had led to quantitative methods being declared a manifestation of formalism. It is well known that classical genetics was also condemned by Lysenko and Prezent as formalistic for its quantitative laws. In Lysenko’s view, only Morganists, lacking proper understanding of the unique properties of life, were seeking to “turn biological science into mere statistics” (Lysenko 1952, p. 579).

The rejection of mathematics had laid the groundwork for the attack on production studies that dealt with values of biomass. Prezent acrimoniously criticized the production studies for their detachment from the practical needs of the time. He asked: “Is any biomass equally important economically?” (Anonymous 1934, p. 61), and gave as an answer an example of biomass of grass and grain. Prezent stated that it is the patterns found in “concrete biological objects” that should be studied instead of the abstract values of production and biomass. For the purposes of this study, I can draw the important conclusion that Prezent differentiated between the economically important biomass and the biomass of objects without practical value. In other words, there are “useful” and “useless” biomass. But what was the attitude towards the productivity problem in hydrobiology?

Discussion on the Problem of Biological Productivity of Bodies of Water in 1936–1937

Hydrobiology was from the outset an interdisciplinary science. This interdisciplinary nature and the complexity of hydrobiology was objectively determined by the multifaceted character of the aquatic environment. It is clear that bodies of water are acted upon not only by living organisms, but are also shaped by physico-chemical conditions. The terrestrial ecosystems, by contrast, are primarily governed by biological factors. In the 1920s, hydrological and biological studies were so closely interlinked that, for example, hydrochemical studies were published in hydrobiological journals. The same specialists often simultaneously conducted biological, chemical, and hydrological studies, and the simultaneous application of techniques from different disciplines was very common. As a result, by the early 1930s, many hydrobiologists began to think about the content of their science and its relationships with associated disciplines.

It should be noted that hydrobiology, both in the Soviet Union and outside of it, has never been unresponsive to economic needs. In fact, the complex of productivity-related problems and the term “production” were themselves born from attempts to address the challenges of commercial fisheries.Footnote 5 In Russia, the first attempts to estimate the productive capacity (productivity) of an aquatic system (or, more precisely, to evaluate possible fish reserves based on the oxygen balance) were made as early as 1908 by Arseniy A. Lebedintsev at the Nikolsky fish rearing plant. Lebedintsev spoke of the oxygen dynamics as a manifestation of lake’s “breath,” i.e., the lake was likened to an organism with all its inherent physiological functions (Lebedintsev 1908). He was not alone in this: in the early twentieth century, the tendency to compare an ecosystem with an organism was quite common in world ecology. With respect to the terrestrial communities, this paradigm was expressed most clearly in the works of Frederick Clements (van der Valk 2014). In the second half of the twentieth century, this idea would continue to develop at the new systemic level (see, for example, Taylor 1988; Worster 1977).

In 1936 and 1937, the Zoological Journal (Zoologicheskij zhurnal) published a series of papers of renowned hydrobiologists Zenkevich (Brotzky and Zenkevich 1936), Muraveiskiy (1936), Karzinkin (1936), Vinberg (1936), and Skadovsky (1937), focused on the problems of understanding biological productivity and methods of studying it. At the time of this discussion, two major avenues of hydrobiological research had emerged in Soviet hydrobiology. One was the ecologico-physiological (hydrophysiological) school founded by Skadovsky, while the other was interdisciplinary limnological studies. Limnological studies were conducted at numerous lake stations and involved comprehensive physico-geographical descriptions of bodies of water in terms of their typology, life activity patterns of organisms, and cycling of matter.

Two different approaches had also been adopted for the assessment of production. Production was evaluated either by measuring the biomass of species and species groups or by using chemical methods of biomass assessment (for example, by measuring consumption or release of oxygen) with subsequent recalculation (for example, for the weight of fish). It is universally known that it was the chemical approach that was used in 1932 by Vinberg to develop a technique of measuring primary production in a water body (the “light and dark bottles method”). These works have gained international acceptance and recognition, and Vinberg’s method is still widely employed by hydrobiologists. Viktor S. Ivlev was developing ideas on the efficiency of energy use by organisms during their growth. All of these studies together laid the groundwork for the energy-oriented approach to the methodology of studying aquatic communities (Lindeman 1942). Soviet hydrobiology and its branch studying productivity progressed to such a high degree that August Thienemann, the famous European limnologist, was prompted to speak about the need for European students to study the Russian language. In 1925, Thienemann wrote in the German newspaper Allgemeine Zeitung:

It is our deepest conviction that the coming generation of Western European scientists—at least, in some disciplines—cannot do without studying the Russian language, unless they want to deprive themselves of the most precious scientific data. (see Behning 1925, p. 220)

However, a general theory of productivity was not created.

The question of how the production process can be controlled for the benefit of economic development of aquatic resources was also left unanswered. This failure was especially important given the existing ideological conditions. The lack of a unified methodology of production studies became the reason for calling the discussion. By the time of this discussion, more or less universally accepted definitions of the terms “productivity” and “production” had been established in world science. Productivity was the ability of a water body to determine a certain type and rate of generation of organic matter in living organisms (Brotzky and Zenkevich 1936). Production was the increase in biomass produced by organisms over a certain period of time. However, as was noted by Leonid L. Rossolimo, it is unclear “which substances and processes must be taken into account, how productivity must be measured and what units must be used” (1934, p. 14). Rossolimo proposed to replace the assessment of productivity with the study of balance of the organic matter in a water body. At the same time, the zoologist and hydrobiologist Brodskiy (1934) argued that only the study of patterns of organization and dynamics of aquatic communities and the biology of commercial species can help in studying productivity.

The participants of the discussion noted a number of theoretical problems, which, in their opinion, engendered the existing ambiguity and uncertainty of the “productivity” concept and methods used in productivity studies. These problems included: insufficient consideration of the issues concerning the disciplines studying the hydrosphere (i.e., whether productivity and production should be studied by biological disciplines alone, together with geographical and chemical sciences, or by a special discipline) (Zenkevich, Muraveyskiy, Vinberg); the use of the term “productivity” to refer both to the water body and the species (Vinberg); and the ambiguity as to what was to be considered the final product of the production process (Karzinkin).

These scientists were almost unanimous in agreeing that biological productivity must be studied by dealing with a water body as an integrated whole. This viewpoint was especially strongly expressed by Muraveyskiy, who considered bodies of water as elements in the global hydrological drainage system. Vinberg spoke of a whole combination of conditions in an aquatic system and regarded the types of bodies of water as representing various developmental stages with respect to the balance of substances they contain. Karzinkin expressed a simplified position, viewing a water body as a producer of economically valuable product (primarily “bioproduct,” or living organisms), because, as he asserted, productivity exists only in relation to the product. The conclusion of the scientists (except for Karzinkin, who did not adopt any unequivocal position in this respect) was that biological productivity must be studied by geographical disciplines with the participation of hydrobiologists. Muraveyskiy even proposed the term “biohydrology” to designate a new branch of hydrology that would study productivity: “The data from hydrobiology cannot be used as the basis for developing the productivity theory” (1936, p. 580).

In this scenario, hydrobiology would remain a branch of ecology. Zenkevich went even further and proposed to make hydrobiology itself a part of hydrophysics. On the other hand, all authors agreed that organisms are active participants in the production process, because the activity of an organism makes it a driving force for the cycling of matter. Therefore, a water body cannot be viewed as a higher integrated entity–“an organism of higher order” or a “microcosmos”–as was done by foreign authors who proposed typological schemes (for example, Thienemann).

Foreign scientists, in the opinion of these authors, had adopted and still retained their idealistic positions. An understanding of a body of water as an integrated whole in terms of productivity implies the knowledge of adaptations of individual species of organisms (Skadovsky) and their biocoenotic relationships. Muraveyskiy noted that in order to study an object as an integrated whole, one has to examine the interactions of this object with its constituent parts (i.e., the water body and organisms).

