Water Resources, Vol. 32, No. 3, 2005, pp. 305313. Translated from Vodnye Resursy, Vol. 32, No.

3, 2005, pp. 337346. Original Russian Text Copyright 2005 by Ostroumov.

WATER QUALITY AND PROTECTION: ENVIRONMENTAL ASPECTS

On Some Issues of Maintaining Water Quality and Self-Purification

S. A. Ostroumov
Moscow State University, Leninskie Gory, Moscow, 119992 Russia
Received August, 4, 2003

AbstractGeneralizations presented in this paper represent, in systematized form, the basic elements of the qualitative theory of water self-purication in freshwater and marine ecosystems. Recommendations are given for maintaining water quality and sustainable development of water resources. Results of experimental studies of the effect exerted by Triton X-100 and OMO synthetic detergent on mollusks Unio tumidus.

INTRODUCTION The role of self-purication increases due to the deterioration of natural water quality [5, 13, 29] and increased anthropogenic load on water bodies and streams [4, 9, 13, 21, 22, 26, 29]. The self-purication of aquatic ecosystems and water quality formation is controlled by many [13, 613, 1519, 21, 22, 24, 33, 35]. The objective of this study is to systematize the knowledge about the polyfunctional role of aquatic biota (aquatic animals) in the self-purication of water bodies and streams and briey present the qualitative theory of the self-purication mechanism of aquatic ecosystems. The synthesis and structurization of material was made at the conceptual level without detailed reviews of works. MAJOR PROCESSES CONTRIBUTING TO WATER SELF-PURIFICATION IN AQUATIC ECOSYSTEMS The formation of water quality and its purication in aquatic ecosystems is governed by physical, chemical [24], and biotic [13, 613, 1519, 21, 22, 24, 33, 35] processes (Table 1). The physical and chemical processes of water selfpurication are often controlled by biological factors or strongly dependent on them. Thus, the redox state of the aquatic environment, which forms with the participation of H2O2 released by microalgae in the light [22, 24], is of importance for a decrease in the toxic effect of some pollutants. The amount of H2O2 released into the aquatic environment was estimated at 105 mol/(l day). The concentration of H2O2 in the Volga was found to equal up to 106105 mol/l, which was supported by measurements made by E.V. Shtamm and other authors [22, 24]. An important process is gravitational sedimentation of suspended particles both of biotic and abiotic nature. The sedimentation of phytoplankton sedimentation depends on water temperature T. It is equal to 0.31.5,

0.41.7, and 0.42.0 m/day at T = 15, 20, and 25C, respectively. According to our data, the sedimentation velocity of Lymnaea stagnalis pellets varies from 0.6 to 1.4 cm/s with a mean value of 0.82 cm/s (at T = 22 24) [22]. Experiments with traps for suspended particles showed that suspended matter precipitates onto the bed of the Moskva River with a mean rate of 2.3 mg per 1 cm2 of the bed surface per day, that is, 23.1 g per 1 m2 of the bed surface per day; the proportion of Corg in these sediments is 64.5% [34]. Organic matter oxidation and water ltration by aquatic animals are among the biotic processes contributing to water purication. The overall oxidation of organic matter by the entire community can be expressed either in absolute or in relative units, for example, as the ratio of energy expenditure to the exchange (total respiration R) by aquatic animals to their total biomass B. This ratio (R/B)e is referred to as Schrodinger ratio. The subscript e is introduced to show that the estimation is made for the ecosystem as a whole. In the water bodies where primary production exceeds the total respiration of the community, this ratio averages 2.996.1 [2], but it can be even greater in some water bodies. For example, the Schrodinger ratio is 17.0 in Lake Lyubevoe in Leningrad province and 33.8 in Lake Zun-Torei east of Lake Baikal [2]. It is believed that the primary production in these lakes is much less than the total respiration and a large amount of organic matter delivered from outside is oxidized here. Many aquatic animals contribute to organic matter oxidation, but particular role in this oxidation belongs to bacteria [19]. The total population of heterotrophic bacterioplankton in the Mozhaisk Reservoir in June and July amounted to (1.365.9) 109 (samples were taken at a depth of 0.11 m), and the population of hydrocarbon-oxidizing bacteria was (0.45) 106 cell/ml [7]. The rate of water ltration by some aquatic animals (sea squirts, barnacles, pearlweeds, echinoderms,

