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The reference for the publication: Ostroumov S.A.

Inhibitory analysis of top-down control: new keys to studying
eutrophication, algal blooms, and water self-purification. Hydrobiologia. 2002. vol. 469, pages 117-129

Inhibitory analysis of top-down control: new keys to studying eutrophication,
algal blooms, and water self-purification

S.A. Ostroumov

Lab of Physico-Chemistry of Biomembranes, Faculty of Biology, Moscow State University,
Moscow 119991, Russia;
e-mail: ar55[at]yandex.ru

Keywords: self-purification, filter-feeders, surfactants, detergents, benthic bivalves, aquatic
ecosystems, eutrophication, algal blooms, hazards of chemical pollution, water quality,
phytoplankton, marine and freshwater invertebrates, clearance rate, biological effects of
xenobiotics, pollutants

Abstract
Top-down control is an important type of interspecies interactions in food webs. It is
especially important for aquatic ecosystems. Phytoplankton grazers contribute to the
top-down control of phytoplankton populations. The paper is focused on the role of
benthic filter feeders in the control of plankton populations as a result of water
filtering and the removal of cells of plankton from the water column. New data on the
inhibitory effects of surfactants and detergents on benthic filter-feeders (Unio tumidus,
U. pictorum, Mytilus galloprovincialis, M. edulis, and Crassostrea gigas) are
presented and discussed. Importance and efficiency of that approach to the problems
of eutrophication and water self-purification is pointed out. Chemical pollution may
pose a threat to the natural top-down control of phytoplankton and water self-
purification process. The latter is considered an important prerequisite for sustainable
use of aquatic resources.
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The reference for the publication: Ostroumov S.A. Inhibitory analysis of top-down control: new keys to studying
eutrophication, algal blooms, and water self-purification. Hydrobiologia. 2002. vol. 469, pages 117-129

Abbreviations: CR - clearance rate; LAS - linear alkylbenzene sulphonate; NOEC - No
observable effect consentration; QSAR – quantitative structure - activity relationship; SFG -
Scope for Growth; SDS – sodium dodecyl sulphate; TDTMA -
tetradecyltrimethylammonium bromide; TX100 - Triton X-100

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The reference for the publication: Ostroumov S.A. Inhibitory analysis of top-down control: new keys to studying
eutrophication, algal blooms, and water self-purification. Hydrobiologia. 2002. vol. 469, pages 117-129

1. Introduction
By definition, the organisms of the two adjacent trophic levels interact with each other so
that the organisms of the higher trophic level may produce some effect on the organisms of
the lower trophic level. If the latter are not too abundant, the effects of the organisms of the
higher level lead to limiting, decreasing or stabilizing the populations of the organisms of
the lower trophic level. These effects might be considered a control or a partial control of
the organisms of the lower trophic level. Many examples of interactions of that type were
studied in various natural and experimental systems (Table 1). The significance of top-down
control attributes additional importance to studies of the grazing activity of crustaceans
(e.g., Sushchenya, 1975; Gutelmaher, 1986), rotifers (e.g., Monakov, 1998; Bul’on et al.,
1999), protozoan plankton (e.g., Bul’on et al., 1999), and benthic invertebrates (e.g.,
Alimov, 1981; Donkin et al., 1989, 1991; Zaika, 1992; Ogilvie & Mitchell, 1995; Widdows
et al., 1995a, Widdows et al., 1995b; Newell, 1999), and other invertebrates (Monakov,
1998).
In aquatic ecosystems, the problem of the control of the organisms of the lower
trophic level (algae) is of outstanding importance because it is relevant to the problem of
eutrophication. Also, control mechanisms are important in better understanding the problem
of algal blooms, including the toxic algae blooms. To avoid over-simplification, we should
realize that there are many factors that regulate the abundance of algal populations; top-
down control is only one of them.
Many species of invertebrates of both plankton and benthos belong to the higher
trophic level as compared with algae and cyanobacteria of phytoplankton. As for
zooplankton species and their filter-feeding activity, an important body of information was
presented and analyzed in (Sushchenya, 1975; Gutelmaher, 1986). Filtering activity of
benthic species has also been studied (e.g., Alimov, 1981; Ostroumov et al., 1997, 1998).
In this paper we focus on some species of invertebrates of benthos, which are filter-
feeders and in this capacity contribute to the top-down control of phytoplankton.

2. Role of benthic invertebrates in filtering water and resulting phytoplankton grazing:
filter-feeders
The diversity of benthic organisms that filter water and remove algal cell and other
particulate matter is broad. Filter-feeders inhabit the bottom of both freshwater and marine
ecosystems. To facilitate broader general conclusions, in this paper we will consider both
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The reference for the publication: Ostroumov S.A. Inhibitory analysis of top-down control: new keys to studying
eutrophication, algal blooms, and water self-purification. Hydrobiologia. 2002. vol. 469, pages 117-129

freshwater and marine organisms. The range of filter-feeders includes sponges, polychaetes,
molluscs, echinoderms, larvae of many insects, ascidians, and some other invertebrates.
There are many examples of massive scale water filtering by benthos (e.g., Table 2;
see also: Alimov, 1981; Ostroumov & Fedorov, 1999). It was shown that in some man-
made reservoirs the total volume of water is filtered by benthic bivalves 2-24 times annually
(e.g., Konstantinov, 1979). In a shallow lake in New Zealand the total volume is filtered
during a time period of less than 2 days (Ogilvie & Mitchell, 1995). Equally massive
filtering activity was discovered for the benthic sponges in the coastal waters of Lake Baikal
which stores 22, 995 km3 of superb clean water (for comparison, the amount of the annual
world consumption of freshwater was 3, 240 km3, and the annual freshwater withdrawal in
Europe was 359 km3, in North and Central America 697 km3; the data for year 1987)
(World Resources 1995-1995).
As a result of water filtering, algal cells are removed from the water column. It is
important that some filter-feeders (e.g., bivalves) remove more algae than they need for
feeding purposes. Excessive amounts of algae biomass and other particulate matter are
excreted in the form of pellets (to distinguish them from regular faeces they are called
pseudofaeces) which are larger in size than the algal cells and therefore they settle to the
bottom rapidly. The amount of pseudofaeces may exceed the amount of the assimilated food
manyfold. As a result, the total activity of bivalve molluscs in removing algal biomass from
the water column and in making water clearer is far beyond just the trophic needs of
bivalves.
The total weight of organic matter that is removed from a water column and
deposited as bottom sediments is measured as high as kilograms per m2 per year. E.g., in the
ecosystem of the man-made reservoir Volgogradskoe, the amount of the formerly suspended
matter that was removed by molluscs from the water column and finally sedimented was 8.3
kg m-2 annually (Kondratiev, 1976; cited in Konstantinov, 1979). For the entire reservoir
that is located in the center of the largest European river, the amount of sedimented matter
was as high as 29 million tons.

3. Inhibitory effects of xenobiotics and pollutants: a decrease in water filtration and
associated phytoplankton grazing
Man-made chemicals can produce strong inhibition of water filtering by benthic
molluscs or impair the normal pattern of opening bivalves which is needed to maintain the
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The reference for the publication: Ostroumov S.A. Inhibitory analysis of top-down control: new keys to studying
eutrophication, algal blooms, and water self-purification. Hydrobiologia. 2002. vol. 469, pages 117-129

efficient filtration of water. Some examples of those effects are given in Table 3. More
examples could be found in literature (e.g., Stuijfzand, 1995; Ostroumov, 1998). The
experiments were usually conducted using some phytoplankton species as the organism that
is being removed from the water column during the filtration experiment. Thus, in
experiments with bivalve Mytilus edulis, the algae Isochrysis galbana are often used
(Donkin et al., 1997; Ostroumov et al., 1997; 1998). In our experiments with M.
galloprovincialis (see below) we have observed a xenobiotic-induced decrease in grazing
phytoplankton cells of Monochrysis lutheri and Dunaliella viridis. In our experiments with
freshwater bivalves Unio tumidus and U. pictorum, we described some pollutant-induced
inhibition of the removal of green algae Scenedesmus quadricauda and cyanobacteria
Synechocystis.
The major part of our experiments were done in the laboratory. Under field
conditions, it was described that in polluted habitats the biomass and vitality of bivalves
declined (Zaika, 1992), which means that their contribution to water filtering is negatively
affected. It was possible to develop an integrative parameter, Scope for Growth (SFG)
which enables the scientist to estimate the total amount of energy available for the
population of mussels for its growth and reproduction (after deduction of the amount of
energy is lost during respiration etc.) (Widdows et al., 1995a, 1995b). It was shown that in
terms of the entire populations, in polluted habitats the reduced filtration and reduced intake
of energy from digested plankton (and seston as a whole) led to the fact that SFG was
reduced.

