Earthworms of different functional groups affect the fate of the Bt-toxin Cry1Ab from transgenic maize in soil

The fate of the insecticidal Cry1Ab protein from crop residues (leaves and roots) of the transgenic maize variety MON810 was studied in the presence and absence of two earthworm species (Lumbricus terrestris, Aporrectodea caliginosa; separate incubations) in soil microcosms. The recombinant Cry1Ab protein was quantified using a highly sensitive ELISA. Control microcosms received corresponding non-transgenic plant material. All earthworms survived in the microcosms over a period of 5 weeks, irrespective of whether they received MON810 or non-transgenic plant material. Weight loss was observed for both earthworm species, independent of the plant material or transgenic modification. A strong decline of immunoreactive Cry1Ab in plant residues (mean initial concentration approx. 5000 ng g⁻¹) of MON810 was observed in all treatments, but in microcosms with earthworms this decline was significantly higher with less than 10% of the initial Cry1Ab concentration remaining after 5 weeks. Cry1Ab concentrations in casts were only 0.1% of those found in remaining plant material of the respective microcosms. No immunoreactive Cry1Ab proteins were found in earthworm tissues (threshold of detection: 0.58 ng g⁻¹ fresh weight). No further decline was found for Cry1Ab concentrations in casts of A. caliginosa during a subsequent period of 3 months of incubation in bulk soil (<0.1 ng g⁻¹) after removal of the earthworms from the microcosms, while in casts of L. terrestris the concentration decreased from 0.4 to below 0.1 ng g⁻¹. In conclusion, this study demonstrates that earthworms enhance the decline of immunoreactive Cry1Ab proteins from maize residues.     

Effects of biomass and size of Millsonia anomala (Oligochaeta: Acanthodrilidae) on particle aggregation in a tropical soil in the presence of Panicum maximum (Gramineae)

Summary The present study assessed the effect of the tropical geophagous earthworm, Millsonia anomala, on the aggregate-size distribution of a sieved (2 mm), tropical ferruginous soil in the presence and absence of the perennial tropical grass Panicum maximum. The effect of two size groupings and graded biomass densities of M. anomala on soil aggregation was analysed in time and with depth in the containers within which the plants were grown. In the absence of earthworms, aggregation was rapid although limited (13.5% of soil as aggregates >2.0 mm), and probably arose from a combination of microbial activity and physical processes (interparticle bonding due to clays and other colloids). The roots of the test species contributed little to aggregation. In contrast, the effect of earthworm activity on soil aggregation was rapid and important. The effects of both biomass and, to a lesser extent, size were significant. After 79 days, aggregation reached a maximum with 3 g per container of small earthworms (ca. 59% of soil as aggregates >2.0 mm) and a minimum with 1 g per container of large earthworms (ca. 35% of aggregates >2.0 mm). Aggregation was considered to occur through three different mechanisms: (1) A rapid aggregation due to the interactions of colloids; (2) an intermediate aggregation due to a combination of unquantified processes related to earthworm activity (mucus secretion, development of fungus hyphae); (3) egestion of soil as earthworm casts which are stable macroaggregates.     

The role of the earthworm Lumbricus terrestris in the transport of bacterial inocula through soil

Two laboratory experiments were used to investigate the effect of the earthworm Lumbricus terrestris on transport of genetically marked Pseudomonas fluorescens inocula through soil microcosms. The microcosms comprised cylindrical cores of repacked soil with or without earthworms. Late log-phase cells of P. fluorescens, chromosomally marked with lux genes encoding bioluminescence, were applied to the surface of soil cores as inoculated filter paper discs. In one experiment, 5 and 10 days after inoculation, cores were destructively harvested to determine concentrations of marked pseudomonads with depth relative to the initial inoculum applied. Transport of the bacteria occurred only in the presence of earthworms. In a second experiment cores were subjected to simulated rainfall events 18 h after inoculation with lux-marked bacteria at 3-day intervals over a 24-day period. Resulting leachates were analysed for the appearance of the marked bacteria, and after 28 days cores were destructively harvested. Although some marked cells (less than 0.1% of the inoculum applied) were leached through soil in percolating water, particularly in the presence of earthworms, the most important effect of earthworms on cell transport was through burial of inoculated litter rather than an increase in bypass flow due to earthworm channels.     