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Table 1. Some factors and processes of water self-purification [13, 22]. (DOM is dissolved organic matter) Factors and processes of water self-purification Dissolution and dilution Comments

Physical and physicochemical processes Mechanical transfer of water masses can depend on the abundance of macrophytes; Transfer to the shore The same Transfer to neighboring water bodies and streams The same Sorption by suspended particles with subsequent sedimentation Depends on the concentration of suspended particles of biogenous nature in water Sorption by bottom sediments Depends of the concentration of organic matter of biogenous nature in bottom sediments Evaporation May depend on the surface film, the properties of which depend on the composition of DOM Chemical processes Hysrolysis May depend on pH which changes during planktonic photosynthesis Photochemical transformations Depends on the concentration of photosensitizers of biogenous nature and water transparency, which, in its turn, depends on plankton Redoxcatalytic transformations Concentration of catalytically active form of metal ion depends on the pH of the environment, which, in its turn, depends on the photosynthetic activity of plankton Transformations involving free radicals Depend on the concentration of H2O2 that forms with the participation of aquatic animals (photosensitized transformation of DOM, release by microalgae) Decrease in the toxicity of pollutants as a result of binding Depends on DOM of biogenous nature; the role of humic with DOM acids is possible Chemical oxidation of pollutant with the participation of oxygen Depends on O2 release into water during photosynthesis Biological processes Release of O2 taking part in many reactions of pollutant The same oxidation Porption and accumulation of pollutants and nutrients by Depends on the population and activity of aquatic animals aquatic animals Biotransformation (redoxreactions, destruction, conjugation) The same Extracellular enzymatic transformation of pollutants Depends on the population and activity of aquatic animals Removal of suspended particles and pollutants from water Is inhibited under the effect of some pollutants (surfactants, column as a result of water filtration by aquatic animals synthetic detergents) [33] Removal of pollutants from water as a result of sorption Pellet formation is reduced when feeding of aquatic animals by pellets excreted by aquatic animals is inhibited Release into water of organic matter, which can be used Organic matter release by phytoplankton was recorded. by bacteria or exert a regulatory effect on them Higher aquatic plants also release exometabolites Release into water of organic matter that serves as a photosensi- Depends on the population and functional activity of aquatic tizer of pollutant photolysis or predecessors of photosensitizers animals Release into water of organic matter (or predecessors of such The same matter), which will bind with pollutants with the formation of less toxic complexes Release into water of organic matter (or predecessors of such Microalgae release hydrogen peroxide matter), which participate in free-radical and redoxcatalytic mechanisms of pollutant destruction Release into water of vitamins, which are necessary for the Water of lakes and pools contains, g/l: vitamin B12 (0.0010.85), vital activity of some aquatic animals, participating in thiamine (0.00112), biotin (0.00010.1), niacin (up to 3.3), self-purification of aquatic ecosystems pantothenic acid (up to 0.26), and others vitamins [35] Release or inactivation of part of bacteria, including pathogenic Takes place at filtration activity of aquatic animals; can take strains place under the effect of metabolites of aquatic animals Prevention or hampering of the release of nutrients and pollut- Depends on the population and functional activity of benthos ants from bottom sediments into water; accumulation and binding of nutrients and pollutants by benthic organisms Biotransformation and sorption of pollutants in soil during Depends on the soil biocenoses soil watering by wastewaters Regulation of the population and activity of organisms partic- Depends on the preservation of intact community ipating in the processes of water purification as a result of organism-to-organism interaction
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Table 2. Characteristics of filtration activity of some aquatic animals [22]. Here and in Tables 3, 6, dash means no data available Animal species Keratella quadrata Filinia sp. Epiphanes brachionus Bosmina longirosris Cladoceras, various species Copepods, various species Unio pictorum, length 91100 mm Dreissena polymorpha Plumatella fungosa Cyprinotus carolinensis
* mg. ** ml/h. *** ml/g per 1 h.