4. More specific examples and new data: inhibitory effects of surfactants
We have initiated a systematic study of the effects of another class of aquatic
pollutants, namely surfactants, on the water-filtering activity of bivalves and on the
resulting removal of algal cells from the water column.
Among the various organic chemicals that are entering the natural environment in
large amounts (Yablokov & Ostroumov, 1983, 1985, 1991), surfactants play a significant
role (Ostroumov, 1986; 1990; 1991; 1994 a; 1994b; Marcomini et al., 1988; Quiroga et al.,
1989; Granmo et al, 1991; Fernandez et al., 199; Lewis, 1991; Takada & Ishiwatari, 1991;
Chalaux et al., 1992; Terzic & Ahel, 1993). It was shown that surfactants produce negative
and sometimes also stimulatory effects on cyanobacteria (Waterbury & Ostroumov, 1994),
green algae (e.g., Goryunova & Ostroumov, 1986), diatoms (Ostroumov & Maertz-Wente,
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The reference for the publication: Ostroumov S.A. Inhibitory analysis of top-down control: new keys to studying
eutrophication, algal blooms, and water self-purification. Hydrobiologia. 2002. vol. 469, pages 117-129

1991), plant seedlings (Ostroumov, 1986; 1990; 1991; Nagel et al., 1987; Maximov et al.,
1988; Telitchenko & Ostroumov, 1990), shrimp (Drewa et al., 1988), Daphnia magna and
D. pulex (e.g., Maki & Bishop, 1979; Martinez et al., 1989), freshwater amphipods
(Pantani et al., 1995), rotifers (Kartasheva & Ostroumov, 1998), fish (e.g., Versteeg &
Shorter, 1992; Malcolm et al., 1995). Some data on the effects of linear alkylbenzene
sulphonate (LAS) on Mytilus galloprovincialis Lmk (Bressan et al., 1989; Marin et al.,
1993), Mytilus edulis (Granmo, 1972) and some other marine benthic species (Marin et al.,
1991) are available. However, almost nothing was known about the effects of
alkylsulfates, nonionic surfactants (derivatives of nonylphenols), and some other surfactants
as well as detergents on the filtering activity of Mytilus edulis, M. galloprovincialis,
Crassostrea gigas, Unio tumidus, and U. pictorum.
The purpose of the experimental part of this work was to obtain data on the effects of
some surfactants and surfactant-containing products including detergents, on the ability of
bivalves (M. edulis, M. galloprovincialis, Unio tumidus, and U. pictorum) to filter water
and remove algal or other cells from it.
Freshwater mussels Unio sp. were collected in the Moscow River. Mytilus
galloprovincialis were collected at the Black Sea. Crassostrea gigas were grown at a
mariculture farm (the Black Sea, Institute of Biology of Southern Seas NANU). M. edulis
were collected at the Exmouth estuary and kept in tanks with aeration , water flow and
periodic automatic imitation of low tide (water was removed out of tanks for 3 h every day)
(Dr. Donkin’s participation and help in the work with M. edulis is acknowledged).
The temperature in experiments with M. galloprovincialis and C. gigas was mostly
22-27 °C, in the experiments with Unio sp. 18-20 °C. The cell removal and the cell density
during the filtration by molluscs was measured using Hitachi 200-20 spectrophotometer
(experiments with Unio sp.) and SF-26 (LOMO) spectrophotometer (experiments with M.
galloprovincialis and C. gigas). In experiments with M. edulis (temperature 16 °C), the
number of cells per unit of volume was measured using the Coulter counter ( Coulter
Electronics, model Industrial D). When a sample of filtered water without adding algae was
used, the Coulter count was usually below 200.
The clearance rate (CR) was calculated according to Widdows & Salkeld (1993)
using the following equation:

CR (l h-1) = (Volume of water e.g. 2 l) x (loge C1 -- logeC2)/time interval in h
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The reference for the publication: Ostroumov S.A. Inhibitory analysis of top-down control: new keys to studying
eutrophication, algal blooms, and water self-purification. Hydrobiologia. 2002. vol. 469, pages 117-129

where C1 and C2 are cell concentrations at the beginning and end of each time increment
(e.g. 0.5 h).
Statistical analysis was performed using EXCEL software. For linear regression
analysis, an option was used which gives an opportunity to fix the intercept at a
predetermined value.
Several chemicals were used. Sodium dodecyl sulfate (SDS) (molecular mass
288.38) was purchased from Fluka. The purity was > 99% (assayed by GC, analysis
number 332533/1 395). Triton X-100 (TX100) (x = 9-10 ethoxy units, H2O < 1 %, residue
on ignition, 0.2%, analysis number 43306/1 795) was purchased also from Fluka.
Tetradecyltrimethylammonium bromide (TDTMA, molecular mass 336.4) was purchased
from Sigma (St.Louis, Missouri, 63178 USA; lot 55H1322). Detergents used were available
commercially.
Results of the experimentation were as following.
Freshwater bivalves, Unio tumidus and U. pictorum removed planktonic cells from
water. The ability to do so was inhibited by surfactants of several types (Table 4),
including TDTMA, and TX100 .
A marine species, M. galloprovincialis, was also efficient in removing from water
cells of phytoplankton and unicellular organisms in general. Several surfactants as well as
detergents which contain surfactants inhibited this ability of M. galloprovincialis (Table 4).
The chemicals tested included surfactants TDTMA, SDS, and several detergents, such as
Tide-Lemon, Lotos-Extra, Losk-Universal.
In experiments with M. edulis, after one hour of filtering, in the control set (clean
water) the number of algal cells per unit of volume decreased to almost 5.6% of the initial
level, which is a good example of how efficiently bivalves can control planktonic
populations (Table 5). This is in accord with the large amount of data on the significant
filtration rates of bivalves (Alimov, 1981; Monakov, 1998) and their impact on ecosystems
(Zaika, 1992). In the important series of measurements, in the control beakers (filtration of
unpolluted water) the number of algal cells decreased by a factor of 15.98, while in the
beakers with SDS (1 mg l–1) the number of cells decreased by a factor of 7.93. Thus, the
algal cell density in control was half that in the system at the initial concentration of 1 mg l–
1
. The difference increased by the end of the experiment.

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The reference for the publication: Ostroumov S.A. Inhibitory analysis of top-down control: new keys to studying
eutrophication, algal blooms, and water self-purification. Hydrobiologia. 2002. vol. 469, pages 117-129

When the initial concentration of SDS was 2 mg l–1, a substantial difference from the
control set was observed after the first half-hour period (Table 6). After 65 min of filtering,
the algal cell density in the control set was almost 1/3 that contained in the system with
SDS.
Further increase of the initial concentration of SDS up to 4 mg l–1 caused a dramatic
3-times increase of the cell density over that in the control set after only 35 min of filtering.
In 65-min of filtering, the difference was 6-fold, and following 95-min filtering - over 14-
fold.
At the initial concentration of SDS 5 mg l–1, the difference between systems with and
without SDS was over 16-fold after 125 min of filtering.
It was possible to calculate the clearance rate (CR), using a standard formula widely
accepted in the literature ( Donkin et al. 1989; 1991; Widdows & Salkeld, 1993).
The summary of the inhibitory effects shows, with a few exceptions, two general
trends:
1) an increase in the initial concentration of SDS in the range of 0.5 to 5 mg l–1 gave rise to
an the increase in the inhibitory effect on CR (Table 7);
2) at any given concentration of SDS, the highest effect took place during the first 30-min
period, with some decrease in the inhibitory effect by the end of the experiment.
The latter trend, however, was not paralleled by a mitigation of the effect on the
residual algal cell density in the water. When the cell density was considered, the difference
from the control was maximal by the end of the experiment.
Using another chemical, a non-ionic surfactant Triton X-100, we obtained similar
data with EC50 close to that of SDS (Table 8). At a concentration of 4 mg l–1, the inhibition
of the clearance rate during the time period of 30 min after the beginning of the experiment
was almost 10-fold, and during the later period of time, the inhibition was about 5-fold.
The data obtained in our study showed that the filtering activity of mussels
demonstrated a more sensitive response than some other biotests we had used in our
experiments in bioassaying SDS, including green algae (Goryunova & Ostroumov, 1986)
and plant seedlings (Nagel et al., 1987). The filtering activity of mussels was also more
sensitive to SDS than some of the traditional lethal biotests with aquatic invertebrates and
fish which had been applied for studies of LAS and alkyl sulfates (Sivak et al., 1982;
Ostroumov, 1991).