Influence of earthworms and leaf litter on edaphic variables in sewage-sludge-treated soil microcosms

Summary A series of 48 greenhouse soil microcosms were established and treated with combinations of sewage sludge, Acer saccharum leaves, and the earthworms Eisenia fetida and Lumbricus terrestris. The microcosms were incubated at constant moisture for 110 days. Samples were then taken and analysed for sludge depth, organic-matter content, and waterstable soil aggregates. Weights of surface leaves and weights and numbers of surviving earthworms were determined for each microcosm. L. terrestris significantly reduced sludge depth and the surface organic-matter content of microcosm soil and significantly increased percentages of 4-mm diameter water-stable aggregates. Leaf litter also significantly reduced sludge depth and increased 4-mm water-stable aggregates. E. fetida inhibited surface feeding by L. terrestris, reduced its 110-day survival rate, and inhibited the production of 4-mm water-stable aggregates in L. terrestris treatments. Numbers of E. fetida increased in L. terrestris treatments. Sludge depth, organic-matter content and water-stable aggregates were not significantly different from controls in E. fetida treatments.     

The impact of earthworms on the abundance of Collembola: improvement of food resources or of habitat?

I assessed the direct influence of earthworm excretions, and the impact of earthworms through their action on the soil structure (increased macroporosity), on the population dynamics of the collembolan species Heteromurus nitidus. The intestinal content of Collembola arising from cultures on different soil types was observed, and two experimental cultures of H. nitidus were run: (1) a culture performed on an inert substrate supplied either with earthworm casts or with soil as food resource, (2) an experiment using microcosms with cores of two humus forms (moder and calcic mull), in the presence or absence of earthworms. The observation of gut contents revealed that H. nitidus feeds on excrements, the composition of which (ratio organic matter/mineral matter) varies according to the humus form where it lived. Slightly aged (10–15 days) organo-mineral casts of earthworms appeared to be a better food than calcic mull aggregates or organic material from moder. Densities of H. nitidus cultured in cores of calcic mull were higher than in moder, except when cores of moder were inhabited by an anecic earthworm for 2 months. The humus form strongly influenced populations of H. nitidus, firstly because densities of predators were higher in moder than in calcic mull, and probably also because of soil macroporosity. It was concluded that earthworms would affect predation on H. nitidus by creating a network of interconnected macropores in which Collembola can move and find shelter.     

Changes in potassium availability and other soil properties due to soil ingestion by earthworms

An incubation experiment was conducted to study the changes that occur in potassium availability and other soil properties with ingestion of soil by earthworms. Two soils were used. Raumai soil with high non-exchangeable K and Milson soil with low non-exchangeable K were incubated with two species of earthworm, Aporrectodea caliginosa and Lumbricus rubellus, for 8 weeks. The casts and soil samples were analysed for exchangeable K, Ca, Mg, Na, and H, pH, organic C, and texture. The results indicated that in Raumai soil, the exchangeable K levels of the casts of both earthworm species were significantly higher than for the control soil, the effect being more marked for L. rubellus than for A. caliginosa. In Milson soil, the exchangeable K levels were significantly lower in the casts of both types of earthworm than in the control soil. The nitric acid-extractable K of the soil and casts was not markedly different for either soil type, but available non-exchangeable K values were significantly higher for the casts of L. rubellus from Milson soil than for the noningested Milson soil. In Raumai soil, the exchangeable Ca was higher in the casts of L. rubellus, exchangeable Mg and H were reduced, and exchangeable Na did not change markedly in the cast compared to the control soil. For Milson soil, the casts contained lower exchangeable Ca and H but higher Na and Mg than the control. The casts of both species of earthworm had significantly higher pH values for both soil types. There was no marked difference in the organic C content of the control soil and cast samples for Milson but a reduction in the casts of A. caliginosa for the Raumai soil. Finer fractions increased in the casts of both earthworm species in both soil types.     