Wet weight (mean), W, g 0.001020* 0.000855* 0.001800* 0.016000* 72.400(with the shell) 0.065 (dry weight without shell)

Water filtration by one animal F, ml/day 0.25 0.40 0.29 6.55 2040 (rarely to 130) 24 (sometimes to 27) 1238** 500700** ~4

Specific filtration activity, ml/mg of wet weight per 1 day, F/W 246.0 468.0 161.0 409.3 17*** 2.2 ml per 1 mg dry weight of the colony per hour

bivalves, gastropods, polychaetes, and sponges) commonly amount to 19 l/h per 1 g of de-ashed dry mass of their body [21, 22]. The dependence of ltration rate FR, l/h, on the mass of the aquatic animal DW, g, can be described by the power function [1, 22] FR = a DWb, (1) were DW is the dry weight of soft tissues, g. The values of coefcient a for some mollusk species vary from 6.8 to 11.6, and those of coefcient b lie between 0.66 and 0.92 [22]. The rate of water ltration by ve mollusk species converted to the area of their gills is about 1.21.9 ml/min per 1 cm2 [22]. The total rate of water ltration by populations of macroinvertebrates (mollusks, sea squirts, polychaetes) was estimated at 110 m3 per 1 m2 of the bed of the aquatic ecosystem per 1 day [13, 22]. Additional data on the ltration activity of aquatic animals is given in Tables 2 and 3. THE MAJOR COMPONENTS OF THE SELF-PURIFICATION MECHANISM OF AQUATIC ECOSYSTEMS The hydrobiological self-purication mechanism of aquatic ecosystems incorporates three types of components [21, 22]: ltration activity of organisms (lters) [30]; the mechanisms of transfer of chemicals from one ecological compartment into another (from one medium into another); splitting pollutant molecules. The processes and aquatic animals that serve as lters [21, 22, 30]: invertebrate lter-feeders [1, 25]; coastal macrophytes, which retain some nutrients and pollutants delivered into water from neighboring areas; benthos, which retains and absorbs part of nutrients and pollutants at the waterbottom sediment interface;
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microorganisms sorbed on particulates that move within water column due to sedimentation of particles under the effect of gravity; as a result, the water mass and microorganisms moves relative to one another, which is equivalent to the situation when water moves through a porous substrate with microorganisms attached to walls [30]. Precipitation of a suspended particle, that is, its movement with respect to water, enhances O2 exchange between the sorbed bacteria and the aquatic medium [6]. The processes and aquatic animals that serve as pumps [21, 22]: facilitating the transfer of part of pollutant from the water column into bottom sediments (e.g., sedimentation, sorption); transferring part of pollutant from the water column into the atmosphere (evaporation); transferring part of nutrients from water into the territory of neighboring terrestrial ecosystems because of the emergence of imago of aquatic insects; transfer of part of nutrients from water onto the territory of neighboring terrestrial ecosystems through sh-eating birds, which withdraw sh biomass from water. The processes and aquatic animals that serve as mills and split pollutants [21, 22]: intracellular enzymatic processes; processes catalyzed by extracellular enzymes; decomposition of pollutants by photolysis: photochemical processes, sensitized by nutrients; destruction of pollutants in the free-radical processes with the participation of biogenous ligands [24]. ENERGY SOURCES FOR BIOTIC SELF-PURIFICATION MECHANISMS OF AQUATIC ECOSYSTEMS The processes of biotic self-purication of water take energy from the following sources: photosynthesis, oxidation of autochthonous and allochthonous organic matter; other redox reactions. Thus, practically

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Table 3. Removal of suspension from water through the filtration activity of freshwater mollusks Unio tumidus. Vessel A contains 10 mollusks with a mean weight (wet weight with the shell) of 18.99 g (12.9725.73 g) and a total biomass of 189.94 g. Mollusks were collected in the Moskva R. (Odintsovo district). Vessel B did non contain mollusks (check experiment). The volume of water in each vessel was 500 ml, T = 19C. Suspension of Saccharomyces cerevisiae Time from the beginning of the experiment, min 10 20 30 40 60 70 85 Optical density at 500 nm (optical path of 10 mm) A 0.660 0.585 0.525 0.442 0.295 0.242 0.184 B 0.713 0.715 0.676 0.666 0.598 0.581 A/B, % 92.57 81.82 77.66 66.37 49.33 41.65 Water turbidity in vessels does not differ Pellet lumps appear in vessel A; the medium is more transparent than in vessel B Pellet lumps are clearly seen in vessel A; water in the vessel with mollusks is notably more transparent than that in vessel B The same " " " Visual observations