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The reference for the publication: Ostroumov S.A. Inhibitory analysis of top-down control: new keys to studying
eutrophication, algal blooms, and water self-purification. Hydrobiologia. 2002. vol. 469, pages 117-129

It is noteworthy that the inhibitory effect of SDS on CR was developed within a
rather narrow range of SDS concentrations (1 to 5 mg l–1). That could be in accord with a
hypothesis that the decrease in CR is at least in part the result of a behavioural response of
mussels.
Our data on effects of SDS are in good agreement with the results obtained by other
authors who studied effects of another anionic surfactant, linear alkylbenzene sulphonate
(LAS) on filtering rate. It was shown that in experiments with exposure for 48 h and 96 h
the filtration rate of mussels Mytilus galloprovincialis was reduced when concentration of
dissolved LAS was higher than 1.5 mg l–1 (Bressan et al., 1989). In our experiments the
biotest was slightly more sensitive as we exposed the animals to the surfactant for 1.5 h
prior to beginning measurements and observed some inhibition at the initial concentration of
1 mg l–1.
Bressan et al. (1989) studied also effects of LAS on the growth of mussels and on
mortality and spermatozoids of freshwater bivalve molluscs. They observed some decrease
in the increment of length of the major axis of the shell of mussels at concentrations of LAS
as low as 0.25 and 0.5 mg l–1, but the effect required up to 70 days to be observed. No
significant effects were found within 30 days of their experiments. The length of time that
was necessary to reveal the effect was a limitation of the technique, however it was
impressive to observe almost a 2-fold decrease in growth when the chronic experiment with
a relatively low level of LAS (0.25 mg l–1 ) lasted for 160 days and more.
In a parallel experiment the same authors observed a 30% increase in the respiration
of LAS-treated (220 days, 0.25 mg l–1 ) young mussels (Bressan et al., 1989). Unfortunately,
they did not specify what they called young mussels.
Some decrease in filtering rate was observed in another set of experiments when the
concentration of LAS was 0.25 mg l–1 , but the duration of the surfactant treatment was
much longer (220 days) than in our experiment, and the size of mussels was again not
specified (Bressan et al., 1989). Also, they have shown that, at a concentration of 1 mg l–1,
LAS inhibited the filtration rate after 7 days of exposure. It seems important that in our
experiments we observed effects after only 1.5 hours of exposure to the anionic surfactant.
The LC50 (48 h) was about 40 mg l–1 and LC50 (96 h) was about 1.7 mg l–1 (Bressan
et al., 1989), which was much lower than in the case of freshwater bivalves Anodonta
cygnea and Unio elongatulus. For the latter two species, LC50 (96 h) was about 200 mg l–1

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The reference for the publication: Ostroumov S.A. Inhibitory analysis of top-down control: new keys to studying
eutrophication, algal blooms, and water self-purification. Hydrobiologia. 2002. vol. 469, pages 117-129

(Bressan et al, 1989). The mobility of spermatozoa of A. cygnea was almost completely
inhibited at a concentration of LAS equal to 20 mg l–1.
Measurements of CR were used to quantify the toxic effects of chemicals and to study
QSAR (Donkin & Widdows, 1990) for various chemicals, including alkanes and phenyl
alkanes (Donkin et al., 1991) as well as such aromatics as toluene, naphthalene, n-
propylbenzene, 1-chloronaphthalene, biphenyl etc. (Donkin et al., 1989). Two
xenobiotics, including an organotin compound, inhibited the fitration rate by Dressena
polymorpha and Crenomytilus grayanus (Mitin, 1984).
The filtering activity of not only bivalves, but also of other filter-feeders is
vulnerable to the inhibition by surfactants. In experiments with rotifers Brachionus
angularis Gosse, we have shown that TDTMA inhibited their filtration rate and the removal
of cells of Chlorella sp. from the water (Kartasheva & Ostroumov, 1998). At a TDTMA
concentration of 0.5 mg l–1, the average efficiency of filtration was 58.5% of that in control.
However important these kinds of studies of CR are, it is also important to consider
the general consequences of a decrease in the CR for the ecosystem.
The role of the filtering activity of mussels is connected with their high population
densities. It was estimated that at Narragansett Bay, Rhode Island, mussels represented
about 77% (11 kg m -2) of the total community dry weight (Nixon et al., 1971), and
numbers of the same order of magnitude were reported for other locations (Seed &
Suchanek, 1992). Taking into account that, in our experiments, one mussel with a total wet
weight about 8.5 g filtered over 1 L of water per hour, it is easy to estimate that, at high
abundancy, a mussel community may filter over 100 L water per hour per 1 m2 of the sea
bottom.
A comparison of the tables for residual cell densities and CR for specific
concentrations of surfactants shows that even a small decrease in CR produces a large
difference in the residual cell density. The latter parameter may be considered as a model
for any kind of particles which are being removed from the seawater by mussel filtering. In
this way we may predict a huge decline in the natural ability of benthic communities to
purify natural water when the water is polluted by surfactants as well as by other chemicals
reducing the CR.
Changes (inhibition) of the filtering activity of bivalves might have many
consequences in changing many parameters and processes in ecosystems, which were
considered in more detail in (Ostroumov et al., 1997; 1998; Ostroumov, 1998).
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The reference for the publication: Ostroumov S.A. Inhibitory analysis of top-down control: new keys to studying
eutrophication, algal blooms, and water self-purification. Hydrobiologia. 2002. vol. 469, pages 117-129

Those considerations show that the inhibition of CR has consequences not limited by
the prosperity of the mussel population, but that it is important for the state of the marine
and estuarine ecosystems in much broader terms. Prospects of chemical-induced inhibition
of water filtration by bivalves poses some ecological hazards in view of the role of bivalves
in eutrophication control. The latter was studied in the case of the ecosystem of Chesapeake
Bay (the Atlantic coast of the U.S.A.) (Newell et al., 1999).

5. Sensitivity of plankton grazers to xenobiotics - analogous evidence for zooplankton
Analysis of the specific LC50 for Cladocera and various species of algae shows that
in case of many pollutants Cladocera are more sensitive than algae. According to the data
disseminated at the recent workshop in Netherlands (9-12 December 1999, Den Helder,
TNO; participants of the research project: M. Scholten, R. Jak, B. Clement, E. Foekema,
P.Hernandez, K.Kaag, H. van Dokkum, M. Smit), in the case of the following pesticides,
species of Cladocera (mainly Daphnia magna, D. pulex, Ceriodaphnia dubia) are more
sensitive: anilazin, benomyl, bentazon, cyfluthrin, dimethoat, lindan, maneb, zineb, and
ziram. In case of several pesticides, it was directly shown that the inhibitory effects on
feeding were observed at lower concentrations, than the concentrations which induced
mortality. EC50 (effects on feeding within 4-24 h) were lower than LC50 (24-48 h) for
endosulfan, diazinon, methyl parathion, lindan, and dichlobenil (according to the data
distributed at the same workshop). In case of atrazine, a concentration of 1.6 mg l–1 within
10 min produced 50% reduction in feeding, which shows again that feeding activity is
inhibited at concentrations lower than those inducing mortality: LC50 (48 h) was 9.88 mg l–1.
Also, NOEC (No observable effect concentration) was the basis for comparing
sensitivities of Cladocera and various species of algae to pesticides. In case of the following
chemicals a higher sensitivity of Cladocera was found: azinfos-methyl, cyromazin,
diazinon, dimethoat, endosulfan, fenpropathrin, malathion, mecoprop, propoxur, trifluralin,
and some other pesticides.
All these data as well as the new evidence in the experiments conducted at TNO
during the project led by Dr. M. Scholten (see Table 1) are in accord with the concept that
pollutants may impair top-down control of algae. This conclusion is analogous to the
conclusion made by us on the basis of our data for benthic filter-feeders.