The authors indicated an undeniable economic value of the production processes. For instance, Zenkevich maintained that the productivity problem is the cornerstone that links science and practice. Muraveyskiy affirmed that productivity is determined not only by natural factors, but also by social interactions. He drew a parallel between the productivity of bodies of water and soil fertility. Vinberg spoke about the possibility of making a prognosis of fishery productivity based on the extent to which the primary organic matter was utilised in a water body. Brodskiy (1934) was already considering the problem of productivity before this discussion and viewed the productivity as a tool for transformation of nature in response to the call of Socialist construction. It should also be mentioned that the terms “productivity” and “fertility” were treated by most hydrobiologists as synonyms; the same was true for the terms “production” and “yield.” Almost all participants in the discussion (except Vinberg) thought that the final product of the production process is an organism, especially an economically valuable one. Vinberg, by contrast, suggested that silt deposits can also be regarded as the final product.

In general, the ideological aspect of this discussion was essentially manifested only in proclaiming the problem of detachment of theory from practice without any serious insinuations or accusations against anyone (except for foreign scientists, who were widely accused of idealism). The examination of materials of this discussion shows the sincere desire of scientists to understand, above all, the system of disciplines studying productivity, the place of hydrobiology in resolving this complex of problems, the methodology of study, and the possibility of using the study results in economic practice. Almost all authors came to the conclusion that the solution to the productivity problems must be found in studying the processes of cycling of matter in connection with physiological adaptations of aquatic organisms. Since productivity is a property of a water body, this problem must become the purview of geographical disciplines.

Nevertheless, the subject matter of the discussion clearly reveals the orientation towards economically valuable biomass, which was especially expressed in Karzinkin’s paper. For him, the most important is question how biomass in a water body is represented (for instance, by worms or harvested mollusks) rather than what is the level of the total biomass in a water body. This was likely the manifestation of Prezent’s influence. The discussion of the problem of biological productivity comes down to the controversy of “organisms versus environmental conditions.” What has to be studied first: organisms or chemical processes? After all, the production process is the process of generation of organic matter, and hence a constituent part of the cycles of matter. From a theoretical viewpoint, the problem of productivity and biomass of organisms is logically integrated into a more global set of problems of the cycles of matter and energy in the biosphere. It should be emphasized that by 1920, the seminal ideas of Vladimir I. Vernadskiy, who regarded the living matter as a geochemical force of planetary scale, were already popular among Soviet scientists (Levit 2011).

By 1940, the discussion generated by this uncertainty had produced its theoretical fruit. This fruit was the theory of biological productivity of bodies of water developed by the Soviet zoologist and hydrobiologist Vladimir I. Zhadin. As mentioned above, this theory was first described in a separate chapter of Zhadin’s 1940 monograph. However, judging from his previous publications, the theory had taken its final form no later than 1937.

Professor Vladimir I. Zhadin (1896–1974)

Vladimir I. (IvanovichFootnote 6) Zhadin primarily studied the fauna of rivers, the systematics and variability of freshwater mollusks, and he was also involved in studying the problem of predicting the dynamics of biological communities in water reservoirs. He was a world-renowned scientist. One of the signs of his international recognition was that in 1965, he won the Medal of Einar Naumann, which is awarded by the International Association of Theoretical and Applied Limnology (Zhadin was the vice-president of this association). The text of his diploma states: “To Professor V.J. Shadin,Footnote 7 Vice-President of the Association, editor of ‘The Freshwater Life of USSR,’ for his outstanding work on rivers and artificial lakes, and on mollusks.”Footnote 8

Zhadin’s official status in the USSR, however, was quite modest. For instance, he was not a member of AS USSR, but only an Honored Science Worker of the Russian Soviet Federated Socialist Republic (RSFSR). From 1934 until almost the end of his life, Zhadin worked at the Zoological Institute AS USSR. For much of that period, he was the head of the still-existing Laboratory of Freshwater and Experimental Hydrobiology.

Zhadin was born and raised in Murom, a town located on the banks of the Oka River, one of the largest Russian rivers. As a boy, Vladimir made natural history excursions together with other children.Footnote 9 In the Murom real school, where he studied, pupils founded a club of natural science lovers.

Zhadin’s scientific career began before the October Revolution. In 1915, he was assigned by the Moscow zemstvo (the district council in the Russian Empire) to study the geographic distribution of the snail Lymnaea truncatula, which had a practical significance as an intermediate host of the digenean Fasciola hepatica. Zhadin then studied at the Kazan and Moscow Universities (with pauses because of his service in the army). He attended the lectures on hydrobiology that were given by academician Sergey A. Zernov at the Petrovskaya Agricultural Academy. In 1920, after the turmoil of the World War I and the Civil War, Zhadin launched the study of various types of small bodies of water associated with the Oka River at the Oka Biological Station, which he founded. Zhadin supervised the study of the flora and fauna of these bodies of water and the variability of invertebrate species. For the purposes of our study, it is important that, as Zhadin himself wrote:

even my first acquaintance with numerous floodplain bodies of water that the Oka River had formed by changing its watercourse set an interesting goal for the station: to study the patterns in the genesis of the biota of the floodplain bodies of water in connection with their geological genesis. (1991, p. 20)

During the subsequent years until 1934, he conducted or supervised the qualitative and quantitative studies of the fauna of various types of bodies of water at the Oka Biological Station. During these fieldworks, Zhadin amassed a significant amount of material that allowed him, for instance, to develop the typology of rivers and other bodies of water (including artificial ones) in terms of water quality. This was especially important since the goals that he had to achieve during his expeditions had an explicitly practical orientation. One of the primary goals was the water supply of towns and cities. The so-called Omutninsky Expedition of 1932 studied the questions of organization of the aquatic economy in connection with construction projects of the second five-year plan. In particular, the study was concerned with “possible changes in water quality as a consequence of river impoundment” (Zhadin 1991, p. 37).

Thus, by the early 1930s, the “riverine” problems became the central theme of Zhadin’s scientific work. He focused on patterns of changes in faunas (primarily, the benthic fauna) and the chemical regime observed when stretches of rivers unaffected by humans were transformed from their pristine condition to ponds created by dams. This study was much in keeping with the goals of the scientific research in the period of industrialization. Apparently, Zhadin was able to understand the imperatives of the times. This was in part facilitated by the need to accomplish those practical tasks that were assigned to him by the economic officials of the region where he worked. This practical orientation, however, as mentioned above, was evident from the beginning of Zhadin’s academic career. A comprehensive, years-long study of one large river (Oka) that has stretches of natural differences in the regime and composition of biological communities allowed him to make confident predictions of the changes occurring as a result of the artificial (technical) intrusion into the communities of different rivers.

Zhadin had the opportunity to present these results at a large Faunistic Conference held at the Zoological Institute AS USSR on 3–8 February 1932. For Zhadin, this conference would be the first step leading towards his outstanding future scientific work. As he later recollected, the conference served for him as “the impetus for moving to Leningrad” (Zhadin 1991, p. 33).

This conference had its own ideological backstory (Weiner 1988). It was convened by academician Zernov, the Director of the Institute, who launched the reorganization of the former Zoological Museum. The conference was held within the walls of the Zoological Institute created from the Museum, but the actual “shadow” initiator of the conference was Prezent. The year 1932 brought an end to the biological “cultural revolution” in the USSR, which had begun in 1928 (Kolchinsky 1999). It was the year when the relative freedom that philosophical discussions had enjoyed during the previous years ceased to exist and plans of socialistic construction began to be forced into the practice of biological research. Numerous discussions organized with Prezent’s participation were called upon to get rid of dissenters and their research projects, but some of the biologists still tried (often with some success) to express publicly their opinions, which often ran against the political agenda of the time. This is exactly what happened at the 1932 Faunistic Conference.

Zhadin was invited by his mentor Zernov to give a presentation at the conference. This and many similar conferences were organized in such a way that presentations were created by teams of several specialists, but were given by one scientist who presented on behalf of the whole team. The title of Zhadin’s presentation was “A biocoenotic and faunistic study of regions and their biotopes in connection with economic operations that change the living conditions of bodies of water” (Zhadin 1934). Zhadin’s team included ichthyologist Mefodiy I. Tikhiy; hydrobiologists Dmitriy E. Beling, G. I. Dolgov, and Gleb Yu. Vereshchagin; and ecologist Stanchinskiy. In his presentation, Zhadin acknowledged the unprecedented rates at which socialism was changing the nature. He praised the “colossal socialist construction” that “has made great strides towards changing the geographic face of the USSR” (1934, p. 33). He mentioned, as if in passing, that “different sewage effluents discharge into bodies of water.” Zhadin also acknowledged that there are only a few publications studying the impact of engineering works on fauna (“faunistic and biocoenotic face”) of the bodies of water. He emphasized the necessity of planning in hydrobiological works and reconciliation of the scientific 5-year plan with economic goals. He urged the conference to study the environmental problems caused by hydrotechnical construction (decline in plankton biomass, difficulties for the reproduction of migratory fish, waterlogging, receding water levels) and outlined ways to resolve them (for instance, fast-growing fish breeds). The presentation showed his concern for nature conservation problems, which he expressed in a very thoughtful and subtle way.