all available energy sources are used. A part of the energy is supplied through oxidation of the components (dissolved and particulate organic matter) which the system gets rid of [32]. Water self-purication is commonly associated with organic matter oxidation by aerobic microorganisms. Equally important are anaerobic processes which receive energy from the transfer of electrons to acceptors other than oxygen. Anaerobic energetics feeds the metabolism of microorganisms of methanogenic community (decomposition of organic matter results in the production of 2S, 2, and 4), and anoxygenic pho2 totrophic community (with the formation of S O 4 , H2S, H2, and CH4) [15]. The products produced by organisms of these communities are used as oxidation substrates by organisms of other communities, including the organisms that form the group referred to as a bacterial oxidation lter. The latter lter functions under aerobic conditions and oxidizes H2 (knellgasbacteria), CH4 (methanotrophs), NH3 (nitrifiers), H2S (thiobacteria), thiosulfate (thionic bacteria) [6]. For example, in Lake Mirror (USA), 19.1 g C/m2 of lake surface is oxidized annually due to phytoplankton respiration, 12.0 due to zooplankton respiration, 1.0 due to macrophytes, 1.16 due to attached plants, 2.8 due to benthic invertebrates, and 0.2 g C/m2 due to sh. Oxidation by bacteria in bottom sediments and by bacterioplankton accounts for 17.3 and 4.9 g C/m2 of lake surface [35]. CONTRIBUTION OF MAJOR TAXA TO SELF-PURIFICATION PROCESSES IN AQUATIC ECOSYSTEMS Practically all major groups of organisms contribute to self-purication of aquatic ecosystems and formation of water quality [6, 914, 1622, 27, 3133, 35].

A signicant role belongs to microorganisms (Table 4) [3, 6, 7, 25], phytoplankton [21, 22], higher plants [21, 22], protozoa [27], zooplankton [21, 22, 35], benthic invertebrates [21, 22, 35], and sh. All these groups contribute largely to the self-purication of aquatic ecosystems, each group taking part in several processes. Microbial processes of water self-purication are associated basically with the activity of heterotrophic aerobic bacteria; however, representatives of practically all major bacterial groups (>30) participate in the key processes of organic matter destruction and selfpurication of water bodies [6]. It is worth mentioning that the microorganisms participating in the destruction of biopolymers and in water self-purication system feature wide taxonomic diversity [6]. An important role in organic matter destruction and self-purication of aquatic ecosystems belongs also to eucaryotic microorganisms (protests), in particular, Diplomonadea, kinetoplastides and euglenes, ameboagellates, dinoagellates, infusoria, heteroagellates, cryptomonades, choanoagellates, and chitrids [6]. An important process of water self purication is water ltration by organisms of many taxa [1, 8, 21, 22, 25]. A detailed list of taxa, including planktonic and benthic lterers in aquatic ecosystems, is given in [16]. The contributions of different groups of organisms to C removal from water of eutrophic Lake Esrum (Denmark) in percent of the total C withdrawn from water are as follows: 24.4% by respiration of producers, 20.9% by bacterial respiration, 30.7% by respiration of consumers, 4.5% (appears to be determined not completely) by the respiration of microorganisms in sediments, 0.14% by the emergence of aquatic insects [35].
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Table 4. Role of microbial communities in the self-purification of aquatic ecosystems (part of examples is given; some bacterial groups are involved in the functioning of communities of several types [6, 22] and others; VFA are volatile fatty acids) Type of community Methanogenic community Sulfidogenic community Anoxigenic phototrophic community (oxidation anoxic phototrophic filter) Oxidation aerobic filter (gasotophs) Substate being transformed (consumed) Decompose biopolymers, form acetate, VFA, hydrogen, CH4 Decompose biopolymers, produce VFA, hydrogen, acetate, lactate, H2S, CH4 Consume VFA, hydrogen, lactate, acetate, H2S; produce 2 hydrogen, acetate, H2S, CH4, SO 4 Consume (oxidize) hydrogen, CH4, VFA, thiosulfate, H2S