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The reference for the publication: Ostroumov S.A. Inhibitory analysis of top-down control: new keys to studying
eutrophication, algal blooms, and water self-purification. Hydrobiologia. 2002. vol. 469, pages 117-129

6. Synoptic Overview and General Conclusions
Some benthic organisms, including spongi, polychets, bivalves, echinoderms, larvae
of insects, ascidia and some others proved to be efficient organisms in filtering water and
thereby in reducing the amount of particulate matter suspended in the water. Benthic filter-
feeders remove from the surrounding water various suspended particles including algal
cells. By doing so, they contribute to natural mechanisms that keep algal populations under
some control. That type of top-down control under some circumstances might become
especially important. The problem of algal blooms in the context of eutrophication is
increasing attention to all mechanisms of control of algal populations including the control
by virtue of water filtering by benthic filter-feeders, including bivalves. Some pollutants
were shown to be efficient inhibitors that decrease water filtering and resulting grazing
phytoplankton. Those chemicals produced a decrease in removal of algae from water
column by bivalves.
The author initiated systematic studies of effects of surfactants and detergents on
filtering activity and removal of algae by freshwater and marine bivalves. Marine and
freeshwater bivalves Mytilus edulis, M. galloprovincialis, and Unio sp. are efficient in
removing unicellular organisms from water in result of their filtration activity. They are
capable of drastically reducing the amount of cells of phytoplankton in water. This is an
important mechanism contributing to natural control of algal populations in ecosystems.
This regulatory mechanism is vulnerable to aquatic pollutants as exemplified by surfactants
and detergents. New data are obtained and presented in this paper on how surfactants
(anionic, non-ionic, and cationic ones) and surfactant-containing detergents inhibit the
ability of marine and freshwater bivalves to remove cells of algae and cyanobacteria from
water. On the basis of our new data, the final conclusion is that the new evidence support
the views proposed in (Ostroumov, 1998; 1999; 2000c; 2000e) about the vulnerability of the
filtration activity of invertebrates (both planktonic and benthic animals) to some pollutants,
including surfactants. Our data and general conclusion are in accord with the idea that
pollutants can induce reduction in grazing efficiency of benthic and planktonic
invertebrates.
We consider the studies of inhibitory effects of chemicals on fiter-feeders as an
effective approach to elucidating the details of filter-feeding and associated removal of
phytoplankton from the water column. The mechanisms and rates of plankton removal are

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The reference for the publication: Ostroumov S.A. Inhibitory analysis of top-down control: new keys to studying
eutrophication, algal blooms, and water self-purification. Hydrobiologia. 2002. vol. 469, pages 117-129

of utmost importance for controlling levels of plankton which are the key parameters in
processes of eutrophication and algal blooms.
Water filtering activity of invertebrates is part of water self-purification in
ecosystems. The self-purification of water is one of preconditions for the sustainable use of
water resources. Therefore, the vulnerability of filter-feeders to aquatic pollutants (including
surfactants and detergents) leads to a potential threat to the sustainable use of aquatic
resources in situations when the ability of ecosystems to purify water is inhibited by
pollutants.
In sum, on the basis of the data presented here and in some of our publications
(Ostroumov, 1998; 1999; 2000a; Ostroumov et al., 1997, 1998; Ostroumov & Fedorov,
1999), the following inferences are to be made:
1. Surfactants inhibit the filtering ability of marine and freshwater bivalves with a drastic
effect on the amount of particulate material (modelled here by algal cells) left in the water.
2. When considering the environmental importance of surfactants and detergents (and of a
broader range of xenobiotics and pollutants as well), the ramifications relevant to
disturbance of the natural ability of the ecosystem to control phytoplankton populations
should be taken into account.
3. Our new data are in accordance with the opinion (Ostroumov, 1990; 1991; 2000b;
2000c; 2000d; Telitchenko & Ostroumov, 1990; Yablokov & Ostroumov, 1991) that
surfactants, if being discharged into the environment at substancial rates, might, under some
circumstances and in some ecosystems, become more significant as environmental
pollutants than it was thought before.
4. We make the prediction that many new examples are to be found of pollutants (both
organic and inorganic) which inhibit filtration rate of filter-feeders (not only bivalves, but
also other benthic and plankton organisms) and by doing so reduce the ability of
invertebrates to control unicellular plankton populations. We predict that new examples are
to be found of pollutants which inhibit the ability of invertebrates to control eutrophication.
5. Sustainable use of resources of aquatic ecosystems requires as an important pre-condition
the efficient functioning of the ecosystems towards self-regulating and water self-
purification. This pre-requisite includes normal functioning of top-down control excercised
by the organisms at the higher levels of the trophic chains of ecosystems.

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The reference for the publication: Ostroumov S.A. Inhibitory analysis of top-down control: new keys to studying
eutrophication, algal blooms, and water self-purification. Hydrobiologia. 2002. vol. 469, pages 117-129

6. Studies of inhibitory effects of chemicals on the top-level organisms (e.g., grazers of
plankton, including benthic filter-feeders) is a useful approach in obtaining information on
the top-down control in trophic chains.

Acknowledgements. The author is grateful to many colleagues at Moscow University, the
Institute of Biology of Southern Seas NANU (Sevastopol), and the Plymouth Marine
Laboratory for assistance, to Prof. V. D. Fedorov, Prof. M.E.Vinogradov, Prof. A.F.Alimov,
Prof. V.V.Malakhov, Prof. V.N.Maximov, Prof. V.E.Zaika, Prof. G. E. Shulman, Prof.
N. Walz, Prof. J. Widdows and Prof. P. Wangersky for stimulating discussions, to Prof.
H. Dumont for reading the manuscript, to Dr. N. N. Kolotilova and Ms. N. Zourabova for
help, to Ms. Saara Turkki for providing copies of useful publications. The grants from the
MacArthur Foundation (the Program on Global Security and Sustainability) and the Open
Society Foundation (RSS grant No. 1306/1999) supported the research with M.
galloprovincialis and Unio sp. The work with M. edulis was done together with Dr. P.
Donkin and funded by EERO and in part by the IBG. After writing the paper, the author got
further encouragement from Dr. M. Scholten, Prof. N. M. van Straalen and other
participants of the workshop on eutrophication and toxification (Netherlands, Den Helder,
TNO).

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The reference for the publication: Ostroumov S.A. Inhibitory analysis of top-down control: new keys to studying
eutrophication, algal blooms, and water self-purification. Hydrobiologia. 2002. vol. 469, pages 117-129

Table 1. Top-down control in various natural and experimental systems (examples)
Systems ‘Top’- ‘Down’- Consequences of References
(in brackets- organisms organisms the removal or
the number of inactivity of top-
trophic levels) organisms
Great Salt Predacious Algae Increase in the Wurtsbaugh,
Lake, Utah, insect, (chl-a was population of 1992
the USA (3) Trichocorixa measured) shrimp Artemia
verticalis franciscana and
(food: 20-fold decrease
Artemia in Chl-a
franciscana)
Intertidal Birds, Larus Algae, Increase in Wootton, 1992
community, glaucescens, Halosaccion populations of
the north-west and glandiforme, intertidal
coast of the Haematopus and Porphyra animals (limpet
USA bachmani spp. Lottia digitalis),
(3) (food: decrease in algae
intertidal
animals)
Eutrophic lake Coregonus Algae increase in Reinertsen et al.,
Haugatjern lavaretus populations of 1990
(Norway) (3) Daphnia
galeata, 4-fold
decrease in the
biomass of algae
Mesotrophic Daphnia Chlorella sp., Increase in Scholten et al.,
Amadorio longispina, Scenedesmus Chl-a 1999 1
reservoir Daphnia sp.,
(Spain), 3000- magna Pediastrum
L enclosures sp.,
(2) Mycrocystis
aeruginosa
Outdoor Ceriodaphnia Chlorella sp.
Increase in algal Scholten et al,
enclosure sp., Daphnia populations 1999 1
experiments magna,
with 80-L test Simocephalus
containers vetulus
(‘plankton
ecoassay’) (2)
Microcosm Bivalves Algae See in the text New data by the
experiments (Unio sp., (see in the author
(2) Mytilus sp.) text)
1
Scholten et al., 1999 - personal communication; the relevant research was done by an
international team (headed by M. Scholten) while working on a project developed by TNO
(Netherlands); the results were presented at the workshop (9-12 December 1999, Den
Helder, Netherlands); the book by M. Scholten et al. (sponsored by CEEP) is in
preparation.
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The reference for the publication: Ostroumov S.A. Inhibitory analysis of top-down control: new keys to studying
eutrophication, algal blooms, and water self-purification. Hydrobiologia. 2002. vol. 469, pages 117-129

Table 2. Water-filtering activity of benthic organisms in some ecosystems (examples)

Ecosystems and water Water-filtering activity of References
bodies benthic organisms
Man-made reservoir Bivalves filter over 840x109 Kondratiev, 1976; cit. in
Volgogradskoe (Russia) m3 water annually, i.e. 24 Konstantinov, 1979
times the volume of the
reservoir
Man-made reservoir Annually bivalves filter two Lvova-Kachanova, 1971;
Uchinskoe (Russia) volumes of the reservoir cit. in Konstantinov,
1979
Estuary of the River Dnepr Annually bivalves filter the Alexeenko, Alexandrova,
(Dneprovsko-Bugskii volume of the estuary over 1995; cit. in Ostroumov
Liman, Ukraine), area 800- 16 times & Fedorov, 1999
928 km2, volume 3-4.1 km3
Lake Tuakitoto (New Bivalves (Hyridella Ogilvie & Mitchell, 1995
Zealand) menziesi) filter the total
volume of the lake during
32 hours