Effects of the earthworm Pontoscolex corethrurus on banana plants infected or not with the plant-parasitic nematode Radopholus similis

Radopholus similis is a worldwide endoparasitic nematode that greatly hampers banana (Musa acuminata, Cavendish subgroup) productivity. Earthworms are known to closely interact with above-ground and under-ground soil biota and particularly with plants and microfaunal communities. This study was aimed at investigating, under greenhouse conditions, the effects of the earthworm Pontoscolex corethrurus on banana growth and nutrient uptake, and assessing the influences of this earthworm on the development of an inoculated population of R. similis. Six-week-old tissue culture banana plants were submitted to four treatments: with P. corethrurus, R. similis, P. corethrurus+R. similis, and a control with no earthworms or nematodes. At the end of the experiment, the P. corethrurus treatments showed significantly higher leaf surface areas, shoot dry root weights, and root fresh weights than those without earthworms. This root growth enhancement probably contributed to the evident but non-significant decrease in the density of nematodes in the roots, even though earthworms did not reduce the total number of nematodes per whole root system. Moreover, the presence of earthworms slightly alleviated the severity of root damage. N bioavailability in the soil, along with N, Ca, and Mg content of banana plants, were also significantly increased in the presence of earthworms. Our results demonstrated that banana plant growth and nutrition were positively influenced by earthworms. Cropping practices that boost the development of earthworm communities in soil should therefore be promoted to enhance sustainability and to naturally alleviate nematode impact.     

Competition between invasive earthworms (Amynthas corticis, Megascolecidae) and native North American millipedes (Pseudopolydesmus erasus, Polydesmidae): Effects on carbon cycling and soil structure

Invasive earthworms can have significant impacts on C dynamics through their feeding, burrowing, and casting activities, including the protection of C in microaggregates and alteration of soil respiration. European earthworm invasion is known to affect soil micro- and mesofauna, but little is known about impacts of invasive earthworms on other soil macrofauna. Asian earthworms (Amynthas spp.) are increasingly being reported in the southern Appalachian Mountains in southeastern North America. This region is home to a diverse assemblage of native millipedes, many of which share niches with earthworm species. This situation indicates potential for earthworm-millipede competition in areas subject to Amynthas invasion.In a laboratory microcosm experiment, we used two ¹³C enriched food sources (red oak, Quercus rubra, and eastern hemlock, Tsuga canadensis) to assess food preferences of millipedes (Pseudopolydesmus erasus), to determine the effects of millipedes and earthworms (Amynthas corticis) on soil structure, and to ascertain the nature and extent of the interactions between earthworms and millipedes. Millipedes consumed both litter species and preferred red oak litter over eastern hemlock litter. Mortality and growth of millipedes were not affected by earthworm presence during the course of the experiment, but millipedes assimilated much less litter-derived C when earthworms were present.Fauna and litter treatments had significant effects on soil respiration. Millipedes alone reduced CO₂ efflux from microcosms relative to no fauna controls, whereas earthworms alone and together with millipedes increased respiration, relative to the no fauna treatment. CO₂ derived from fresh litter was repressed by the presence of macrofauna. The presence of red oak litter increased CO₂ efflux considerably, compared to hemlock litter treatments.Millipedes, earthworms, and both together reduced particulate organic matter. Additionally, earthworms created significant shifts in soil aggregates from the 2000-250 and 250-53 μm fractions to the >2000 μm size class. Earthworm-induced soil aggregation was lessened in the 0-2 cm layer in the presence of millipedes. Earthworms translocated litter-derived C to soil throughout the microcosm.Our results suggest that invasion of ecosystems by A. corticis in the southern Appalachian Mountains is unlikely to be limited by litter species and these earthworms are likely to compete directly for food resources with native millipedes. Widespread invasion could cause a net loss of C due to increased respiration rates, but this may be offset by C protected in water-stable soil aggregates.     