Virtually all groups of organisms belonging to procaryotes and eucaryotes are necessary for water selfpurication. THE RELIABILITY OF WATER SELF-PURIFICATION SYSTEM The reliability of a technical system often relies on the presence of back-up components. Analysis of aquatic ecosystems shows a similar principle to govern their functioning. For example, the ltration activity of aquatic animals is doubled so that it is implemented by two large groups of organisms, i.e., plankton and benthos. Both groups lter water with a considerable rate [1, 8, 13, 25]. Additionally, benthos duplicates the activity of planktonic organisms permanently inhabiting the pelagic zone, since the larvae of many benthic lterers follow the planktonic pattern. Plankton incorporates two large groups of many-celled invertebrate lterers, i.e., crustaceans [25] and rotifers [8], both of which implement water ltration. One more large group of organisms (protozoa), which has somewhat different type of nutrition, also duplicates the ltration activity of many-celled lterers (crustaceans and rotifers). The fermentative decomposition of pollutants is partially duplicated by the activity of bacteria and fungi. Almost all aquatic animals, which are, in some way or another, capable of consuming and oxidizing dissolved organic matter, perform this function. Self-regulation of biota is an important component of the reliability of water self-purication mechanism. The organisms that took active part in water self-purication are subject to control of organisms of both lower and higher trophic levels in the food chain. The regulating role of organisms can be effectively studied with the use of the authors method of inhibitor analysis of regulatory interactions in trophic chains [11, 31]. Various forms of signaling, including informationbearing chemicals (environmental chemoregulators and chemomediators [9, 19, 20]) play an important role in the regulation of ecosystems. Water purication and permanent reproduction of its quality is an important component for ecosystem self-stabilization. This enables the regeneration of the
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normal state of habitats for the species represented in the ecosystem. Such regeneration of water quality is necessary for ecosystem stability, because autochthonous and allochthonous organic matter and nutrients are permanently delivered into water from land by water of tributaries, atmospheric precipitation, and solid particles carried by air [35]. Therefore, water selfpurication is as important for an aquatic ecosystem as DNA repair is for the heredity system. This allows us to regard water self-purication as an environmental repair in aquatic ecosystems. The wide range of variations in the ltration activity rates suggests the need to regulate this activity. The volume of water ltered within an hour and measured in the body volumes of the lterer amounts to 5 106 for nanoagellates and 5 105 for ciliates [35]. Cladocerans lter up to 414 ml per one organism per day [35] (according to [25], 20130 ml). Copepods and rotifers lter 227 [35] and 0.070.3 ml/day per animal, respectively [8]. All these aquatic animals and other lterers remove suspension from water. Thus, all forms of regulation and communication of organisms within community are of importance for maintaining the reliability of ecosystem functioning. An important role in the regulation and communication in aquatic communities belongs to dissolved substances, ecological chemoregulators and chemomediators [9, 19, 20]. ON THE RELATIONSHIP BETWEEN THE RELIABILITY OF WATER SELF-PURIFICATION SYSTEM AND AQUATIC ECOSYSTEM STABILITY Filtration activity is not only a part of water selfpurication process and water quality repair, but also a part of processes that maintain the stability of the aquatic ecosystem. The latter is performed through the conditioning of water, which serves as a habitat for many other aquatic species, and the environmental tax for the environmental stability that lterers pay in the form of pellets of organic material. These pellets form in the organisms of lterers (mollusks) from particulate organic matter they lter from water and release into the environment in the form of lumps. Pellets precipitate onto the bed of water bodies or streams. They are

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Table 5. The effect of surfactant Triton X-100 (TX-100), 5 mg/l, to the withdrawal of suspension of unicellular organisms from water because of inhibition of filtration activity of mollusks Unio tumidus. Here and in Table 6, EEW is the effect on the efficiency of suspension withdrawal [13, 15]. Vessel A contains 10 mollusks with a mean weight (wet weight with a shell) of 19.6 g, the total biomass of 196.25 g. Vessel B contains 10 mollusks with a mean weight (wet weight with a shell) of 19.4 g, the total biomass of 194.2 g. The mollusks were conducted in the Moskva R. (Odintsovo district). Vessels C and D contained no mollusks. Water volume in each vessel was 500 ml. T = 20C. Suspension of Scenedesmus quadricauda was added into each vessel (the age of the culture was 5 day; on KratsMayers medium, growth at T = 21C) Optical density at 500 nm (optical path of 10 mm) Time from the begin- solution solution without without ning of the of TX-100 of TX-100 TX-100 TX-100 EEF, % experi- was added was added ment, min A B C D 10 15 0.099 0.104 0.068 0.054 0.106 0.106 0.101 0.102 145.59 192.59

Comment (visual observations)