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The reference for the publication: Ostroumov S.A. Inhibitory analysis of top-down control: new keys to studying
eutrophication, algal blooms, and water self-purification. Hydrobiologia. 2002. vol. 469, pages 117-129

Table 3. Xenobiotics and contaminants that were shown to inhibit water-filtering activity
of bivalves

No. Contaminants and xenobiotics References
1 Organochlorine pesticides, lindane Donkin et al., 1997
and endrine
2 Organophosphate pesticide, Varanka, 1987 1 ; McHenery et al.,
dichlorvos 1991 2 ; Donkin et al., 1997
3 Organophosphate pesticide, Varanka, 1987 1
malathione
4 Pyrethroids permethrin, deltamethrin Varanka, 1987 1
5 Organotin compound Mitin, 1984
6 Dibutyltin (DBT), tributyltin (TBT) Widdows & Page, 1993
7 Synthetic surfactants Ostroumov et al., 1997, 1998; this
study - see tables below
8 Detergents This study - see next tables
1
The chemical altered the periodicity of opening freshwater bivalves, which could diminish
the normal filtration rate and decrease grazing efficiency;
2
The chemical made bivalves to close, which led to a decrease in filtration rate.

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The reference for the publication: Ostroumov S.A. Inhibitory analysis of top-down control: new keys to studying
eutrophication, algal blooms, and water self-purification. Hydrobiologia. 2002. vol. 469, pages 117-129

Table 4. New data on the inhibitory effect of surfactants and products that contain surfactants on
the filtration efficiency of bivalve molluscs1

No Chemicals Concent- Species of bivalves Unicellular organisms that
which caused rations, were being removed from
inhibition mg l–1 the water

1 TX100 5 Unio tumidus Scenedesmus
quadricauda,
Synechocystis sp.6803;
2 TX100 1 U. tumidus Synechocystis sp.6803;
3 TDTMA 1-2 U. pictorum Synechocystis sp.6803
4 TDTMA 1-2 U. pictorum Saccharomyces cerevisiae
5 SDS 1.7 M. Dunaliella viridis
galloprovincialis
6 TDTMA 1 M. Monochrysis lutheri
galloprovincialis
7 AHC 5-60 M. Monochrysis lutheri
galloprovincialis
8 Detergents Tide- 6.7-50 Mytilus Monochrysis lutheri
Lemon, Losk- galloprovincialis
Universal,
Lotos-Extra
9 Detergent 10-20 Mytilus Saccharomyces cerevisiae
IXI galloprovincialis
10 Detergents 1-30 Crassostrea gigas Saccharomyces cerevisiae
Vesna, Lanza,
Deni
11 SDS 0.5 Crassostrea gigas Saccharomyces cerevisiae
12 SDS 1-5 M. edulis Isochrysis galbana
13 TX100 0.5-5 M. edulis Isochrysis galbana

1
AHC - Avon Herbal Care (Avon Cosmetics). Authors thanks Dr. P. Donkin, Mr. F. Staff,
Dr. N. N. Kolotilova, Ms. N. E. Zourabova for help in taking some measurements. Besides
phytoplankton, S. cerevisiae was used in some experiments as a model for suspended matter.

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The reference for the publication: Ostroumov S.A. Inhibitory analysis of top-down control: new keys to studying
eutrophication, algal blooms, and water self-purification. Hydrobiologia. 2002. vol. 469, pages 117-129

Table 5. Decrease in Isochrysis galbana cell density (per 0.5 ml) during filtering by
Mytilus edulis in clean water (control beakers, A) and at 1 mg l–1 SDS (experimental
beakers, B) (S.E., standard error) (after Ostroumov et al., 1997, with some changes)1

Experimental Control beakers
beakers with SDS (no SDS added)
Time Cells S.E. Cells S.E. Ratio
after ad- count, count, A/B
ding average average
algae, (expl) (con-
min A trol)
B
5 17440.8 363.27 17416 1373.3 1.001
35 6593.33 1072.22 4881.58 856.36 1.351
65 2199 789.1 1089.83 247.08 2.018
95 1003 448.33 370.25 66.89 2.709

1
Cell density at the beginning of the experiment: 19533 per 0.5 ml.
Average Coulter count in filtered sea water: 131.7.

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The reference for the publication: Ostroumov S.A. Inhibitory analysis of top-down control: new keys to studying
eutrophication, algal blooms, and water self-purification. Hydrobiologia. 2002. vol. 469, pages 117-129

Table 6. Effect of SDS (2 mg l–1) on the efficiency of water filtering measured as the the
number of cells of Isochrysis galbana (per 0.5 mL) in the water after the 30-min period of
filtering by Mytilus edulis 1

No. of Presence (+) Number of cells measured Average number of cells
beaker or absence (-) by the Coulter counter (S.E., standard error)
of SDS
1 + 9307; 9537; 9311 9385.0
3 + 7475; 7455; 7384 7438.0
5 + 5880; 5779; 5678 5779.0
7 + 10608; 10508; 10467 10527.7
1,3,5,7 + Average for the four beakers: 8282.4 (S.E. 549.0)
2 - 5019; 4993; 4843 4951.7
4 - 4007; 3938; 4007 3984.0
6 - 4806; 4739; 4668 4737.7
8 - 3126; 2931; 3055 3037.3
2,4,6,8 - Average for the four beakers: 4177.7 (S.E. 227.0)
9 - 17875; 17881; 17697 17817.7
(without
mussels
1
The difference of the averages for the control and experimental sets was statistically
significant (the significance level >99.9%); p=2 x 10 -6; the concentration of the cells in the
beginning of the experiment: 17809.7; temperature 16° C; 16 animals were used in the
experiment, the average wet weight of one animal was 8.8 g (with the shell). Dr. P. Donkin
participated in the experiment.

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The reference for the publication: Ostroumov S.A. Inhibitory analysis of top-down control: new keys to studying
eutrophication, algal blooms, and water self-purification. Hydrobiologia. 2002. vol. 469, pages 117-129

Table 7.
Inhibition (%) of the clearance rate (CR) of Isochrysis galbana during filtering by
Mytilus edulis at various concentrations of SDS (after Ostroumov et al., 1998, with
some changes)

Concentrations of SDS, mg l–1

Time 0 0.5 1 2 4 5

period,

min

5 - 35 0 4.6 22.8 44.7 76.8 95.7

35 - 0 -1.0 19.2 27.7 82.1 88.1
65

65 - 0 -11.9 11.8 -2.7 69.5 89.7
95

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The reference for the publication: Ostroumov S.A. Inhibitory analysis of top-down control: new keys to studying
eutrophication, algal blooms, and water self-purification. Hydrobiologia. 2002. vol. 469, pages 117-129

Table 8. Effect of Triton X-100 on the clearance rate during filtering algae
Isochrysis galbana by mussels Mytilus edulis (after Ostroumov et al., 1998, with
some changes)

Concentration of Clearance rate, l h-1 (in brackets - % of control)
Triton X-100, during various time periods
mg l–1

0 - 60 min 60 - 120 min
0 (control) 9.53 (100%) 4.85 (100%)

4 1.04 (10.9%) 1.02 (21.0%)

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The reference for the publication: Ostroumov S.A. Inhibitory analysis of top-down control: new keys to studying
eutrophication, algal blooms, and water self-purification. Hydrobiologia. 2002. vol. 469, pages 117-129

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ADDENDUM
(This addendum was added after publishing this paper, when the paper was put at the website)

The relevant bibliography that includes a broader list of the author’s publications is given below.
After publishing this paper the author did several new research projects that confirmed the main
conclusions.
Additional evidence was obtained that the inhibition of water filtration by surfactants and
detergents was a typical response of a broad spectrum of filter – feeders including both marine and
freshwater bivalves [1], rotifers [24], and crustaceans [34]. Filtration activity of invertebrates is a
component of the contribution of aquatic biota to water self-purification. It is a part of ecosystem
services towards maintaining and improving water quality.
The results of those detailed studies that used a variety of filter-feeders (suspension feeders) were
summarized in the monographs [1, 36-38]. Those publications got positive evaluations in the
reviews ([39-54] and others), and won several awards and diplomas (the list of them see below,
after the list of the publications).
All in all, this paper together with other publications, including the more recent ones, contribute to
better understanding of how aquatic biota participate in ecosystem services towards environmental
and ecological stability, as well as environmental safety. The role of biota is necessary to
understand and to protect in order to maintain the sustainable use of aquatic and biological
resources.
1. Ostroumov S. A. Biological Effects of Surfactants. CRC Press. Taylor & Francis. Boca
Raton, London, New York. 2006. 279 p. ISBN 0-8493-2526-9 [new facts and concepts on
assessment of hazards from chemicals, new look on the factors important to water quality,
to sustainability; new priorities in environmental safety]