The results of Zhadin’s own observations and studies formed a separate part of the presentation, although he did not emphasize it explicitly. For instance, he spoke (albeit only in broad outlines and abstract terms) about changes in qualitative and quantitative composition of the fauna living on granulometrically different types of substrate. He associated this problem with blocking rivers with dams, which resulted in precipitation of the suspended particles and siltation downstream from the dam.

Zhadin’s presentation was met with overwhelming approval. Prezent was naturally one of the participants in the discussion, and he acknowledged that the presentation was the response to the calls of the five-year plan. Zhadin completed his presentation at the end of the discussion by saying that “faunists must work in close cooperation with industry. This is the path that we must follow, the path along which the communist party will lead us” (1934, p. 47). All of Zhadin’s proposals were included in the resolution adopted by the conference. The resolution proclaimed that the main goal of hydrobiological studies was “to make a prognosis for the biological use of all categories of engineering structures … and to find ways to overcome difficulties caused by construction” (Zhadin 1934, p. 48).

Zhadin’s presentation attracted attention to nature protection problems, showing his great personal courage. This courage had also been shown during the discussion, when he mentioned the name of the already-reproved Stanchinskiy, a fact especially emphasized by Weiner (1988). This presentation became programmatic both for Soviet hydrobiology in general and for Zhadin’s own works.

To apply his ideas in practice and verify his conclusions, Zhadin had to expand the scope of his research by studying rivers in various geographical zones. This became possible when Zhadin took up a position at the Zoological Institute. While there, Zhadin was able to launch the study of the fauna of rivers and water reservoirs in connection with hydro-construction. His goal was to find general patterns in this phenomenon, which had to be achieved on rivers and water reservoirs in different geographic landscapes, because “depending on the geographic (in the wide sense of the word) conditions, hydrotechnical construction can have a different impact on the regime of the bodies of water, and by extension on their fauna.” According to Zhadin:

When we study the impact of the hydrotechnical construction on the riverine fauna, we can distinguish two phases of the study. The first phase is the study conducted before the dam was built […] The second phase is the study of the water body after the construction was completed, the study of the river downstream from the dam… (1940, p. 527)

Throughout the 1930s (especially in 1934 and 1935), Zhadin made numerous expeditions to study large rivers and river systems of various landscapes of the USSR. All of these landscapes were affected to a greater or lesser extent by hydrotechnical construction: they either had water reservoirs on their territory or the construction of these reservoirs was underway. Zhadin was astonished by differences in the diversity of the animal populations in the rivers with different water transparency. He found that at the bottom of the Rion reservoir, in which “our legs sank into the mud deeper than our knees,” “there is no organic life” (Zhadin 1991, p. 43). On the stones of the transparent Zangu River, Zhadin recognized the presence of “a rich and very diverse fauna consisting of larvae of insects, crustaceans, mollusks, worms” (1991, p. 44). In 1939, he became involved in resolving the problem of “Big Volga,” the creation of a system of water reservoirs on the largest river of European USSR.

All of this fieldwork allowed Zhadin to gather a colossal amount of material on the qualitative and quantitative composition of the fauna from different types of rivers and sections of rivers. As he wrote: “Thus our works covered the rivers located at various latitudes ranging from the deep south of the USSR to the Subarctic” (Zhadin 1940, p. 545). As he explained:

We have a group of southern mountain rivers of the Transcaucasian region (Rion, Kura, Zangu), rivers of the Russian plains (Dnieper, Oka), north-western lacustrine (flowing from lakes) rivers (Volkhov, Yavan, Pestovka, Valdayka), and Subarctic lacustrine rivers (Varzuga, Vyala, Muna). (Zhadin 1940, p. 722)

In total, Zhadin and his co-workers collected “over a thousand benthic samples and several thousand plankton samples” (1940, p. 545). These results supplemented significantly the studies he had begun on the Oka River. This material also enabled him to establish general patterns in the transformation and succession of benthic riverine communities.

Zhadin argued that the leading factor in this process was the accumulation of sediments suspended in water and carried by currents, which in its extreme form can lead to siltation. Zhadin concluded that “the suspended matter continuously precipitating from the turbid water impedes the development of the fauna” (Zhadin 1991, p. 43). According to Zhadin, “Suspended substances carried by river waters reflect the characteristics of the geological structure of the catchment, the structure of the soils and human’s economic activity on them…” (1940, p. 727).

Suspended particles can play completely opposite roles in the life of hydrobionts. Their primary role is to serve as food, but this is only possible if suspended particles are organic. On the other hand, if a high concentration of suspended matter reduces water transparency, the suspended particles have a negative effect on lighting conditions, limiting the penetration of light energy into the water column. The accumulation of suspended organic substances in the bottom sediments (siltation) results in deterioration of the oxygen regime and, by extension, of the breathing conditions of aquatic animals (Zhadin 1946a).

Zhadin classified benthic biocoenoses by the substrate type and by their relation to the speed of the current (Zhadin 1940). He presumed that these factors play a leading role in the distribution of the fauna. He subdivided the biocoenoses into lithorheophilic (biocoenosis of the hard rock bottom with fast currents), phytorheophilic (biocoenosis of plant thickets with fast currents), psammorheophilic (biocoenosis of sand with fast currents), pelorheophilic (biocoenosis of silt with fast currents), pelophilic (biocoenosis of silt in lentic or slow-moving waters), and phytophilic (biocoenosis of plant thickets in lentic waters). Zhadin noted that this classification was to a significant extent developed by his wife and colleague, Ekaterina S. Neizvestnova-Zhadina (1937). Most importantly, Zhadin posed the question about the origin of the riverine fauna by considering the proportion of the primary and secondary bodies of water (Zhadin 1946b). Rheophilic components of biocoenoses belong predominantly to the primary bodies of water, while the organisms of the lentic waters are secondary aquatic.

Zhadin traced the faunal composition of bodies of water in a series of their successive transformations (1940). He found it easier to illustrate this series using as case studies those bodies of water that have or had various degrees of connections with the watercourse of a river. As the currents in the bodies of water grow slower and the water level drops, the rheophilic biocoenoses become replaced by pelophilic and phytophilic ones. In this way, as accumulation increases, the succession of benthic biocoenoses proceeds from a complex of primary aquatic to that of the secondary aquatic species.

The conclusions drawn from Zhadin’s collected material allowed him to develop the prognostic principles for water reservoirs. The detailed discussion of these principles is beyond the scope of this article. For the present study, the most important aspect is that a water reservoir is considered as a derivative of the landscape in the whole catchment area of the reservoir (Zhadin 1940). In addition to the prognostic principles, Zhadin also formulated his theory of biological productivity for bodies of water, which is discussed below.

Theory of Biological Productivity of Water Bodies

While considering the problem of productivity of bodies of water as part of the processes involved in the cycles of organic substances, Zhadin proposed using the term “accumulation” to denote the build-up processes of these substances. He regarded the accumulation as the foundation for the relationships between the water body and the catchment: “Rivers of different geographic landscapes have also different magnitudes of accumulation processes” (Zhadin 1940, p. 946).

Production is calculated from an increase in biomass of living organisms, and it is organisms that, according to Zhadin, determine “the degree of quantitative perfection and qualitative value of accumulation of organic substances,” with the organic substances being “primarily a function of the species (or group) composition of an animal (and vegetative) inhabitants.” He stressed that different types of production can result from the same degree of accumulation, “depending on the composition of the fauna and flora that assimilate the accumulating organic substances” (Zhadin 1940, p. 946). In other words, animals and plants (“bionts” in Zhadin’s terminology) “are responsible” for the use of resources of the organic substances supplied in the process of accumulation (“organic food”). Therefore:

I use the term bioecological provision to denote the provision of a water body or any of its parts with animals (and, in other cases, plants), whose ecological nature reflects the living conditions in it. (Zhadin 1940, pp. 946–947)

This provision (the composition of the fauna and flora of a given water body) is the result, on the one hand, of adaptation of organisms to the conditions of the water body and, on the other hand, of the biogeographic history of the region and its water areas. Therefore, in Zhadin’s opinion, the organism and the environment complement each other; the organism reflects the environment.

