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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.


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 within 15 s (fraction 1), was separated from the remain-
consist of products of incomplete digestion and pseud- ing suspension of the pellet material per se. The latter
ofeces. Different taxons are characterized by the fol- was transferred into another glass cylinder. From this
lowing values of food assimilability (in percents): suspension, pellets precipitated within 3 h (fraction 2);
Rotatoria, 48–80; Bryozoa, 41.6; Gastropoda, 42–82; this precipitate was separated by means of decanting the
Bivalvia, 40–47; Cladocera, 50.5–85.5; Copepoda, 30– supernatant. The dry weight of fraction 2 was 1434 mg.
88; Mysidacea, 84.2–95; Isopoda, 68; Amphipoda, 5.5– To determine the carbon content in the vegetative
98; Decapoda, 38.7–96.1; larvae of Odonata, 20–97.2; material and in pellets, they were oxidized by 10%
Ephemeroptera, 41–72; Plecoptera, 9–73; Trichoptera, K2Cr2O7 in the presence of a mixture of concentrated
5–51; and Diptera, 1–31.4 [2]. When settling to the bot- H2SO4 and H3PO4 [5]; the CO2 formed was trapped by
tom of water bodies by gravity, pellets contribute to 0.5 M NaOH, and the remaining alkaline was titered
vertical fluxes of chemical elements through the eco- with hydrochloric acid. The photometric method was
system level of the biospheric biogeochemical cycles or also used; the optic density of the solution was mea-
into “biogenic migration of atoms in the biosphere” [3]. sured at a wavelength of 590 nm after oxidation of the
The purpose of this study was to estimate the capac- material with bichromate. The amount of organic nitro-
ity of mollusk pellets for contributing into the vertical gen was measured using a Kjeltec Auto 1030 Analyzer
transfer of chemical elements through an aquatic eco- (Tecator, Sweden) by the Kjeldahl method after miner-
system, using Limnaea stagnalis (L.) and bivalves alization with a mixture of H2SO4 and H2O2. Phospho-
(Unionidae) as examples. rus was assayed using phosphoromolybdenum blue
(PMB) in a working solution containing ions of ortho-
The mollusks L. stagnalis were collected in June in phosphate and orthosilicic acid in 15.0 ml of 1 N H2SO4
a pond in the floodplain of the upper Moskva River and [6]. The sum of organogenic and soluble (mineral) sili-
were kept afterwards as described in [4]. Bivalve mol- con was determined using the modified silicomolybde-
lusks (Unionidae) collected from the partly silted sand num blue method without preliminary ashing of the
bottom of the Moskva River upstream of the town of vegetative material [6]. Aluminum was determined in
Zelinograd represented a sample from a natural benthic an aliquot of HNO3 hydrolyzate (1.5–2.5 ml) neutral-
community dominated by Unio tumidus and U. pic- ized with ammonium and dissolved in water to a final
torum (63.21 and 27.36%, respectively, of the overall volume of 25.0 ml; afterwards, 2.0 ml of 1% ascorbic
number of specimens in the sample). The proportions acid were added and pH 2.5 was obtained. Then, 0.04%
of Crassiana crassa and Anodonta cygnea were lower aquatic solution of eriochromcyanogen R (pH 2.5) and
(7.55 and 1.89%, respectively). The bivalves sampled 5.0 ml of CH3COONH4 (50% solution, pH 7.0) were
from the natural ecosystem (the total biomass was added. After addition of water to obtain a final volume
3302 g wet weight, including the shells; the average of 50.0 ml, the optical density was measured at 535 nm.
weight of one mollusk was 21.9 g), where they filtered Chemical analysis is described in detail elsewhere [4, 6].
natural seston, were incubated for 24 h in a wide flask
containing settled tap water to obtain the pellet precip- The element composition of L. stagnalis pellets
itate, which was afterwards resuspended in a 300-ml after the mollusks were fed on the leaves of Nuphar
glass cylinder. The sand fraction, which precipitated 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
Moscow State University, Vorob’evy gory, Moscow, phytomass of N. lutea, whereas the content of Si was
119899 Russia noticeably higher than that in the phytomass of both
Bakh Institute of Biochemistry, Russian Academy of Sciences, plant species that served as a food for the mollusks. Pel-
Leninskii pr. 33, Moscow, 117071 Russia lets of the bivalves were similar in composition to those

0012-4966/01/0708-0378$25.00 © 2001 MAIK “Nauka /Interperiodica”

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


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- cies. During 23 h of incubation, the degree of inhibition
culate both specific rates of element transfer (per 1 g of was 36% at the detergent concentration of 75 mg/l.
mollusk biomass) and the magnitude of the transfer Phytomass consumption was estimated per unit biomass
(per unit area of the ecosystem). In L. stagnalis, the of the mollusks (per 1 g of wet weight of L. stagnalis).
specific rate of matter transfer (dry weight of pellets per Thus, trophic activity of molluscs and the related
1 g of mollusk biomass) (Table 4) was significantly transfer of matter and energy along the trophic chain
higher than in bivalve mollusks (Table 3). However, the (phytomass–phytophage–excreted pellets) are inhib-
magnitudes of transfer (as calculated per unit area of ited by surfactants. These results, as well as other data
the ecosystem) differed less, because the biomass den- [7–12], indicate one more potential anthropogenic haz-
sity of bivalves (per 1 m2) was significantly higher than ard to biogeochemical flows in the biosphere.
that of L. stagnalis. However, this is only a conservative
estimate, because, in natural ecosystems, the mollusk
biomass density considerably varies, and, hence, the ACKNOWLEDGMENTS
matter transfer also varies. Therefore, we regard these We are grateful to V.D. Fedorov, A.F. Alimov,
estimates with caution and do not extrapolate them to V.V. Malakhov, E.A. Kuznetsov, M. V. Chertoprud, and
other ecosystems or other areas of the same ecosystem other researchers at the Moscow State University and
that differ in the mollusk population density. Russian Academy of Sciences for their help and useful
In previous experiments with microcosms, we stud- comments on this study.
ied the effect of the surfactant tetradecyltrimethylam- The work of S.A. Ostroumov was supported by the
monium bromide (TDTMA) on the trophic activity of Open Society Support Foundation (RSS grant
L. stagnalis, which was measured from the consump- no. 1306/1999).
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 RFERENCES
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