30
The reference for the publication: Ostroumov S.A. Inhibitory analysis of top-down control: new keys to studying
eutrophication, algal blooms, and water self-purification. Hydrobiologia. 2002. vol. 469, pages 117-129

2. Ostroumov S. A. Biological filtering and ecological machinery for self-purification and
bioremediation in aquatic ecosystems: towards a holistic view // Rivista di Biologia /
Biology Forum. 1998. V. 91(2). P.221-232.
3. Ostroumov S. A., Donkin P., Staff F. Filtration inhibition induced by two classes of
synthetic surfactants in the bivalve mollusk Mytilus edulis // Doklady Biological Sciences,
1998. Vol. 362, P. 454-456.
4. Ostroumov S. A. The concept of aquatic biota as a labile and vulnerable component of
the water self-purification system - Doklady Biological Sciences, Vol. 372, 2000, pp. 286–
289. http://sites.google.com/site/2000dbs372p286biotalabil/
5. Ostroumov S. A., Kolesnikov M. P. Biocatalysis of Matter Transfer in a Microcosm Is
Inhibited by a Contaminant: Effects of a Surfactant on Limnea stagnalis. - Doklady
Biological Sciences, 2000, 373: 397–399. Translated from Doklady Akademii Nauk, 2000,
Vol. 373, No. 2, pp. 278–280. http://sites.google.com/site/2000dbs373p397biocatallstag/
6. Ostroumov S. A. An aquatic ecosystem: a large-scale diversified bioreactor with a water
self-purification function. - Doklady Biological Sciences, 2000. Vol. 374, P. 514-516.
http://sites.google.com/site/2000dbs374p514bioreactor/
7. Ostroumov SA. Criteria of ecological hazards due to anthropogenic effects on the biota:
searching for a system. - Dokl Biol Sci (Doklady Biological Sciences). 2000; 371:204-206.
http://sites.google.com/site/2000dbs371p204criteria/
8. Ostroumov S. A. An amphiphilic substance inhibits the mollusk capacity to filter out
phytoplankton cells from water. - Biology Bulletin, 2001, Volume 28, Number 1, p. 95-
102.
[Biology Bulletin: ISSN 1062-3590 (Print) 1608-3059 (Online); DOI
10.1023/A:1026671024000; http://www.springerlink.com/content/l665628020163255/;]
9. Ostroumov S. A. Inhibitory Analysis of Regulatory Interactions in Trophic Webs.
-Doklady Biological Sciences, 2001, Vol. 377, pp. 139–141. Translated from Doklady
Akademii Nauk, 2000, Vol. 375, No. 6, pp. 847–849. [In the paper, the author developed a
new approach to analyze the fundamental ecological issue, the interactions between
organisms in ecosystems. He suggested to use the methodology of inhibitory analysis to
study interactions in trophic chains. Important situation is the top–down control of
plankton by benthic filter-feeders. This control, as author’s experiments have shown,
might be removed by chemical inhibitors (the latter may enter the ecosystem as
pollutants)]. http://sites.google.com/site/2001dbs377p139inhibitory/;
http://blog.researchgate.net/masterblog/299_Inhibitory_Analysis_A_new_method_for_an
alyzing_interactions_between_organisms_in_ecosystems;
10. Ostroumov SA. The synecological approach to the problem of eutrophication. - Dokl Biol
Sci. (Doklady Biological Sciences). 2001; 381:559-562.
11. Ostroumov SA. The hazard of a two-level synergism of synecological summation of
anthropogenic effects. - Dokl Biol Sci. (Doklady Biological Sciences). 2001; 380:499-501.
http://sites.google.com/site/2001dbs380p499synerg/
12. Ostroumov SA. Responses of Unio tumidus to mixed chemical preparations and the
hazard of synecological summation of anthropogenic effects. - Dokl Biol Sci (Doklady
Biological Sciences). 2001; 380: 492-495. http://sites.google.com/site/2001dbs380p492unio/
13. Ostroumov SA, Kolesnikov MP. Pellets of some mollusks in the biogeochemical flows of
C, N, P, Si, and Al. - Dokl Biol Sci (Doklady Biological Sciences). 2001; 379:378-381.
http://sites.google.com/site/2001dbs379p378pellets/

31
The reference for the publication: Ostroumov S.A. Inhibitory analysis of top-down control: new keys to studying
eutrophication, algal blooms, and water self-purification. Hydrobiologia. 2002. vol. 469, pages 117-129

14. Ostroumov SA. Imbalance of factors providing control of unicellular plankton
populations exposed to anthropogenic impact. - Dokl Biol Sci (Doklady Biological
Sciences). 2001; 379:341-343. http://sites.google.com/site/1dbs379p341imbalance/;
https://www.researchgate.net/profile/Sergei_Ostroumov/blog/321_Discovery_of_the_syste
m_where_chemical_pollution_could_impair_the_regulation_and_balance_in_the_abunda
nce_of_phytoplankton_threat_to_ecological_stability_
15. Ostroumov SA. Effect of amphiphilic chemicals on filter-feeding marine organisms. -
Dokl Biol Sci (Doklady Biological Sciences). 2001; 378:248-250. For the first time, oysters
were used as the test-organisms in bioassay of synthetic surfactants and detergents. The new
data revealed a novel type of hazard to mariculture. New negative effects of surfactants and
chemical mixtures on water filtering activity of Crassostrea gigas were discovered.
http://sites.google.com/site/2001dbs378p248effammaroyst/
16. Ostroumov S. A. Inhibitory analysis of top-down control: new keys to studying
eutrophication, algal blooms, and water self-purification // Hydrobiologia. 2002. vol. 469.
P.117-129.
17. Ostroumov S. A. Polyfunctional role of biodiversity in processes leading to water
purification: current conceptualizations and concluding remarks // Hydrobiologia. 2002.
v. 469 (1-3): P.203-204.
18. Ostroumov SA. Identification of a new type of ecological hazard of chemicals: inhibition
of processes of ecological remediation. - Dokl Biol Sci (Doklady Biological Sciences).
2002; 385:377-379.
19. Ostroumov SA. System of principles for conservation of the biogeocenotic function and
the biodiversity of filter-feeders. - Dokl Biol Sci (Doklady Biological Sciences). 2002;
383:147-150.
20. Ostroumov SA. A new type of effect of potentially hazardous substances: uncouplers of
pelagial-benthal coupling. - Dokl Biol Sci (Doklady Biological Sciences). 2002; 383:127-
130.
21. Ostroumov SA. Biodiversity protection and quality of water: the role of feedbacks in
ecosystems. - Dokl Biol Sci (Doklady Biological Sciences). 2002; 382:18-21.
22. Ostroumov S. A. Studying effects of some surfactants and detergents on filter-feeding
bivalves // Hydrobiologia. 2003. Vol. 500. P. 341-344.
23. Ostroumov S.A. Anthropogenic effects on the biota: towards a new system of principles
and criteria for analysis of ecological hazards. - Rivista di Biologia/Biology Forum. 2003.
96: 159-170. Review. PMID: 12852181 [PubMed - indexed for MEDLINE]
http://sites.google.com/site/ostroumovsergei/publications-1/rivista2003criteria
24. Ostroumov SA, Walz N, Rusche R. Effect of a cationic amphiphilic compound on rotifers.
- Dokl Biol Sci. (Doklady Biological Sciences) 2003; 390: 252-255.
25. Ostroumov S. A. On the biotic self-purification of aquatic ecosystems: elements of the
theory. - Doklady Biological Sciences, 2004, Vol. 396, Numbers 1-6, p. 206-211.
26. Ostroumov S. A. Suspension-feeders as factors influencing water quality in aquatic
ecosystems. - In: The Comparative Roles of Suspension-Feeders in Ecosystems, R.F.
Dame, S. Olenin (Eds), Springer, Dordrecht, 2004. p. 147-164.
27. Ostroumov S. A. Some aspects of water filtering activity of filter-feeders // Hydrobiologia,
2005. Vol. 542, No. 1. P. 275 – 286.