While developing his productivity theory, Zhadin formulated the rule of productivity. According to this rule, productivity of a body of water is a function of the accumulation of organic substances and bioecological provision in this water body. Zhadin’s goal, however, was deeper. Under the state-planned economy, he was to engage in “developing methods for controlling productivity” (Zhadin 1940, p. 944). He had to find a way to explain and direct the production process. To achieve that, he needed to determine exactly how the biomass of benthic animals depends upon the level of accumulation and on the number of species. Zhadin suggested a diagram showing this three-pronged association and the rule of productivity (Fig. 1). When accumulation is low, both the number and biomass of organisms are also low. As accumulation rises, the species diversity grows rapidly and so does the biomass. A further increase in accumulation leads to a decline in species diversity, but the biomass keeps growing to a certain critical point, although at a slower rate. If there is an even higher and more rapid increase in accumulation processes, the biomass shows a steady decline. Consequently, the highest biomass (the highest productivity) is observed when the species number is relatively low. Thus, the optimum of species diversity and the optimum of biomass do not coincide. Zhadin gave the following commentary on these patterns:

Fig. 1
figure 1

Diagram illustrating the underlying principles of the biological productivity theory: accumulation and bioecological provision (after: Zhadin 1940, p. 950). The X axis shows the number of species, while the Y axis shows the mass (of accumulating substances or organisms; “гp” is the Russian abbreviation for “gram”). Dashed line: biomass; solid line: bioecological provision (number of species); bold line: accumulation. The numbers in the circles are not explained by Zhadin

Full size image

Under a moderate accumulation of organic substances, which promotes the expansion of bioecological provision of the biocoenosis, biological productivity is increased because of the involvement in the production process of a large number of ecologically diverse elements; if the degree of accumulation is increased somewhat more, this time causing a reduction in biological provision, biological productivity continues to grow, but this happens due to an acceleration in growth and reproduction of individual components of the biocoenosis; if accumulation crosses a certain critical point causing a further decrease in biological provision, biological productivity shows a decline and in the case of hyper-accumulation can fall down to zero. (1940, p. 949)

Zhadin revealed two fundamental ecological patterns concerning the production process. First, his diagram provides an illustration of the general principle of optimum: to obtain a system with the maximum productivity, the level of accumulation should be selected in such a way as to provide the highest level of production. Second, the highest level of productivity is possible only at low levels of species diversity, in relatively simple, species-poor communities. When there are only a few species, their populations can increase their biomass by an intensive growth and reproduction because of the lack of competitors. This pattern is now well known and was expressed in a quantitative form (Alimov 2003).

As follows from the chapter in Zhadin’s monograph in which he expounds upon his theory, the rule of accumulation with its explanatory diagram lies at the core of the theory. The following text in the monograph contains no further discussion of the theory or any explicitly emphasized concepts. All explanations that follow serve to clarify the application of his concepts of productivity to the theory of hydrobiology and to the practice of aquatic economy. In particular, Zhadin believed that his theory could explain the distribution of organisms in bodies of water. For instance, relict forms leave water reservoirs because of the high level of accumulation of organic substances in these bodies of water. The endemic fauna “retreats” from those parts of bodies of water that have increased levels of siltation (which happens, for instance, in the inlets of Lake Baikal). In all these examples, Zhadin clearly and persistently associated the accumulation with siltation. He thought it could also be successfully applied to explain the distribution and quantitative development of other groups of aquatic organisms. When applied to fish, this theory explains migration patterns and the origin of early and late runs of fish. In this case, water turbidity becomes the key factor, because highly turbid waters prevent the spawning of migratory fish.

Zhadin rightfully viewed his theory as an answer to the questions posed over the course of the discussion of 1936–1937. First of all, Zhadin consistently advocated the biological nature of the productivity problem. This problem could and must be studied only by hydrobiology. He gave the definition of hydrobiology as a biological science. The duality of his production theory entailed two lines of research. He believed that, first, the water body should be studied as an integrated whole by using a combined effort of hydrologists, hydrochemists, and hydrobiologists of different specialties (botanists, zoologists, microbiologists) under the supervision of a hydrobiologist “with a sufficiently wide scientific field of expertise” (Zhadin 1940, p. 959) (study of accumulation). Second, the research should involve the study of geography and ecology of individual organisms and biocoenoses, and the metabolism of organisms in connection with the environmental conditions, as well as the development of ecological spectra of species (study of provision). All of this could be done only within the field of hydrobiology.

While addressing the problem of relationships in the “organism-environment” system, Zhadin persistently promoted the idea of their unity. Historical traits that determined the formation of the fauna were the underlying causes of this unity. For instance, the genesis of fauna of the bodies of water in the European part of the USSR had been shaped by the impact of the ice age. In Zhadin’s view, this impact was manifested in a higher intensity of erosion processes. In turn, this increased the turbidity of the river water and forced many organisms to leave rivers for other bodies of water. As a result, riverine biocoenoses became unsaturated by species and, for this reason, led to the reverse migration of organisms that successfully proceed in the upstream direction from river mouths (the most striking example was the mollusk Dreissena).

On the other hand, Zhadin considered the organism as a constituent part of biocoenosis. He rejected the notion of this supraorganismal system as a certain integrated union, which is treated almost as a “superorganism.” Zhadin argued that biocoenoses are highly dynamic systems and the links between species are not too rigid. The elements of one type of biocoenosis can be present in another.

According to Zhadin, the rate at which biocoenoses replace one another had increased many times over by economic activity. He was convinced of this from observations over the transformation of the riverine fauna during the construction of water reservoirs. The intensity of the erosion processes had also increased, and so there was now an urgent need to harness these spontaneous processes. This clearly frames the problem of managing productivity for economic purposes.

Zhadin argued that the control over productivity was only possible along two major lines of his theory: accumulation and bioecological provision. Zhadin provided the necessary recommendations. For instance, he stressed the importance of addressing the problems of pond fertilization and reducing soil erosion in the catchment in order to control accumulation. The development of acclimatization methods for organisms was essential to solving the problem of the formation of provision. This was only possible if aquatic biocoenoses were not saturated with species. Nevertheless, acclimatization of individual species should be planned by taking into account the relationships of these species with other components of biocoenoses. Scientists should also be able to predict those implications that can arise with the invasion of new species.

While addressing the economic challenges of that period, Zhadin at the same time paid much attention to the question of species conservation. He spoke of the necessity to preserve the most vulnerable primary aquatic organisms sensitive to the oxygen concentration in water (lithorheophilic fauna). To achieve this, hard-rock-bottom habitats should be left intact. If this is impossible, they should be artificially constructed. In his opinion, in the case of fishes, the measures should include the construction of fish passes and fish nurseries, and the organization of artificial spawning grounds.

Zhadin considered the final product of the production process to be the biomass of an economically valuable species (for instance, fish or mollusks). Zhadin had an acrimonious argument in the press with Vinberg, and regarded this method of assessing the primary production as “strange.” In Zhadin’s opinion, the results of measuring the rate of photosynthesis did not have anything in common with measuring production. This conclusion was based on the observation that a high level of photosynthesis in a lake can be associated with fish production, which was “represented by small-sized crucian carps” (Zhadin 1949, p. 208). By contrast, a lake with a low photosynthetic production can have a significant amount of valuable and large-sized fish. The same approach can be applied to benthic organisms.

Whether or not a live form qualifies as productive depends on its importance for economy. For instance, oligochaetes are often buried deep in the substrate, and hence are unavailable for fish. Consequently, they cannot be regarded as a productive group of zoobenthos. Large bivalves with a massive shell are well protected from being eaten by commercial fish, but the commercial harvesting of these animals can provide the pearl industry with high-quality material. For this reason, mollusks cannot be viewed as non-productive organisms. This example demonstrates Zhadin’s deeply economical approach to biological problems. As we will see, in his later publications Zhadin took these ideas even further by dividing production into “useful” and “harmful.”