30

0.066

0.023

0.107

0.113

286.96

50

0.052

0.012

0.092

0.103

433.33

No difference was observed between the vessels; the vessels contain no pellets First lumps of microppellets appear in vessels A and B, water in vessel B becomes more transparent than in all other vessels Pellets, which are clearly seen in vessel B, are larger than the micropellets in vessel A; water in vessel B is more transparent than in vessel A and other vessels Pellets are clearly seen in vessels A and B; water in vessel B is more transparent than in vessel A and other vessels

used as food by many other aquatic animals, including zoobenthos and bacteria. The environmental tax is surprisingly high as compared with the share of C of the organic matter included in production. In some cases, it can be >100%, when calculated as the ratio of the amount of C not assimilated from the food (that is, C from fecal and pseudofecal pellets) to the amount of C consumed for production. The formation of pseudofeces by molluskslterers (that is, the process in which part of the ltered seston does not pass through the digestive tract of the mollusk but is prepared to the release into the environment in the form of pellets) begins at low seston concentration. Thus, at the concentration of seston as low as 2.6 mg/l (the concentration of seston is commonly much greater), mollusks Mytilus edulis (shell size of 1.7 cm) started releasing pseudofecal pellets [22]. Therefore, the formation of pseudofeces is not the result of excessive concentration of organic matter in the aquatic environment. The high environmental tax is justied because the lterers will eventually benet from the high level of stability of water quality characteristics. The entire system of water self-purication also benets from this, because it requires the wide diversity of aquatic species to maintain its stability. Aquatic ecosystems serve as one of the most important regulators of global geochemical cycles (e.g., of water and C), the stability of which withstands the hazard of global disturbances. Therefore, the reliability of water self-purication system is of importance for the global stability in the biosphere [19].

RESPONSE OF THE ENTIRE SYSTEM OF WATER SELF-PURIFICATION TO EXTERNAL (ANTHROPOGENIC) IMPACTS ON THE WATER BODY The author has found an essential element of lability in one of the processes involved in water self-purication, i.e., water ltration by aquatic animals (mollusks and rotifers) [1023, 3033]. Water ltration was inhibited in experiments by sublethal concentrations of anthropogenic pollutants, such as surfactants, surfactant-containing mixed preparations, and heavy metals (Tables 57). Other pollutants were found to have similar effect on mollusks and zooplanktonic lterers [22, 28]. The population biomass of lterers in polluted aquatic ecosystems decreases, the result of which is an additional drop in the total ltration activity in such ecosystems [22]. Therefore, the system of water self-purication processes and its quality formation is labile [12, 21, 22] and quickly rearranges to adjust to changes in the environment. The obtained data demonstrate the hazard of a decrease in the efciency of water self-purication system in aquatic ecosystems subject to anthropogenic impacts (chemical pollution of water bodies and streams) [1014, 1723, 3033]. RELATIONSHIP BETWEEN THIS THEORY AND FUNDAMENTAL ECOLOGICAL CONCEPTS An important principle in the organization of ecosystems is the interdependence and mutual usefulness of the organisms involved. This principle is conrmed
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Table 6. Effect of surfactant-containing mixture (OMO, 50 mg/l) on the withdrawal of suspension of unicellular organisms from water through inhibition of the filtration activity of Unio tumidus. Vessel A contains 10 mollusks with a mean weight (wet weight with the shell) of 19.16 g and a total biomass of 192 g. Vessel B contains 10 mollusks with a mean weight (wet weight with the shell) of 19.0 g and a total biomass of 190 g. The mollusks were collected in the Moskva R. (Odintsovo district). Vessels C and D contain no mollusks. Water volume in each vessel is 500 ml. T = 19.5C. Suspension of S. cerevisiae was added to both vessels Optical density at 500 nm (optical path of 10 mm) Time from the begin- solution solution without without ning of the of OMO of OMO OMO OMO EEF, % experi- was added was added ment, min A B C D 5 20 0.711 0.644 0.794 0.556 0.693 0.702 0.671 0.671 89.55 115.83

Comment (visual observations)

35 60 165

0.614 0.557

0.448 0.373

0.648 0.655

0.660 0.629

137.05 149.33

No visual distinctions between vessels Pellet lumps appear in vessel B; water in this vessel becomes more transparent than in all other vessels; vessel A does not contain pellets Pellets are clearly seen in vessel B, water is more transparent than in any other vessel The same Pellet lumps are also seen in vessel A, though not for all mollusks and with a lesser extent than in vessel B

