Doklady Biological Sciences, Vol. 379, 2001, pp. 378–381. Translated from Doklady Akademii Nauk, Vol.
379, No. 3, 2001, pp. 426–429.
Original Russian Text Copyright © 2001 by Ostroumov, Kolesnikov.
GENERAL BIOLOGY
Pellets of Some Mollusks in the Biogeochemical Flows
of C, N, P, Si, and Al
S. A. Ostroumov and M. P. Kolesnikov
Presented by Academician M.E. Vinogradov February 1, 2001
Received February 1, 2001
Aquatic invertebrates excrete pellets [1,2], which
consist of products of incomplete digestion and pseud-
ofeces. Different taxons are characterized by the fol-
lowing values of food assimilability (in percents):
Rotatoria, 48–80; Bryozoa, 41.6; Gastropoda, 42–82;
Bivalvia, 40–47; Cladocera, 50.5–85.5; Copepoda, 30–
88; Mysidacea, 84.2–95; Isopoda, 68; Amphipoda, 5.5–
98; Decapoda, 38.7–96.1; larvae of Odonata, 20–97.2;
Ephemeroptera, 41–72; Plecoptera, 9–73; Trichoptera,
5–51; and Diptera, 1–31.4 [2]. When settling to the bot-
tom of water bodies by gravity, pellets contribute to
vertical fluxes of chemical elements through the eco-
system level of the biospheric biogeochemical cycles or
into “biogenic migration of atoms in the biosphere” [3].
The purpose of this study was to estimate the capac-
ity of mollusk pellets for contributing into the vertical
transfer of chemical elements through an aquatic eco-
system, using Limnaea stagnalis (L.) and bivalves
(Unionidae) as examples.
The mollusks L. stagnalis were collected in June in
a pond in the floodplain of the upper Moskva River and
were kept afterwards as described in [4]. Bivalve mol-
lusks (Unionidae) collected from the partly silted sand
bottom of the Moskva River upstream of the town of
Zelinograd represented a sample from a natural benthic
community dominated by Unio tumidus and U. pic-
torum (63.21 and 27.36%, respectively, of the overall
number of specimens in the sample). The proportions
of Crassiana crassa and Anodonta cygnea were lower
(7.55 and 1.89%, respectively). The bivalves sampled
from the natural ecosystem (the total biomass was
3302 g wet weight, including the shells; the average
weight of one mollusk was 21.9 g), where they filtered
natural seston, were incubated for 24 h in a wide flask
containing settled tap water to obtain the pellet precip-
itate, which was afterwards resuspended in a 300-ml
glass cylinder. The sand fraction, which precipitated
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within 15 s (fraction 1), was separated from the remain-
ing suspension of the pellet material per se. The latter
was transferred into another glass cylinder. From this
suspension, pellets precipitated within 3 h (fraction 2);
this precipitate was separated by means of decanting the
supernatant. The dry weight of fraction 2 was 1434 mg.
To determine the carbon content in the vegetative
material and in pellets, they were oxidized by 10%
K2Cr2O7 in the presence of a mixture of concentrated
H2SO4 and H3PO4 [5]; the CO2 formed was trapped by
0.5 M NaOH, and the remaining alkaline was titered
with hydrochloric acid. The photometric method was
also used; the optic density of the solution was mea-
sured at a wavelength of 590 nm after oxidation of the
material with bichromate. The amount of organic nitro-
gen was measured using a Kjeltec Auto 1030 Analyzer
(Tecator, Sweden) by the Kjeldahl method after miner-
alization with a mixture of H2SO4 and H2O2. Phospho-
rus was assayed using phosphoromolybdenum blue
(PMB) in a working solution containing ions of ortho-
phosphate and orthosilicic acid in 15.0 ml of 1 N H2SO4
[6]. The sum of organogenic and soluble (mineral) sili-
con was determined using the modified silicomolybde-
num blue method without preliminary ashing of the
vegetative material [6]. Aluminum was determined in
an aliquot of HNO3 hydrolyzate (1.5–2.5 ml) neutral-
ized with ammonium and dissolved in water to a final
volume of 25.0 ml; afterwards, 2.0 ml of 1% ascorbic
acid were added and pH 2.5 was obtained. Then, 0.04%
aquatic solution of eriochromcyanogen R (pH 2.5) and
5.0 ml of CH3COONH4 (50% solution, pH 7.0) were
added. After addition of water to obtain a final volume
of 50.0 ml, the optical density was measured at 535 nm.
Chemical analysis is described in detail elsewhere [4, 6].
The element composition of L. stagnalis pellets
after the mollusks were fed on the leaves of Nuphar
lutea Smith and Taraxacum officinale Wigg is shown in
Table 1. The compositions of L. stagnalis pellets were
similar in both cases. The contents of N and P were
slightly higher in the pellets of L. stagnalis than in the
phytomass of N. lutea, whereas the content of Si was
noticeably higher than that in the phytomass of both
plant species that served as a food for the mollusks. Pel-
lets of the bivalves were similar in composition to those
 PELLETS OF SOME MOLLUSKS 379