Zhadin’s economic orientation led him to criticize the balance-oriented approach to the study of bodies of water. He described it as “formally correct,” but stressed its “lack of goal orientation” (Zhadin 1940, p. 958). The notion of the negative balance (when destruction prevails over production) is illogical, because in this case, in Zhadin’s opinion, the secondary production (of harvested species) may in fact be rather high.

The system of Zhadin’s views on production can be summarized as follows. A water body should be regarded in terms of productivity as an integrated whole to be studied by hydrobiology as a biological science. However, the integrity of the water body has a dynamic, rather than an absolute, nature. It emerges through the processes of accumulation that link together the water body and the catchment. Accumulation is responsible directly (through the granulometric composition of the substrate) and indirectly (through nutritional and gas conditions) for the succession of biocoenoses (at least, benthic biocoenoses). This succession results in the condition with a certain level of species diversity of the fauna. The species composition of hydrobionts “provides” the processing of inflowing organic substances. The organisms generate the highest biomass under a certain intermediate level of accumulation and a relatively low number of species. The active accumulation of the organic matter in the bodies of organisms during their growth is the production process. The final product of this process is the biomass of economically valuable species.

Thus, Zhadin’s main achievement in his attempt to resolve the problem of productivity is that, unlike other hydrobiologists and limnologists, he was able to add real context to the idea of “a water body as an integrated whole.” A water body is not simply a part of the runoff process (Muraveyskiy), an element of the water balance of the Earth (Rossolimo), a part of cycles of matter (Zenkevich), or the maker of products (Karzinkin). Rather, the water body according to Zhadin is a part of the “water body-catchment” system united by the process of accumulation. Accumulation is the factor that ensures the integrity of the water body by uniting all its parts. Accumulation is equally applicable to rivers and lakes. Zhadin was able to show a regular nature of the dynamics of biocoenoses in the river viewed as a single unified system. Other hydrobiologists before Zhadin saw only one (water body) or the other (organism) side of the production process, apparently due to their professional specializations. In his attempt to address practical economic problems, Zhadin had sought and found the zone of the maximum productivity. Rather than using empty rhetoric, he demonstrated functional interrelationships (even if only at the qualitative level) and developed the logical model of the production process. This opened possibilities for practical manipulation of the production processes.

What ideological aspects can be identified in Zhadin’s theoretical concept of the productive phenomena? First of all, the theory is clearly narrative in nature and lacks quantitative reasoning. In the illustrating diagram (Fig. 1), the Y-axis shows the quantities of the mass, but they appear “suspended in the air” in the context of the whole discourse. It is likely that in so doing, Zhadin took into account the ideological situation by deliberately dismissing mathematical reasoning, because otherwise he could have become the target of serious criticism with all its ensuing consequences.

Zhadin then gave clear practical recommendations, but he stressed the necessity of conservation measures for pinpointing the organisms that appeared to be most vulnerable. Zhadin believed that not only nature is spontaneous, but that human activities also can be haphazard and therefore require control and direction. At the same time, Zhadin did not think that the links between organisms in the biocoenosis are so tight that they cannot be directed for practical purposes. This opened the possibilities for reconstruction of biocoenoses, in particular, by using acclimatization, but he warned against its indiscriminate and inconsiderate use. Zhadin thus dismissed the notion of the balance and harmony in natural communities, which appears in line with the ideological precepts. He also presumed that an organism and the environment correspond to each other historically. This concurs with one of the notions of Lysenkoism about the “tight-fittedness” of the organism to the environment. Finally, Zhadin accorded the prime importance to the environmental conditions (nutritional and gas conditions), rather than to the interactions of organisms. This idea also agrees with views of Lysenko, who rejected, for instance, intraspecific competition. In this way, Zhadin kept a balance between ideological needs and purely scientific ideas. And, while he was oriented towards the economic product, he also did not forget about species conservation.

The extent to which Zhadin was not only close to the practical needs of that period, but was also sincere in trying to respond to the challenges of the time, can be illustrated by a draft note entitled “On accumulation for fish” discovered in his personal archive. In 1940, Zhadin worked on Lake Ilmen, where he studied and tested measures against fouling of hydrotechnical structures by the mollusk Dreissena. During his fieldworks, he attended one of the production meetings at the kolkhoz (Soviet collective farm) “Trudovoy rybak” (“Working fisherman”). Zhadin wrote that one of the fishermen said the following: “Why can’t we catch large fish in Lake Ilmen? Because the wind muddies the water of the lake and the fish escapes to the clean water of rivers.”Footnote 10 This note shows how much he appreciated even the commonsensical observations of ordinary workers. Judging from the fact that this was kept “for personal use,” his interest was sincere. For him, the observation of a fisherman was a source of information and food for thought. This is a true example of surmounting “the rupture between intellectual and physical labor,” as proclaimed by Nikolay I. Bukharin (Roll-Hansen 2005).

It would be wrong to assume that Zhadin’s views in hydrobiology were purely practical, aimed only at improving the product yield. Zhadin was also a theoretician. Considering siltation as the leading process caused by accumulation, he proceeded to the discussion of the faunal genesis in the bodies of water. He further elaborated his views on productivity in the publications of the post-war period.

Creative Adoption of the Elements of Lysenkoism’s Biology: The Genesis of the Fauna of Continental Waters and New Goals of Hydrobiology

As discussed above, the theoretical significance of Zhadin’s ideas on production is expressed more strongly and at a new level in his treatment of the problems of evolution of bodies of water and the origin of their faunas. The problem of the fauna’s genesis is the cornerstone of Zhadin’s productivity theory of bodies of water (Zhadin 1946b). This is an obvious consequence of addressing the productivity problems based on the observations of changes in the riverine fauna during the process of hydrotechnical construction.

During that period, the search for pathways and patterns in the origin of faunas was one of the objectives of the ecological research program in the USSR, which was outlined by Daniil N. Kashkarov (1937). This leader of Soviet ecology of animals urged the use of the dynamic approach to the natural phenomena: “The understanding of the need to study not only statics, but also dynamics is increasingly acknowledged in biology” (Kashkarov 1937, p. 120). Mirzoyan wrote that “introduction of faunocenogenesis [genesis of faunas] into the sphere of interest of ecology was the principled position of Kashkarov” (2013, p. 25). However, Kashkarov emphasized the primacy of climate’s influence, while Zhadin stressed the key role of geological processes (notably, erosion).

Zhadin considered the accumulation process to be the leading factor in the evolution of bodies of water:

As the processes of accumulation of organic and organogenic substances tend to accelerate with the age of the Earth and reach their maximum intensity in our epoch, when, in addition to the products of erosion of spontaneous origin, tremendous amounts of substances set in motion by humans… settle down in the bodies of water, there occurs secular succession of various bodies of water with their biomes. (1940, p. 956)

Thus, accumulation is the result of erosion. Zhadin then expounded on the patterns of the genesis and replacement of faunas:

In the water courses and bodies of water with an uneroded catchment… there is an abundant development of the primary aquatic fauna… whereas the water courses and bodies of water with an eroded catchment… show the depression of the primary aquatic fauna, its replacement by the secondary aquatic fauna, and at the final phases of this sequence (marshes) the replacement of the aquatic fauna by amphibious and terrestrial. (1940, p. 957)

Zhadin developed the typology of bodies of water based on their genetic relations. The basis for this classification was the type of the catchment that determined the level of accumulation. In the works published after 1940, Zhadin unequivocally emphasized that the shallowing and waterlogging of bodies of water are the phenomena that help resolve theoretical problems of transition of organisms from the aquatic to terrestrial lifestyle (Zhadin 1946b).

Since accumulation is most often manifested as siltation and silt removes oxygen from water, oxygen conditions and respiratory adaptations are the high-priority aspects in assessing the productive ability of a water body. Zhadin’s main concern in further developing his ideas is the need to conduct ecologico-physiological experiments on respiration of hydrobionts. The study of oxygen consumption reveals “pathways of adaptation of aquatic animals to life in variously silted bodies of water” (Zhadin 1949, p. 200). This research program can clearly be seen in the abstract of Zhadin’s presentation given during a hydrobiological seminar at the Zoological InstituteFootnote 11 (see also the full text and detailed discussion in Rizhinashvili 2017b). The presentation was scheduled for late July of 1948 and was entitled “Oxygen consumption by aquatic animals as an indicator of the genesis pathway of the fauna of continental bodies of water.”