Mucus excretion and carbon turnover of endogeic earthworms

Summary Mucus excretion of endogeic earthworms, by the body surface to burrow walls and by the intestine to casts, was investigated using uniformly ¹⁴C-labelled adult or subadult specimens of Octolasion lacteum (Örley) in laboratory incubations in soil from a beechwood on limestone. The daily loss of C due to mucus excretion from the body surface and in casts was calculated as 0.2 and 0.5% of total animal C, respectively. The C loss due to mucus excretion by subadult or adult individuals of O. lacteum is assumed to account for 63% of total C losses (including mucus excretion and respiration) of the earthworms. In a second experiment we studied the incorporation of ¹⁴C from labelled soil, again from a beechwood on limestone, into the tissue of the endogeic earthworm species Aporrectodea caliginosa (Savigny). The results of this experiment indicate the existence of two C pools, one more labile and one more stable, in earthworms. It is assumed that the C investment for respiration and mucus excretion is derived from the labile C pool of endogeic earthworms.     

Effect of food type and placement on earthworm (Aporrectodea tuberculata) burrowing and soil turnover

Subsurface-dwelling Aporrectodea tuberculata, a common earthworm in Upper-Midwest (USA) agricultural fields, may be a significant component of agroeco-systems with regard to soil mixing and preferential transport of water and chemicals. In this study we looked at effects of food residue placement and food type on A. tuberculata burrowing and soil turnover in two-dimensional Evans box microcosms. Four food residue placements mimicked patterns induced by primary tillage and two food types, readily available and natural food sources, with no food as a control. An average earthworm population of 100 earthworms m^-2 was calculated to generate 1058 km ha^-1 of new burrows and turnover 7.9 Mg ha^-1 of soil in 1 week of activity at 20°C. Burrowing was random until food sources were encountered, at which time burrowing appeared to center around the food source.     

Influence of earthworm activity and rice straw application on the soil microbial community structure analyzed by PLFA pattern

The earthworm Pheretima hilgendorfi, one of the most common anecic species in Japan, abounds in soils with applied rice-straw residues. The influences of the worm’s activity and/or those of rice-straw application on the soil microbial community structure were studied using a microcosm approach. Low Humic Andosol was incubated with or without earthworms in a jar for a month after the following treatments: 1) no treatment, 2) chopped rice straw top-dressed on the soil and 3) rice straw incorporated into the soil. The soil NO₃^–-N level increased with the earthworms’ presence even in the soil without rice straw. The soil NH₄⁺-N level was by far the highest in the soil treated with worms and no rice straw. Amounts of total and bacterial PLFAs increased due to the earthworms’ presence when rice straw was incorporated. The proportion of unsaturated fatty acids increased in the earthworm treatment while the saturated fatty acids decreased, suggesting an increase in the Gram-negative bacterial proportion. The results of principal component analysis indicated that the rice straw and the earthworms affected the soil microbial community structure independently. Any direct influence of the PLFAs contained in the worms’ bodies and in the rice straw on the soil’s PLFA profile was thought to be small.     

Assessing the impact of Diplocardia ornata on physical and chemical properties of compacted forest soil in microcosms

 The influence of compaction on Diplocardia ornata (Smith) burrowing and casting activities, soil aggregation, and nutrient changes in a forest soil were investigated using pot microcosms. Treatments included two levels each of compaction, organic matter, and earthworms. Both burrowing and casting activities were more abundant in uncompacted soil than in compacted soil. Bulk density decreased in microcosms of compacted soil containing D. ornata from 1.76 g cm^–3 to 1.49 g cm^–3 over the study period. The overall percent of aggregates in the same size classes in compacted soil was less than the percent of aggregates in uncompacted soil. The mean percent of aggregates in earthworm casts for size classes 0.25–1.00 mm was higher for compacted soil than for uncompacted soil. The reverse was true for aggregates in class sizes 2.00–4.00 mm. Soil compaction also affected soil microbial biomass carbon and soil inorganic N concentrations. These results indicate that the burrowing and casting activities of earthworms in compacted forest soils, as in soils of agricultural and pastured lands, can help ameliorate disturbed soils by improving aggregation, reducing bulk density, and increasing nutrient availability. Received: 1 September 1999     