Table 1. Element composition of pellets (% of dry weight) formed by the mollusk L. stagnalis feeding on the leaves of
N. lutea and T. officinale
N. lutea T. officinale
Element
leaves pellets leaves pellets
C – 69.4 – 67.5
N 2.11–2.19 2.83 2.57 2.89
P 0.34–0.39 0.5 0.44 0.48
Si 0.81–0.85 1.73 1.15 1.87
Al 0.043–0.047 0.054–0.059 0.076 0.094
The sum of ash elements 4.17 5.97 5.43 –
(ash content)
Note: The element composition of leaves (in percents of the leaf dry weight) is shown for comparison.

Table 2. Formation of pellets by the mollusk L. stagnalis feeding on the leaves of various plant species

Parameter N. lutea T. officinale

Wet weight of the biomass consumed, g 15.89 9.66

Dry weight of the pellets formed, mg 2434 1387
Dry weight of the pellets formed per 1 g of wet weight of the leaf mass 153.2 143.6
Number of mollusks 30 55
Incubation time, h 309 48
Note: The incubation conditions when N. lutea was used as food: 309 h, 22–24°C, eight flasks; the incubation conditions when T. officinale
was used as food: 48 h, 22–23°C, nine flasks.

Table 3. Transfer of matter and chemical elements (mg) with pellets of bivalves (a sample from a natural community of
Unionidae) per unit biomass of mollusks and per unit area of the ecosystem from which the mollusks were collected

mg/1 kg of mollusk biomass

Pellets and chemical elements mg/1 m2 of the ecosystem
(wet weight including shells)
(% of dry weight)
per day 120 days per day 120 days

Pellets (total dry weight, mg) 434.28 52114 717.00 86040

C 279.24 3509 461.03 55324
N 11.86 1423 19.57 2349
P 1.69 203 2.80 336
Si 4.95 594 8.17 981
Al 0.31 37 0.51 61
The sum of ash elements (ash content), 7.11 30.88 3705 50.98 6117
Note: Pellets (fraction 2) were obtained from mollusks (total biomass, 3302 g of wet weight with shells) collected from the area of 2 m 2
in August in the Moskva River. The element composition of pellets (% of dry weight) was as follows: C, 64.3; N, 2.73; P, 0.39;
Si, 1.14; Al, 0.071.

of L. stagnalis. Note that pellets of bivalves were char-
acterized by a somewhat higher ash content and lower
silicon content.
Based on data on L. stagnalis nutrition in a micro-
cosm, it was calculated that 143.6 to 153.2 mg of pellets
(dry weight) were formed per 1 g of consumed plant bio-
mass, and carbon accounts for as much as 67.5 to 69.7%
of this amount (from 96.9 to 106.8 mg) (Table 2). It is
also interesting that pellet formation occurred at a sim-
ilar rate when the mollusks were fed on the phytomass
of different plant species.

DOKLADY BIOLOGICAL SCIENCES Vol. 379 2001

380 OSTROUMOV, KOLESNIKOV

Table 4. Chemical elements (mg) with pellets of the mollusk L. stagnalis per unit biomass of mollusks and per unit area of
the ecosystem from which the mollusks were collected

mg/1 kg of mollusk biomass

mg/1 m2 of the ecosystem
(wet weight with shells)
Pellets and chemical elements, % of dry weight
per day 120 days per day 120 days

Pellets (total dry weight, mg) 3372.3 404676 178.42 21410

C 2351.8 282221 124.43 14931
N 95.44 11452 5.05 606
P 6.86 2023 0.89 107
Si 58.34 7001 3.09 370
Al 2.02 243 0.11 13
The sum of ash elements (ash content), 5.97 201.33 24160 10.65 1278
Note: Pellets from mollusks (total biomass, 158.72 g of wet weight with shells) were collected from an area of 3 m 2 in August in a pond
in the floodplain of the upper Moskva River. The element composition of pellets (% of dry weight) was as follows: C, 69.74; N, 2.83;
P, 0.5; Si, 1.73; Al, 0.06.