In this presentation, Zhadin examined a wide range of biochemical aspects of “the battle for oxygen.” Zhadin also intended to touch on the question of the origin of the flying stage in insects. In his research program, in keeping with the spirit of ideology, he proclaimed “the unity of field observations and laboratory experiments,” with field observations defined as the study of “the distribution of species of the study animals in bodies of water of different nature” (apparently, with different magnitudes of bottom siltation).

Zhadin went even further and proposed to establish a new branch of hydrobiology, evolutionary hydrobiology. Quite unexpectedly, he presented the previously denounced analogy between the development of organisms and the bodies of water by using the well-known Haeckel-Müller’s biogenetic law as a model. In February 1948, Zhadin wrote in the abstract of the presentation (unpublished):

Both natural and artificial changes in ecological characteristics of bodies of water are accompanied by a brief and fast relative repetition of the historical process of the faunal genesis, the process of transitioning from the prevalence of the primary aquatic fauna to that of the secondary aquatic fauna. We thus observe in the genesis of the fauna of continental waters the operation of a law essentially similar to Haeckel’s fundamental biogenetic law.Footnote 12

He spoke of the saltatory nature of faunal succession along the accumulation gradient.

Zhadin reformulated his productivity rule in line with these concepts, employing the terminology of the categories of organisms (primary and secondary aquatic). The gradual accumulation of organic matter initially leads to an increase in productivity, but once the level of accumulation passes a certain critical point with a rapid reduction in near-bottom oxygen content,

the fauna is abruptly transformed to the secondary aquatic fauna, with its quantitative development reaching significant values; however, if accumulation of organic substances continues… biological productivity decreases, also resulting in an abrupt transition from aquatic to terrestrial life.Footnote 13

He specified that there is a turning point in the proportion of primary and secondary bodies of water on sandy-silty bottoms (in a series from hard-rock to clay substrates). Zhadin concluded that the analysis of the faunal origin

has revealed the same patterns that had been established by Darwinism in the process of speciation. These facts show that the whole genesis process of the fauna of bodies of water occurs according to the strict laws of dialectical development.Footnote 14 This circumstance gives us the right to treat the branch of science we have introduced — genetic (or evolutionary) hydrobiology — as a constituent part of the science that studies the laws of life development, i.e. Darwinism.Footnote 15

These evolutionary studies, in Zhadin’s opinion, are essential for the long-term prediction of changes in the bodies of water, including those associated with the human impact that accelerates the events occurring in the bodies of water.

Unfortunately, Zhadin (like all biologists of that period) had to deal with the consequences of the VASKhNIL session of August 1948. Lysenko maintained that the nature of an organism can be changed by affecting its physiological properties: “The source, the material, from which the organisms create or build themselves, are the environmental conditions, the food in the general sense of the word” (1946a, b, p. 337). The “food,” as it was understood by Lysenko, included abiotic conditions such as sunlight, temperature, air humidity, chemical elements of soils and nutrition, and atmospheric gases. But nutrition in the strict sense also held a place of special importance in Lysenko’s system of views: “Nutrition is also a mentor” (Prezent 1948, p. 52). All these influences tend to “shake down” the inheritance, and further modifications caused by the environment can become inheritable. From this standpoint, the organism is “the mold” created by the environment.

After the VASKhNIL session, Zhadin devoted a special publication to the problems of applying the ideas of Lysenkoism to hydrobiology (Zhadin 1949). In this publication, Zhadin used Lysenko’s ideas on the mechanism of origin of new species by gradual accumulation of the modifying environmental changes (“persisting variability”). Zhadin put forward the idea that “the speciation-inducing role of changes in the aquatic environment” (1949, p. 200) should be studied by hydrobiologists. Zhadin stated that the speciation rate can be estimated by “examining the manifestation of the so-called persisting variability in bodies of water, which at a certain stage changes abruptly into an entirely new inheritable form” (1949, p. 201). Changes occurring in the bodies of water themselves reinforce the importance of such studies: “These changes happen so fast that many bodies of water turn into land before our very eyes” (Zhadin 1949, p. 200). This situation is explained by intensification of soil erosion caused by economic development in the catchments.

Zhadin then proceeded to the analysis of biocoenoses (communities). He distinguished two significant traits in the organization of the biocoenosis. First, the biocoenosis is formed in a regular fashion over the course of historical processes in a water body. According to Zhadin, the biocoenosis is “a historically established, sustainable grouping of organisms” (1949, p. 203), formed by adaptation of organisms to the environmental conditions. Second, the prime importance in the system of relationships in the biocoenosis is given to the external physico-chemical conditions (current velocity and characteristics of the substrate), rather than to the interspecific relations (in particular, competition). For this reason, both biocoenosis and the water body in general (as a biocoenosis of higher order, i.e. biome) cannot be compared to an organism of higher order, which has some kind of harmonic organization.

Once again, Zhadin approached the question of integrity of the water body from the historical position. He pointed out that “the unity of a water body is not determined by some kind of vitalistic ‘self-organisation,’ but rather by its history and by the processes in the cycles of matter” (Zhadin 1949, p. 205), which tie together all parts of the water body and its biocoenoses. Zhadin proposed an additional argument for the dismissal of likening of the water body to a superorganism, because from a standpoint of the evolution of a water body, its different parts can be inhabited by communities of different ages.Footnote 16 In Zhadin’s opinion, biocoenoses should be given names that “reflect the unity of the biocoenosis and the environment.” In so doing, he called for the removal of the excessive foreign terminology, which he likened to “a weed of sycophancy towards the corrupt foreign influence” (Zhadin 1949, p. 204). Zhadin also criticized the use of quantitative indices in studying biocoenoses such as prevalence, species abundance, etc.

Compared to his earlier works, Zhadin placed even more emphasis on his utilitarian approach to the production studies. As he wrote, “we, as I think, must follow the same path as agriculture and forest husbandry, i.e. to judge productivity based on the magnitude of production, the amount of produce that the human takes from a unit area per unit time” (Zhadin 1949, p. 207). He proposed a purely economic division of production into “useful” and “harmful.” According to Zhadin, it is important “whether the water body produces a tonne of organic matter in the form of larvae of malaria mosquitos or the same tonne in the form of tendipedid larvae eaten by fish” (1949, p. 208). And the ton of crucian carps is incomparable to the ton of sturgeons or salmons. It should be noted that all this phraseology is very similar to the statements given by Prezent in 1934 (see above).

In his criticism of “the formal bottle method of Vinberg,” Zhadin differentiated between the intensity of photosynthesis, which is measured by the bottle method, and the biomass of economically valuable organisms. Zhadin stressed that the whole production process of organic matter cannot be reduced only to the photosynthesis of the phytoplankton. One should take into account the substances flushed from the catchment and stirred up from the bottom. For instance, if the inflow of such substances is put under control, one would be able to influence the growth rates of hydrobionts.

In this respect, Zhadin supported the creation of protective tree belts,Footnote 17 designed to prevent soil erosion. He sought to encourage “a course of purposeful biological production” as opposed to “spontaneous production.” Methods of such purposeful production (including acclimatization) are part of the toolbox of Soviet biology. Zhadin said they will allow us “to give the production, which is useful for humans, and to suppress one, which is harmful” (1949, p. 209). In matters of acclimatization, Zhadin was sufficiently prudent. He suggested that organisms should be introduced in the bodies of water that have free habitats, vacated as a result of certain adverse factors (including human activity). The biological characteristics of the species to be introduced should match the conditions of the water body.

In general, Zhadin thought that the problem of productivity was central for hydrobiology at the new phase of Soviet biology. He also believed that it was necessary to use direct methods of studying the processes of photosynthesis, feeding, reproduction and growth of organisms, and the phenomena of cycles of matter. By stating this, Zhadin seems to somewhat contradict his own negation of Vinberg’s “light and dark bottle method,” which involved the direct measurement of oxygen generated by photosynthetic activity of the phytoplankton. The examination of a draft version of Zhadin’s paper explains this contradiction. In the draft,Footnote 18 Zhadin crossed out a whole paragraph devoted to the “bottle” method, which Zhadin categorized as an indirect method of studying production. Seeing Vinberg’s approach as “oversimplified” and “formalistic,” he criticized it not only on ideological grounds, but also using logical reasoning. Zhadin wrote:

While rejecting indirect methods of studying biological productivity for their inefficiency and even outright harmfulness at this point in the development of hydrobiology, I admit the complete application of this kind of methods at that stage in the development of science, when the exact qualitative and quantitative understanding of the relationship between life activity of organisms and the reflection of this activity in the chemistry of the water body will be achieved.… Formalists, on the other hand, use indirect methods of studying biological productivity seeking to understand a complex object /producing organism/ from a simple one /change in water chemistry/, but in actual fact produce only an approximate idea about the amount of some featureless organic matter /that could be useful, but could also be harmful/.Footnote 19

When Zhadin mentioned the “featureless” matter, he repeated the words of Prezent in 1934 during the discussion on the goals and the content of ecology.