The fate of catechol in soil as affected by earthworms and clay

The effect of endogeic earthworms (Octolasion tyrtaeum) and the availability of clay (Montmorillonite) on the mobilization and stabilization of uniformly ¹⁴C-labelled catechol mixed into arable and forest soil was investigated in a short- and a long-term microcosm experiment. By using arable and forest soil the effect of earthworms and clay in soils differing in the saturation of the mineral matrix with organic matter was investigated. In the short-term experiment microcosms were destructively sampled when the soil had been transformed into casts. In the long-term experiment earthworm casts produced during 7 days and non-processed soil were incubated for three further months. Production of CO₂ and ¹⁴CO₂ were measured at regular intervals. Accumulation of ¹⁴C in humic fractions (DOM, fulvic acids, humic acids and humin) of the casts and the non-processed soil and incorporation of ¹⁴C into earthworm tissue were determined.Incorporation of ¹⁴C into earthworm tissue was low, with 0.1 and 0.44% recovered in the short- and long-term experiment, respectively, suggesting that endogeic earthworms preferentially assimilate non-phenolic soil carbon. Cumulative production of CO₂-C was significantly increased in casts produced from the arable soil, but lower in casts produced from the forest soil; generally, the production of CO₂-C was higher in forest than in arable soil. Both soils differed in the pattern of ¹⁴CO₂-C production; initially it was higher in the forest soil than in the arable soil, whereas later the opposite was true. Octolasion tyrtaeum did not affect ¹⁴CO₂-C production in the forest soil, but increased it in the arable soil early in the experiment; clay counteracted this effect. Clay and O. tyrtaeum did not affect integration of ¹⁴C into humic fractions of the forest soil. In contrast, in the arable soil O. tyrtaeum increased the amount of ¹⁴C in the labile fractions, whereas clay increased it in the humin fraction.The results indicate that endogeic earthworms increase microbial activity and thus mineralization of phenolic compounds, whereas clay decreases it presumably by binding phenolic compounds to clay particles when passing through the earthworm gut. Endogeic earthworms and clay are only of minor importance for the fate of catechol in soils with high organic matter, clay and microbial biomass concentrations, but in contrast affect the fate of phenolic compounds in low clay soils.     

The effect of earthworms (Lumbricidae) on nitrogenase activity in soil

Summary Nitrogenase activity associated with earthworms, their faeces and activity in soil was measured by the acetylene reduction technique. A clear increase in nitrogenase activity was found in field-deposited casts of Aporrectodea caliginosa in comparison with surrounding soil, although potential nitrogenase activity was significantly higher in soil than in casts. Nitrogenase activity associated directly with earthworms (Lumbricus rubellus ) was detected, indicating the presence of active N₂-fixing bacteria on the body surface and/or in the gut. Laboratory experiments showed that nitrogenase activity in the casts of L. rubellus was higher than in unmodified soil, and that nitrogenase activity in soil was significantly increased by the burrowing and feeding activity of these worms. This paper discusses the possible causes of these earthworm effects on soil nitrogenase activity and some methodological problems of determining the nitrogenase activity.     