32
The reference for the publication: Ostroumov S.A. Inhibitory analysis of top-down control: new keys to studying
eutrophication, algal blooms, and water self-purification. Hydrobiologia. 2002. vol. 469, pages 117-129

28. Ostroumov S. A. On some issues of maintaining water quality and self-purification. -
Water Resources [(Vodnye Resursy) ISSN PRINT: 0097-8078; ISSN ONLINE: 1608-
344X; http://www.maik.rssi.ru/cgi-perl/journal.pl?name=watres&lang=eng]. 2005,
Volume 32, Number 3, p. 305-313.
29. Ostroumov S. A. On the multifunctional role of the biota in the self-purification of
aquatic ecosystems // Russian Journal of Ecology, Vol. 36, No. 6, 2005, p. 414–420.
30. Ostroumov S. A. Biomachinery for maintaining water quality and natural water self-
purification in marine and estuarine systems: elements of a qualitative theory //
International Journal of Oceans and Oceanography. 2006. Volume 1, No.1. p.111-118.
[ISSN 0973-2667]. Publisher: Research India Publications, Dehli]. Basic elements are
formulated for a qualitative theory of the polyfunctional role of the biota in maintaining
self-purification and water quality in aquatic ecosystems. The elements of the theory
covers the following: (1) sources of energy for the mechanisms of selfpurification; (2) the
main functional blocks of the system of self-purification; (3) the list of the main processes
that are involved; (4) analysis of the degree of participation of the main large taxons; (5)
degree of reliability and the main mechanisms providing the reliability; (6) regulation of
the processes; (7) the response of the system towards the external influences (man-made
impacts); (8) the analogy between ecosystems and a bioreactor; and (9) conclusions
relevant to the practice of biodiversity conservation. In support of the theory, results are
given of the author's experiments which demonstrated the ability of some pollutants
(surfactants, detergents, and some others) to inhibit the water filtration activity of marine
filter-feeders (namely, the bivalve mollusks Mytilus galloprovincialis, Mytilus edulis, and
Crassostrea gigas).
31. Ostroumov S. A., Widdows J. Inhibition of mussel suspension feeding by surfactants of
three classes. // Hydrobiologia. 2006. Vol. 556, No. 1. Pages: 381 – 386. 3 Tables. Bibliogr.
37 refs. [For the first time the negative effects of the three surfactants on the filtration
rates by marine mussels were presented in one paper. The xenobiotics tested represented
anionic, cationic and non-ionic surfactants (tetradecyltrimethylammonium bromide,
TDTMA, a representative of a class of cationic surfactants; sodium dodecyl sulphate,
SDS, a representative of anionic alkyl sulfates; and Triton X-100, a representative of non-
ionic hydroxyethylated alkyl phenols). Negative effects of SDS, TDTMA, and Triton X-
100 on the filtration activity of marine mussels M. edulis and M. edulis / M.
galloprovincialis were discovered. All three surfactants inhibited the clearance rates. This
is the first publication of the negative effects of a cationic surfactant on Atlantic mussels
Mytilus. The significance of the results for the ecology of marine ecosystems is discussed].
DOI 10.1007/s10750-005-1200-7; http://sites.google.com/site/ostroumovsergei/publications-
1/hydrobiologia2006ostwidd; http://sites.google.com/site/3surfactantsfiltrationmytilus/;
http://www.springerlink.com/content/7166067538534421/
32. Ostroumov S. A. Biotic self-purification of aquatic ecosystems: from the theory to
ecotechnologies. - Ecologica, 2007. vol. 15 (50), p.15-23. (ISSN 0354-3285; Belgrade). Some
basic elements of a new theory for the biological mechanism for water self-purification are
presented. Hydrobionts (aquatic organisms) are actively involved in various processes
leading to water purification. Not only microorganisms (bacteria, cyanobacteria and
fungi), but also algae, plants, invertebrates, and many other groups of organisms are
involved, which is discussed and analyzed in the paper. Results of the author's
experiments that study the effects of various pollutants on aquatic organisms (freshwater
and marine bivalves) are given. The theory is an innovative basis for developing ecological
technologies to clean water and to upgrade its quality by using organisms and ecosystems
[http://scindeks.nb.rs/article.aspx?artid=0354-32850750015O].

33
The reference for the publication: Ostroumov S.A. Inhibitory analysis of top-down control: new keys to studying
eutrophication, algal blooms, and water self-purification. Hydrobiologia. 2002. vol. 469, pages 117-129

33. Ostroumov S. A. Basics of the molecular-ecological mechanism of water quality formation
and water self-purification. - Contemporary Problems of Ecology, 2008, Vol. 1, No. 1, p.
147-152.
34. Vorozhun I. M., S. A. Ostroumov. On studying the hazards of pollution of the biosphere:
effects of sodium dodecylsulfate (SDS) on planktonic filter-feeders. - Doklady Biological
Sciences, 2009, Vol. 425, p. 133–134.
35. Ostroumov S.A. Towards the general theory of ecosystem-depended control of water
quality. - Ecologica, [ISSN 0354-3285], 2009, vol. 16, No. 54, p. 25-32. Abstract: A new set
of ecological generalizations formulated in this paper represent, in a systematized form,
the basic elements of the qualitative theory of biotic control of water quality and water
self-purification in freshwater and marine ecosystems. The theory contributes to a better
understanding of the issues of stability and regulation in the biosphere. The theory is
supported by the results of the author’s experimental studies of the effects exerted by
surfactants, detergents and other pollutants on aquatic organisms.
http://sites.google.com/site/9enecologica16p25theory/
36. Ostroumov S.A. Biotic Mechanism of Self-purification of Freshwater and Marine Water.
(Ecological Studies, Hazards, Solutions, vol. 9) Мoscow: МAX Press. 2004. IV. 96 p., tab.
Bibliogr. 59-85. Abstract in English. Section in English: p.53-58; ISBN 5-317-01120-5.
[Diploma of the Academy of Aquatic Sciences, awarded in 2006; another Diploma to the
book was awarded at the 7th International Conference ‘Aquatic Ecosystems, Organisms,
Innovations’ (2005)].
37. Ostroumov S.A. Pollution, Self-purification and Restoration of Aquatic Ecosystems.
Мoscow: МAX Press. 2005. 100 p., tab. ISBN 5-317-01213-9. (Diploma of the Academy
of Aquatic Sciences, awarded in 2006).
38. Ostroumov S.A. Aquatic organisms in water self-purification and biogenic migration of
elements. Moscow. MAX Press. 2008. 200 p. ISBN 978-5-317-02625-7.
39. Prof. Yakovlev S.V. (Full Member, Acad. Sci.) Review of the book: Ostroumov S.A.
Biological Effects of Surfactants on Organisms (2001) // Vestnik Russian Academy of
Sciences. [ISSN 0869-5873; http://www.maik.ru/cgi-bin/list.pl?page=vestnik ], 2002. v.72,
No.11, p. 1038-1047.
40. Prof. Vasiliev О.F. (Full Member, Acad. Sci.) Review of the book: Ostroumov S.A.
Biological Effects of Surfactants on Organisms (2001) // Vestnik Russian Academy of
Natural Sciences, [ISSN 1682-1696], 2002, v.2, No.3, p. 65.
41. Prof. Braginsky L.P., Prof. Sirenko L.A. Review of the book: Ostroumov S.A. Biological
Effects of Surfactants on Organisms (2001) // Hydrobiological Journal (published in
Kiev, Ukraine ISSN: for PRINT: 0018-8166). 2003, v.39, No.3, p.115-118.
42. Prof. Rozenberg G.S. (Corresponding Member of the Russian Academy of Sciences,
Director of the Institute of Ecology of the Volga Basin, Russ. Academy of Sciences)
Review of the book: Ostroumov S.A. Biological Effects of Surfactants on Organisms
(2001) // Uspekhi Sovremennoi Biologii (Advances of Modern Biology) [ISSN PRINT:
0042-1324; http://www.maik.ru/cgi-bin/list.pl?page=uspbio]. 2003. No. 6, p. 618-619.
43. Malakhov V.V. (Corresponding Member of the Russian Academy of Sciences; Chair of
the Dept. of Zoology of Invertebrates, Moscow State University), Review of the book
(Ostroumov, S.A., 2004. Biotic Mechanism of Self-purification of Freshwater and Marine
Water. MAX Press, Moscow) // Ecol. Stud. Haz. Solutions. 2004. Vol.10. p.138.