On the surface, Zhadin’s paper abounded with ideological slogans. Zhadin called Lysenko’s presentation at the VASKhNIL session “a powerful invocation to the Soviet creative Darwinism, to the revision of ideological positions in various fields of biology…. The time has now come for this kind of revision in hydrobiology” (1949, pp. 197–198). Therefore, at this time, “any ideological indecision, any subjugation to corrupt bourgeois beliefs, any veneration of foreign ideas can be treated only as a step towards defecting to the enemy camp” (1949, p. 197).

This kind of rhetoric was very common in scientific works of that period. Zhadin generally accepted the principles of Lysenkoisms biology, but, in many respects, he also repeated and supported the views on productivity detailed in his 1940 monograph. He advocated for the unity of an organism and the environment determined by their historical relationships and defended his view on a water body as an integrated system of the biocoenoses tied together by their common genesis. Interestingly, in this major publication, Zhadin almost did not elaborate on the genetic approach to the bodies of water that he had recently proposed. Instead, he suggested the consideration of the role of the environment in speciation, much in keeping with the beliefs of Lysenkoists.

In spite of the accentuated economic utilitarianism of this paper and support of the measures for transformation of nature that had been implemented in the country by high authorities, Zhadin continued to give entirely rational recommendations on acclimatization. The text of the paper gives an impression of a very sensible and well-reasoned program. Zhadin refrained from abandoning his previous views and did not regret his convictions. As mentioned above, he only allowed a modicum of self-criticism with respect to his nonessential publications on variation in mollusks.

In Zhadin’s personal archive, there is a draft note (typed on a half of a page, without a date or a title) in which he wrote:

The osmotic pressure of the internal milieu seems to protect the animals from the direct effect of the environment. However, the processes of feeding and respiration open up the possibility to influence the organisms and change their biochemical properties. T.D. Lysenko is quite right in this respect when he speaks that the impact of the external environment on the organism is mediated by metabolism.Footnote 20

These words, written “for the drawer,” give an illustration of Zhadin’s sincere desire to find a rational grain in Lysenko’s words rather than simply to cite his works. The key phrase for this conclusion is the collocation “quite right.” I believe that in the general context, these words do not indicate the unconditional acknowledgement by Zhadin of the academician’s correctness, but rather an attempt to understand and adapt his views.

The Fate of Zhadin’s Theory and Its Place in Hydrobiology

Since Zhadin’s theory was published in the prewar years, one might expect that it would not have received an immediate response, but as early as 1945 the renowned marine biologist Vladimir A. Vodyanitskiy published a paper devoted specifically to Zhadin’s theory. Vodyanitskiy did not believe that Zhadin’s theory should be called a theory. He did not see in it any aspects explaining known facts or new steps towards building the concept of the production process. As the common saying goes: “everything new in it is incorrect and everything correct is not new.” According to Vodyanitskiy, the only positive aspect of the theory is that Zhadin’s principles allow him “to express in terse words what used to be articulated in more verbose terms” (1945, p. 75). So what approach did Vodyanitskiy propose to develop a theory of bioproductivity? He stressed the need to study the biology of “index dominant species.” In his opinion, this was exactly what constitutes “the correct scientific approach to understanding the productivity of bodies of water” (1945, p. 75). The only question that remained unanswered is how our knowledge of the biology of “index dominant species” all by itself can help in formulating the productivity theory.

Ivlev (1945) devoted a few words to Zhadin’s ideas on the productivity problem. In his opinion, Zhadin “substituted ‘productivity’ with the notion of fauna richness, i.e. its biomass and species diversity” (Ivlev 1945, p. 101). I believe that Ivlev failed to understand the meaning of Zhadin’s term “bioecological provision,” which was specifically meant to denote species diversity. This diversity is only a factor of productivity. The productivity itself is manifested in a certain level of biomass production.

In another part of his paper, Ivlev asserted that Zhadin understood productivity in the opposite way: as organic matter represented by organisms or as the “product” according to Karzinkin. But, in fact, Zhadin clearly stated that production is an increase in biomass. The product according to Karzinkin is an abstract concept, and Zhadin only stressed that “we should not forget about product,” “as was correctly reminded by Karzinkin” (Zhadin 1940, p. 943). Therefore, Ivlev’s remarks should not be viewed as bearing on the subject matter of the theory, but as a reflection of his misinterpretation of Zhadin’s ideas.

The comparison of Ivlev’s and Zhadin’s ideas reveals two major viewpoints on productivity. Zhadin (1947) criticized Ivlev’s ideas on productivity in terms of energy sources in the water body. Zhadin maintained that solar radiation cannot be viewed as the only energy source without considering the products of soil erosion, because the substrate for bacteria is in fact provided by this terrigenous material. Zhadin thought that Ivlev denied the dynamism in a water body, and thus he essentially “separated” it from the landscape. Therefore, Zhadin regarded the catchment and the processes of erosion and accumulation as the primary energy sources. In contrast, for Ivlev and Vinberg, the foundations for all energy-generating processes in a body of water were solar radiation and algal photosynthesis.

In later publications of Soviet hydrobiologists, Zhadin’s theory and his ideas about production were almost never mentioned or discussed. The only exception is a monograph by Skadovsky (1955). Skadovsky emphasized the need to study changes of biocoenoses in bodies of water and praised the accumulation principle proposed by Zhadin. Skadovsky also listed Zhadin’s works among those that laid the theoretical foundation for the concept of biological productivity of bodies of water. During a major discussion of the essence and goals of hydrobiology held in 1951 “in the wake of” the VASKhNIL session, there were only a few references to Zhadin’s publications or ideas (see Pavlovskiy 1953). The participants in the discussion mentioned only his works on water reservoirs and discussed the definition of hydrobiology that he suggested. He himself, for unknown reasons, did not participate in the discussion on productivity problems at a specially designated section of the All-Union Fisheries Conference in December 1951,Footnote 21 and he published only several commentaries on these topics.

Nevertheless, one of the active participants of this discussion, Karzinkin, devoted some words to Zhadin’s theory (Karzinkin 1952). Like Vodyanitskiy, he did not believe that Zhadin’s theory was a real theory and criticized it in a rather erratic and inconsistent fashion. To give just one example, Karzinkin asserted that Zhadin spoke of biomass in general, but ignored the problem of producing economically valuable organisms. However, further on, Karzinkin noted that a positive aspect of Zhadin’s “theory” was in fact that it drew attention to the possibility of acclimatization of economically valuable organisms.

In the 1950s and 1960s, Zhadin did not return to his theory of biological productivity of bodies of water or to the concepts of “accumulation” and “bioecological provision.” At the same time, he continued to conduct productivity studies. In the early 1950s, he initiated research on the theory and practice of fertilization of fish-rearing ponds. Later on, he became interested in the use of radioactive isotopes in hydrobiology, most notably in measuring the primary production.

In a paper by the marine biologist Evpraksiya F. Gurianova (1968), special attention was paid to Zhadin’s 1940 monograph and the productivity theory, which she called the ABEP (accumulation and biological and ecological provision) theory. Gurianova amended this theory, differentiating the biological provision (“the totality of living populations”) and ecological provision as the sum total of all conditions in the water body. Gurianova noted that Zhadin’s theory “provides hydrobiologists with the most fruitful idea of creating a classification of natural bodies of water” (1968, p. 11). This is essentially the only reference to the theory in hydrobiological publications. At the same time, Gurianova admitted (in the context of the correctness of Zhadin’s ideas) that Zhadin’s views had been criticized by “some researchers.”