so often that it has almost become an axiom and does not attract particular attention. However, its signicance manifests itself in a new way in the analysis of water self-purication processes in aquatic ecosystems. The cooperative functioning of procaryiote communities is one example. Another example is the high activity of lterers for removing suspension from water, during which the amount of suspended organic matter extracted from water is much greater than it is required for the lterers organism [1, 21, 22]. The environmental signicance of suspension removal from water and pellet formation is analyzed in detail in [22]. The assimilability of food by lterers in laboratory experiments was ~5060% [8], however it can be much lower in nature. Thus, mollusks Mytilus galloprovincialis (with a biomass of 2 g) featured the assimilability that varied within the year from 4.8 to 51% [22], that is, in some cases >95% of ltered out material was released by the mussels in the form of pellets. Thus, the synecological cooperation is one of the functional principles of the biotic system of water selfpurication. Purication of aquatic ecosystems is accompanied by transfer of chemical substances and their constituents from one point of the aquatic ecosystem into another. The results of data analysis support the earlier formulated proposition that a competitive unity of vector and stochastic motion of chemical elements and the regulation of these processes based on biological matter exists in aquatic ecosystems [18]. Conrmations were also obtained for the assumption that competitive unity and biological-matter-controlled regulation of cyclic and noncyclic paths of chemical elements, representing chains of successive transitions of
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chemical elements from one phase into another (interphase transfer) and from one organism into another (organism-to-organism transfers) take place in aquatic ecosystems [18]. The author emphasized that the regulation of many processes of transfer of chemical elements in aquatic ecosystems is biologically controlled, and the roles of both componentsbiotic and abiotic are equally important.
Table 7. Effect of various pollutants on suspension withdrawal from water by filterers [21, 22]. (The maximum value of EEF for the entire period of experiment is given. LD is liquid detergent, SDS is sodium dodecyl sulfate, TDTMA is tetradecyl trimethyl ammonium bromide, TMOC trimethyltinchloride) Substance TX-100 TDTMA SDS The same Copper sulfate Lead nitrate LD E LD Fairy TMOC TDTMA The same The same Organism Unio tumidus Crassostrea gigas M. edulis M. galloprovincialis C. gigas M. galloprovincialis M. galloprovincialis C. gigas C. gigas Dreissena polymorpha Brachionus angularis B. plicatilis B. calyciflorus Concentration, mg/l 5 0.5 >1 0.5 2 20 2 2 0.0110 0.5 0.5 0.5

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CONCLUSIONS When improving the methods and processes of wastewater treatment and the methods and processes of water treatment, one should take into account and use the natural principles of water purication under natural conditions. The preservation of water-purication potential of water bodies and streams is an important prerequisite of stable use of waterbiological and water resources (an essential condition for sustainable economic development and growth in RF). Practically all the biodiversity participates in the processes that lead to water quality formation, selfpurication of aquatic ecosystems or regulation of these processes. This demonstrates the need to preserve the biodiversity in aquatic ecosystems [16]. Onshore ecosystems and habitats actively participate in water purication processes. Therefore, it is necessary to preserve the biodiversity of these onshore ecosystems as well. The concept of preservation of biodiversity now differs from the previous approach based on the preservation of the gene pool of species. The nature protection objectives and the regime in the land and water area of the territory to be preserved should include the preservation of not only gene pool and populations of species, but also the level of functional activity of these populations (we mean the functional activity that contributes to the maintaining of water quality and, thus, the stability of the entire aquatic ecosystem) [16, 17]. The consistent application of the proposed theory allowed us to establish new types of hazards of chemical pollution in aquatic ecosystems [21, 22]. The theory of the polyfunctional role of aquatic biota suggests new proposals regarding the synecological approach to solving the problem of eutrophication of aquatic ecosystems [14]. We can expect the appearance of new pollutants, which may cause a hazardous drop in the self-purication capacity of aquatic ecosystems (water bodies and streams). ACKNOWLEDGMENTS The author is grateful to A.F. Alimov and M.E. Vinogradov (Institute of Oceanology), V.L. Kasyanov and T.I. Moiseenko (Water Problems Institute, RAS), V.V. Malakhov, E.A. Kriksunov, and V.D. Fedorov (Moscow State University), G.E. Shulman and G.A. Finenko (Institute of Biology of Southern Seas), C. Steinberg and N. Walz (Institute of Freshwater Ecology, Germany), J. Widdows (Plimooth Marine Laboratory) for discussion; N.E. Zurabova, M.P. Kolesnikov, N.N. Kolotilova, E.A. Kuznetsov, V.A. Kuznetsov, E.A. Matveeva, T.A. Ostroumova, L.V. Podshchekoldina, and L.I. Shpitonova (Moscow State University) for their help.

This study was partially supported by the Macarthur Foundation, and Open Society Institute. REFERENCES
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