The data shown in Tables 3 and 4 can be used to cal-
culate both specific rates of element transfer (per 1 g of
mollusk biomass) and the magnitude of the transfer
(per unit area of the ecosystem). In L. stagnalis, the
specific rate of matter transfer (dry weight of pellets per
1 g of mollusk biomass) (Table 4) was significantly
higher than in bivalve mollusks (Table 3). However, the
magnitudes of transfer (as calculated per unit area of
the ecosystem) differed less, because the biomass den-
sity of bivalves (per 1 m2) was significantly higher than
that of L. stagnalis. However, this is only a conservative
estimate, because, in natural ecosystems, the mollusk
biomass density considerably varies, and, hence, the
matter transfer also varies. Therefore, we regard these
estimates with caution and do not extrapolate them to
other ecosystems or other areas of the same ecosystem
that differ in the mollusk population density.
In previous experiments with microcosms, we stud-
ied the effect of the surfactant tetradecyltrimethylam-
monium bromide (TDTMA) on the trophic activity of
L. stagnalis, which was measured from the consump-
tion of leaf phytomass for three days of incubation [4].
At the surfactant concentration of 2 mg/l, the feeding
rate decreased with time, as well as the pellet excretion
by the mollusks. The inhibition of trophic activity
ranged from 27.9 to 70.9%, depending on time. Under
the exposure to TDTMA, formation and total accumu-
lation of pellets on the bottom per unit biomass of mol-
lusks was only 58.3% of the control value [4]. In simi-
lar experiments, the surfactant sodium dodecylsulfate
(SDS) also inhibited the trophic activity of L. stagnalis
feeding on T. officinale biomass during 17 h. The
degree of inhibition was 52.2 and 44.9% at SDS con-
centrations of 1 and 2 mg/l, respectively. Similarly, the
synthetic detergent Tyde-Lemon inhibited the trophic
activity of L. stagnalis feeding on the same plant spe-
cies. During 23 h of incubation, the degree of inhibition
was 36% at the detergent concentration of 75 mg/l.
Phytomass consumption was estimated per unit biomass
of the mollusks (per 1 g of wet weight of L. stagnalis).
Thus, trophic activity of molluscs and the related
transfer of matter and energy along the trophic chain
(phytomass–phytophage–excreted pellets) are inhib-
ited by surfactants. These results, as well as other data
[7–12], indicate one more potential anthropogenic haz-
ard to biogeochemical flows in the biosphere.
ACKNOWLEDGMENTS
We are grateful to V.D. Fedorov, A.F. Alimov,
V.V. Malakhov, E.A. Kuznetsov, M. V. Chertoprud, and
other researchers at the Moscow State University and
Russian Academy of Sciences for their help and useful
comments on this study.
The work of S.A. Ostroumov was supported by the
Open Society Support Foundation (RSS grant
no. 1306/1999).
RFERENCES
1. Alimov, A.F., Funktsional’naya ekologiya presnovod-
nykh dvustvorchatykh mollyuskov (Functional Ecology
of Fresh-Water Bivalve Mollusks), Leningrad: Nauka,
1981.
2. Monakov, A.V., Pitanie presnovodnykh bespozvonoch-
nykh (Feeding of Freshwater Invertebrates), Moscow,
1998.
3. Vernadskii, V.I., Khimicheskoe stroenie biosfery Zemli i
ee okruzheniya (Chemical Structure of the Biosphere of
the Earth and Its Surroundings), Moscow: Nauka, 1965.
4. Ostroumov, S.A. and Kolesnikov, M.P., Dokl. Akad.
Nauk, 2000, vol. 373, no. 2, pp. 278–280.

DOKLADY BIOLOGICAL SCIENCES Vol. 379 2001

 PELLETS OF SOME MOLLUSKS
5. Dalal, R.C., Analyst (Cambridge, UK), 1979, vol. 104,
pp. 151–154.
6. Kolesnikov, M.P. and Abaturov, B.D., Usp. Sovrem.
Biol., 1997, vol. 117, no. 5, pp. 534–548.
7. Ostroumov, S.A., Biologicheskie effekty poverkhnostno-
aktivnykh veshchestv v svyazi s antropogennymi vozde-
istviyami na biosferu (Biological Effects of Surfactants
in Connection with the Anthropogenic Impact of the
Biosphere), Moscow: MAKS, 2000.
DOKLADY BIOLOGICAL SCIENCES Vol. 379 2001
381
8. Ostroumov, S.A., Donkin, P., and Staff, F., Dokl. Akad.
Nauk, 1998, vol. 362, no. 4, pp. 574–576.
9. Ostroumov, S.A., Dokl. Akad. Nauk, 2000, vol. 372,
no. 2, pp. 279–282.
10. Ostroumov, S.A., Dokl. Akad. Nauk, 2000, vol. 374,
no. 3, pp. 427–429.
11. Ostroumov, S.A., Dokl. Akad. Nauk, 2000, vol. 371,
no. 6, pp. 844–846.
12. Ostroumov, S.A., Dokl. Akad. Nauk, 2000, vol. 375,
no. 6, pp. 847–849.