Zhadin’s views on production have found no response in the modern theory of aquatic ecosystems for a seemingly obvious reason: the seminal work of Lindeman (1942) was published almost simultaneously with Zhadin’s theory. Lindeman’s work marked a true breakthrough in ecology. After the Second World War, ecology and, in particular, aquatic ecology (hydrobiology) had risen to a whole new ecosystemic and quantitative level of research. The study of quantitative patterns in energy transport along the trophic chains (Taylor 1988) has become and still is the groundwork of the modern productivity studies. The productivity of an ecosystem is no longer necessarily associated with the yield of the economically valuable biomass.

For these reasons, the head of the Russian scientific school of productivity-based hydrobiology, RAS academician Alexander F. Alimov, who was Zhadin’s disciple in malacology, does not consider his mentor to be a member of the productivity branch of hydrobiology (Rizhinashvili 2017a). Alimov believes that only the scientific school created by Vinberg can be regarded as productivity-based. Zhadin’s hydrobiological works are now mentioned only occasionally and only in the Russian literature, although his malacological publications are commonly cited in the international scientific literature even today.

Nonetheless, Zhadin’s theory has an especially promising side: the notion of a river as an integrated whole. The concept of a lake as an integrated and relatively closed ecosystem is readily understood, and this was the reason why the theoretical ideas of Lindeman, who worked on lakes, initially appeared so well argued and complete. The task of studying patterns in the cycles of matter and energy in rivers is, for obvious reasons, much more difficult. In Zhadin’s view, the process that unites the parts of the river into a single whole is accumulation, which in turn is a function of the processes of the cycles of matter in the catchment. Rivers can be subdivided into sections according to the extent and type of accumulation. The idea of the river’s non-uniformity, the gradient nature of its conditions, and corresponding succession of communities were established almost half a century later as the foundation for the concept of the riverine continuum (Vannote et al. 1980). The authors of this concept, however, have never mentioned Zhadin’s works. On the other hand, Zhadin has recently been named a forerunner of the ideas closely related to this concept (Protasov 2008).

The question that remains is why Zhadin’s highly promising ideas (if not the productivity theory itself) happened to be “thrown away.” Was it the ideological component that proved to be an impediment for the acceptance of these ideas?

Traces of Lysenkoism (by Way of Conclusion)

In general, hydrobiologists were among those biologists who most persistently opposed the influence of Lysenko and Prezent and the plans of transformation of nature. This can be seen from Zhadin’s own works. Later in the early 1950s, during a well-known discussion on biological productivity, hydrobiologists showed their firm commitment and managed to save the integrity of hydrobiological research (Zaika 2002). But, needless to say, they were also forced to adapt to the ideology.

As they tried to adapt to the Soviet regime, many scientists (certainly not only hydrobiologists) found that the dialectical way of thinking imposed by governmental officials could be turned into a useful tool of intellectual inquiry (Graham 1987; Weiner 1988). Zhadin’s theory is an excellent example of the dialectization of biology and of this style of thinking. Zhadin demonstrated how quantity (input intensity of organic compounds) can be transformed into quality (replacement of the primary aquatic with the secondary aquatic fauna).

One of the central elements of Stalin’s science was its close ties with practice (Roll-Hansen 2008). This practice-oriented approach, however, was not uncommon even before the October Revolution, because in pre-revolutionary times limnological studies were often arranged and carried out with the participation of regional authorities that were interested in making an inventory of local resources. Scientists all over the world were also always interested in such coordination and were enthusiastic about helping the practical life of the country (Kolchinsky 2014; Schwarz and Jax 2011). As discussed above, Zhadin never shied away from the practical and economic challenges. In the atmosphere of an extremely utilitarian science of the time, he managed to choose a direction for his research that was oriented towards practical ends. This approach eventually found its expression in his productivity theory. Zhadin saw the biomass of an economically valuable product as the ultimate goal of the production process, and he showed the way to find the optimum product output and methods of controlling it.

Were Lysenkoism ideas congenial to Zhadin’s own style of thinking and his studies? Alimov noted that Zhadin had a “riverine” style of thinking (Rizhinashvili 2017a), and a river is rather dynamic in its hydrological characteristics. This dynamism raises abiotic conditions to a place of prime importance in the life of hydrobionts. For this reason, Zhadin had almost no interest in analyzing interspecific and especially intraspecific relationships. This approach, which stresses the primacy of abiotic conditions, is well in line with Lysenkoist ideas.

Finally, Zhadin widely proclaimed the dismissal of mathematical reasoning in describing biological processes. His theory lacks mathematical apparatus even in its simplest form. It is still unclear whether it was Zhadin’s sincere position or his compliance to the ideologically-driven rejection of mathematics. In his earlier works, too, Zhadin almost never used mathematical methods to any significant extent.Footnote 22 In any event, this defining characteristic of Zhadin’s theory could indeed have prevented its adequate reception. But it stands in full agreement with the Lysenkoist ideology.

I believe that Zhadin was sincere in seeking to apply Lysenkoism beliefs to his research program. This sincerity is underscored by the fact that many of those beliefs were in accord with his own ideas and notions. These aspects of Zhadin’s style of thinking were determined by his professional experience. This, in my opinion, constitutes the interrelation between the cognitive and ideological components in Zhadin’s theory. The reasons why this theory was forgotten are not entirely clear. They can lie both in the ideological aspects and in the particular character of the theory, which explained production processes specifically for riverine ecosystems.

This analysis of Zhadin’s theory, with the aim of identifying social and cognitive components of scientific knowledge, has demonstrated the complexity and contradictory nature of the scientist’s work under the totalitarian regime. In our study, the scientist’s own views and ideologically influenced (imposed) elements proved difficult to differentiate. In this case, the behavioral pattern of “protective coloration” proposed by Weiner (1988) proved to be very efficient. I should also stress Zhadin’s lack of criticism towards official authorities in his everyday life. It was probably this attitude that saved him; he was fortunately never arrested and, so far as is known, never suffered even the slightest persecution from the government. Zhadin, however, seemed to understand the negative processes in Soviet biology of that period. In his memoir My Path to Hydrobiology, he mentioned one well-recognized scientist (botanist Sergey S. StankovFootnote 23), who later “played a certain role in the normalization of situation in biological science” (Zhadin 1991, p. 41).

In my view, the present analysis highlights a rather unconventional aspect of Lysenkoism that can be called its rational grain. While in genetics Lysenko’s and Prezent’s ideas were unquestionably pseudoscientific, the situation in ecology was more complicated. Lysenkoism was represented in ecology by reasonably sensible scientific principles pushed to the extreme for the sake of ideology, and by the commitment to the practical realization of the study results. These principles include the unity of an organism and its environment, irreducibility of the processes in supra-organismal systems to the functions of an organism, characteristic properties of the living matter, importance of accounting for regular patterns in metabolism, combination of an experiment and observation, and importance of understanding of the phenomena’s dynamics. It is important to note that these principles were known in science before Lysenkoists and without them.Footnote 24

For this reason, it is quite understandable that in dealing with the sufficiently professional studies of that period, it is difficult to distinguish between the common sense of the author and the ideas borrowed in one form or another from Lysenkoism. In fact, this is one of the reasons why Lysenko’s ideas could have been appealing for genuine scientists, who were unaware of their ideological underpinnings.Footnote 25 This has already been noted by several authors, and it also explains the phenomenon of the high efficiency and productivity of science in the USSR despite several dozen years of severe pressure of ideology, a fact which was noted by Graham (1987). Scientists learned to use the imposed ideology for their own benefit by modifying it in a creative way. It does not imply, of course, that Lysenkoism is a truly scientific discipline. Using seemingly correct ideas, the adepts of Lysenko and Prezent (with the support of governmental officials) were forcibly imposing their principles on ecology and hydrobiology, without paying much consideration to the specifics of particular scientific problems. In any case, the question of the cognitive foundations of Lysenkoism in ecology and hydrobiology is still a long way from being fully explored.

Born in the depths of ideologized science, Zhadin’s theory of biological productivity of bodies of water must not only find its place in the history of hydrobiology, but should also be critically analyzed by modern-day science. My results call for a more careful and deep evaluation of the scientific works of the period of the personality cult in the USSR. The “Lysenko affair” in different branches of biology other than genetics is more complicated and needs appropriate examination. This approach will also be useful in analyzing modern-day pseudoscientific ideas and theories; otherwise “Lysenko’s ghost” will unfortunately continue to return over and over again.