34
The reference for the publication: Ostroumov S.A. Inhibitory analysis of top-down control: new keys to studying
eutrophication, algal blooms, and water self-purification. Hydrobiologia. 2002. vol. 469, pages 117-129

44. Petrosyan V.S. Review of the book: Biological Effects of Surfactants. CRC Press. Taylor
& Francis. - Ecological Studies, Hazards, Solutions, 2007. vol. 12, p.117-119 (in English).
45. Review of the book: Ostroumov S.A. Biological Effects of Surfactants. CRC Press. Taylor
& Francis. Boca Raton, London, New York. 2006. 279 p. – Bulletin Samarskaya Luka
[http://www.ssc.smr.ru/ssc_sl.html] - 2007. - V. 16, No. 4(22). - P. 864-867. Bibliogr. 10
refs.; http://www.ssc.smr.ru/media/journals/samluka/2007/16_4_22.pdf;
46. Review of the book: Biological Effects of Surfactants. CRC Press. Taylor & Francis. Boca
Raton, London, New York. 2006. 279 p. // Problems of Biogeochemistry and Geochemical
Ecology. 2007. No. 2 (4). p.108.
47. Review of the book: S.A.Ostroumov. Biological Effects of Surfactants (2006). - Ecologica,
2008. т.15, No. 51, p. 71-72. (ISSN 0354-3285; in English).
48. Ermakov V.V. Review of the book: Ostroumov S.A. Biological Effects of Surfactants.
CRC Press. Taylor & Francis. Boca Raton, London, New York. 2006. 279 p. –
Toxicological Review [Toksikologicheskij Vestnik; ISSN 0869-7922;
http://www.rpohv.ru/magazin/], 2009, No. 2, p. 40.
49. Toderas I.K. (Academician-Secretary of the Section of the Biological, Chemical, and
Ecological Sciences of the Academy of Sciences of Moldova) et al. Novelty about ecological
hazards of the chemicals that pollute aquatic environment. A review of the book:
Ostroumov S.A. Biological Effects of Surfactants. CRC Press. Taylor & Francis. Boca
Raton, London, New York. 2006. 279 p.). – // Bulletin of the Academy of Sciences of
Moldova. Life Sciences (Buletinul Academiei de Stiinte a Moldovei. Stiintele Vietii; ISSN
1857-064X). 2007, No. 2, p.169-172. Bibliogr. 10 refs. ["The book is a new significant step
toward better knowledge and understanding the effects of chemical pollution on the
biosphere" (p. 172)].
50. Ecological sciences: from theory to practice and sustainable development. Review of the
books by S.A.Ostroumov 'Pollution, Self-purification and Restoration of Aquatic
Ecosystems' and 'Ecology and Hydrobiology'. - Ecological Systems and Instruments.
[ISSN 2072-9952; http://www.ecoguild.ru/docs/2007esip.htm] 2006. No. 4. P.38-39.
51. Prof. Rumyantzev I.S., Zimnyukov V.A. Review of the book: Ostroumov S.A. Pollution,
Self-Purification, and Restoration of Aquatic Ecosystems. Moscow. MAX Press, 2005, 100
p. ISBN 5-317-01213-9. – Ecological Chemistry [ISSN 0869-3498;
http://www.thesa.ru/ecology/], 2006, v. 15, No. 3, p. 211-212 [on the author: Prof. I.S.
Rumyantzev is the President, The Academy of Water Science].
52. Prof. Abakumov V.A. Review of the book "Pollution, Self-Purification, and Restoration
of Aquatic Ecosystems". Moscow. MAX Press, 2005, 100 p. – Ecology of the Environment
and Safety of Life Activity, [ISSN 1726-5428], No. 4 (34), 2006, p.88-89 [the reviewer
recommended to publish a new edition of the book].
53. Prof. Ermakov V.V. Review of the book "Pollution, Self-Purification, and Restoration of
Aquatic Ecosystems. Moscow. MAX Press, 2005, 100 p." and of the collection of
educational programs "Ecology and Hydrobiology" (Moscow. MAX Press, 2005). –
Problems of Biogeochemistry and Geochemical Ecology. 2007. v. 1(3). p.122-124.
54. Prof. Braginsky L.P., Kalenichenko K.P., Ignatyuk A.A. Generalized mechanisms of self-
purification of natural waters. Review of the book: Ostroumov S.A. Pollution, Self-
Purification, and Restoration of Aquatic Ecosystems. Moscow. MAX Press, 2005, 100 p. //
Hydrobiological Journal. [ISSN: for PRINT: 0018-8166] 2007, v. 43, No. 6. P.111-113.

35
The reference for the publication: Ostroumov S.A. Inhibitory analysis of top-down control: new keys to studying
eutrophication, algal blooms, and water self-purification. Hydrobiologia. 2002. vol. 469, pages 117-129

The publications above [1-38] and the results in them won some awards and Diplomas,
including:
Scientific Discovery Diploma No. 274 "Ability of synthetic surfactants to inhibit the filtration
activity of bivalves"; priority of the discovery 9 August, 2000, according to the date of publishing
the book: S.A.Ostroumov "Biological effects of surfactants in connection with the anthropogenic
impact on the biosphere" (2000, MAX Press, Moscow). Diploma of 31.01.2005, registration
number 332; (about the discovery: http://idea.emind.ru/discovery/show/67);
P.L.Kapitza Medal for the Scientific Discovery;
Diploma of the National Ecological Award 2004, for contribution to ecological security and
sustainable development of Russia;
Diploma of the First Degree for books "Biological effects of surfactants in connection with the
anthropogenic impact on the biosphere" and "Biological effects of surfactants on organisms". The
books were displayed at the exhibition of new books at the conference Aquatic Ecosystems and
Organisms (2004);
Diploma at the 1st International Forum on Conservancy of Nature (Moscow, 7-9th September
2005), signed by Co-Chairs of the Forum, Deputy Minister of Natural Resources of Russian
Federation and Vice-President of the Trade Chamber of Russian Federation).
Diploma of the journal 'Ecology and Life' (awarded 15 October 2005).
Diploma of the competition 'Sustainable use of natural resources and environmental protection –
strategy of sustainable development of Russia in the 21st century' at the international conference
'Sustainable Development: Nature-Society-Man' (organized in 2006, Moscow, by the Ministry of
Natural Resources) for the paper by Ostroumov S.A. and Solomonova E.A. "On studies of water
self-purification and interactions between pollutants (surfactants) and biota: searching approaches
to issues relevant to sustainable use of water resources " (Directive of the Ministry of Natural
Resources No.126 of 02 June 2006; the Diploma was signed by the Minister).
Diploma to the book 'Biotic Mechanism of Self-purification of Freshwater and Marine Water' by
Dr. S. A. Ostroumov, awarded at the 7th International Conference ‘Aquatic Ecosystems,
Organisms, Innovations’ (2005). Diploma of the Academy of Aquatic Sciences for the series of
innovative publications on aquatic ecology, interactions between chemicals and organisms, and
water self-purification, including the books 'Biological Effects of Surfactants in Connection with
the Anthropogenic Impact on the Biosphere'; 'Biological Effects of Surfactants on Organisms';
'Biotic Mechanism of Self-Purification of Freshwater and Marine Water'; 'Pollution, Self-
Purification and Restoration of Aquatic Ecosystems'; 'Biological Effects of Surfactants', (awarded
in 2006).
Diploma of the Academy of Aquatic Sciences 'for his innovative contribution to improvement of
environmental and ecological education', including the book 'Ecology and Hydrobiology. Curricula
of Lecture Courses' (awarded in 2006).
Diploma of the Moscow Society of the Researchers of Nature (awarded in 2007).

**
More recent publications, sites and posts which supported the conclusions and
discoveries presented in this paper:
http://www.researchgate.net/profile/Sergei_Ostroumov/blog/11203_New_links_amon
g_toxic_pollution_and_eutrophication; New links among toxic pollution and
eutrophication.
http://www.scribd.com/doc/51414359; Ecology. Key Innovations, Discoveries. The
material include two updated tables. They are based on the book [Ecology and
Life Sciences: Bibliography of S. A. Ostroumov. Compiled by A. V. Viktorova
and J. Robertson. 4th ed., updated. Мoscow: MAX-Press, 2010].
http://www.researchgate.net/profile/Sergei_Ostroumov/blog/9142_FAQBiotic_Self-
purification_of_Aquatic_Ecosystems; FAQ on the paper: S. A. Ostroumov. On
the Biotic Self-purification of Aquatic Ecosystems: Elements of the Theory.

36
The reference for the publication: Ostroumov S.A. Inhibitory analysis of top-down control: new keys to studying
eutrophication, algal blooms, and water self-purification. Hydrobiologia. 2002. vol. 469, pages 117-129

http://www.researchgate.net/profile/Sergei_Ostroumov/blog/6882_scribd_fulltexts:
Texts online free.
**

Address of Dr. S. A. Ostroumov:

Lab of Physico-Chemistry of Biomembranes,
Faculty of Biology,
Moscow State University,
Moscow 119991, Russia;
e-mail: ar55[at]yandex.ru

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