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Applied Soil Ecology 16 2001 97–107 The influence of dietary nitrogen and phosphorus on Cd accumulation in the woodlouse Porcellio scaber Latr Andrei D. Pokarzhevskii a,∗ , Nico M. van Straalen b a Institute of Ecology and Evolution of the Russian Academy of Sciences, Leninsky prospekt 33, 117071 Moscow, Russia b Institute of Ecological Science, Vrije Universiteit, De Boelelaan 1087, 1081 HV Amsterdam, The Netherlands Received 11 February 1999; received in revised form 20 July 2000; accepted 20 July 2000 Abstract Bioaccumulation of potentially toxic metals by litter-feeding invertebrates is determined not only by the concentration of metal in the diet, but also by the flux of food through the body. Nitrogen and phosphorus are the main elements regulating food consumption and so are expected to affect the bioaccumulation of trace elements such as cadmium. To test this idea, we applied a three-factor orthogonal experimental design to estimate the effects of nitrogen, phosphorus and cadmium additions to the food on cadmium accumulation by the terrestrial isopod Porcellio scaber. Cd, N and P were added to milled poplar litter in concentrations of 0, 10 and 20 mg g − 1 for cadmium, 0, 0.875 and 1.75 for nitrogen and 0, 0.2 and 0.4 for phosphorus. Observations were made for daily food consumption estimated from faecal pellet production and weekly Cd accumulation over a period of 4 weeks. Dietary Cd decreased consumption in the first 2 weeks of the experiment, but this effect disappeared later. Phosphorus had a significant positive effect on consumption in the second and the third week of the experiment. Accumulation of Cd was determined mainly by the Cd concentration in the food, but the effect was stimulated by P and diminished by N additions. Analysis of the data by regression using response surfaces confirmed that N and P had opposite influences on both Cd accumulation and food consumption. The data underline the importance of measuring consumption and food quality when conducting bioaccumulation and ecotoxicity experiments with soil invertebrates exposed through the diet. © 2001 Elsevier Science B.V. All rights reserved. Keywords: Bioaccumulation; Cadmium; Heavy metals; Isopoda; Nutrition; Soil invertebrates

1. Introduction

Food is an important source of contaminants as well as nutrients for soil animals. An elevated con- centration of heavy metals in food commonly leads to an increased concentration in the body Reichle et al., 1970; Martin and Coughtrey, 1982; Pokarzhevskii, 1985; Hopkin, 1989. However, metal accumulation depends not only on the concentration in food but ∗ Corresponding author. Tel.: +7-095-1359816; fax: +7-095-9545534. E-mail address: apokarzhonline.ru A.D. Pokarzhevskii. also on the rate of food passage through the body. Metal accumulation should be considered as a balance between input and output processes Pokarzhevskii, 1985; Hopkin, 1989; Fågerström, 1991. Nitrogen and phosphorus are the main elements determining animal productivity and food require- ments for animals, including invertebrates Dunger, 1958a,b; Prosser, 1973; House, 1974; Satchell, 1983; Gunnarsson, 1987. Within certain limits, low con- centrations of N and P in the food will stimulate con- sumption, as is well known from livestock farming practice. We have suggested earlier Pokarzhevskii, 1985; Pokarzhevskii and Van Straalen, 1996 that N 0929-139301 – see front matter © 2001 Elsevier Science B.V. All rights reserved. PII: S 0 9 2 9 - 1 3 9 3 0 0 0 0 1 1 2 - 8 98 A.D. Pokarzhevskii, N.M. van Straalen Applied Soil Ecology 16 2001 97–107 and P regulate the passage of food through the animal body and that hence these elements could also affect heavy metal uptake and accumulation. To test the idea of N and P regulating trace metal accumulation it is necessary to manipulate dietary N and P concentrations at a constant level of heavy metal. In most studies these factors are considered separately. This is due to the problem of how to deter- mine the influence of more than two interacting fac- tors simultaneously in a single experiment. A suitable method for such an experiment is a multi-factorial design. This approach has been used in many fields of science especially in the determination of optimal con- ditions for technological processes including indus- trial microbiology and agrochemistry Nalimov and Chernova, 1965; Adler et al., 1973; Mead, 1988. We have used the method for ecotoxicological and radioe- cological studies some years ago Terytze et al., 1989; Pokarzhevskii et al., 1993. A similar approach was applied by Loch et al. 1993 to assess the influence of different factors on Cr accumulation by ryegrass. Multifactorial response surface analysis assumes that the distances between the levels of a factor are equally spaced and can be represented by codes such as −1, 0 and +1. This simplifies the construction of orthogonal regression models of interactions between factors. Soil animals such as earthworms, woodlice and diplopods are good subjects for testing the hypothesis concerning the influence of N and P on trace metal accumulation. These invertebrates consume soil and litter and, although the attractiveness of the food de- pends on microbial infestation, they do not pick out particular species growing on the food bacteria, mi- crofungi, etc., as do microarthropods. This allows N and P compounds to be added directly to soil or litter without taking into account reallocations within the food, as in the case with microarthropods. In a pilot study Pokarzhevskii et al., 1994 we found that N and P influenced Cu accumulation in the earthworm Octolasium lacteum Oerley. Woodlice Crustacea: Isopoda: Oniscoidea are convenient animals for such ecotoxicological experi- ments as conditions for their rearing and breeding and their physiological features are well known Sutton, 1972; Warburg, 1987; Hopkin, 1989; Van Straalen and Donker, 1994. They survive well in captivity, readily consume litter including contaminated materials and are one of the best investigated soil invertebrates in ecotoxicology Drobne, 1997. Hopkin et al. 1993 suggested the use of the woodlouse Porcellio scaber Latreille as a monitor of bioavailability of metals in terrestrial ecosystems. Based on the hypothesis formulated above, we ex- pected that manipulation of the N and P contents in the food would affect accumulation of trace metals. We conducted a multi-factorial experiment, exposing isopods to cadmium-contaminated food at various levels of N and P, to test this expectation.

2. Materials and methods

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