Effects of earthworms on nitrogen mineralization

The influence of earthworms (Lumbricus terrestris and Aporrectodea tuberculata) on the rate of net N mineralization was studied, both in soil columns with intact soil structure (partly influenced by past earthworm activity) and in columns with sieved soil. Soil columns were collected from a well drained silt loam soil, and before the experiment all earthworms present were removed. Next, either new earthworms (at the rate of five earthworms per 1200 cm³, which was only slightly higher than field numbers and biomass) were added or they were left out. At five points in time, the columns were analyzed for NH ₄ ⁺ , NO ₃ ^– , and microbial biomass in separate samples from the upper and lower layers of the columns. N mineralization was estimated from these measurements. The total C and N content and the microbial biomass in the upper 5 cm of the intact soil columns was higher than in the lower layer. In the homogenized columns, the C and N content and the microbial biomass were equally divided over both layers. In all columns, the concentration of NH ₄ ⁺ was small at the start of the experiment and decreased over time. No earthworm effects on extractable NH ₄ ⁺ were observed. However, when earthworms were present, the concentration of NO ₃ ^– increased in both intact and homogenized cores. The microbial biomass content did not change significantly with time in any of the treatments. In both intact and homogenized soil, N mineralization increased when earthworms were present. Without earthworms, both type of cores mineralized comparable amounts of N, which indicates that mainly direct and indirect biological effects are responsible for the increase in mineralization in the presence of earthworms. The results of this study indicate that earthworm activity can result in considerable amounts of N being mineralized, up to 90 kg N ha^–1 year^–1, at the density used in this experiment.     

不同秸秆施用方式下接种蚯蚓对土壤团聚体及其中碳分布的影响

通过室内培养试验,研究在施用有机物条件下接种蚯蚓对土壤团聚体的分布、团聚体的水稳性以及不同粒径的水稳性团聚体中有机碳含量的影响.干筛结果表明,不同秸秆施用方式下,蚯蚓接种能显著促进各个处理中＞2 mm团聚体含量的增加,且在秸秆混施的处理中表现得尤为明显,团聚体含量增加了2.95倍;湿筛结果表明,蚯蚓在不施和混施秸秆的处理中能显著降低土壤黏砂粒含量,即增加土壤中水稳性团聚体的含量,但是在表施秸秆的处理中显著降低了0.25～ 0.053 mm粒级团聚体含量,使之分散为黏砂粒.蚯蚓和秸秆对土壤团聚体分布和水稳性的影响都达到显著水平.蚯蚓对水稳性微团聚体的影响极显著,而秸秆的作用更多地表现在水稳性大团聚体上.在秸秆表施和秸秆混施条件下,接种蚯蚓均显著促进了微团聚体碳含量的增加,分别为相应对照的2.1和1.2倍.蚯蚓作用能显著降低黏砂粒有机碳在全碳中含量,增加团聚体有机碳含量,主要是由于蚯蚓的作用能促进黏砂粒黏结为团聚体.     

Novel method for the study of the population dynamics of a genetically modified microorganism in the gut of the earthworm Lumbricus terrestris

Summary A laboratory microcosm study was used to investigate the survival and population dynamics of genetically modified microorganisms (GMM) in the gut of Lumbricus terrestris. Three methods of axenic earthworm production were investigated. An antibiotic mixture of streptomycin and cycloheximide was introduced either passively, mixed with sterile soil or cellulose, or actively, by teflon catheter. Worms treated by all methods lost weight but this was least for the catheter method which was also the only method to produce axenic earthworms. Axenic earthworms were used to determine the effect of competition with indigenous gut bacteria on ingested GMM. The GMM used was Pseudomonas fluorescens, strain 10586/FAC510, with chromosomally inserted Lux genes for bioluminescence, and chromosomal resistance to rifampicin. The bacteria were grown up to the mid-exponential phase before inoculation into earthworms. Bacteria in faecal material were enumerated by dilution plate counting using selective agar. The GMM were re-isolated from the casts of both antibiotic-treated and untreated earthworms. Lower concentrations of GMM and higher concentrations of indigenous bacteria in the casts of untreated compared to antibiotic-treated earthworms suggested that competition is a fundamental control on population dynamics of the introduced bacterial inocula ingested by earthworms.The catheter method, developed in this study, is proposed as a technique to contribute to the risk assessment of environmental release of GMM.     

Impacts of earthworms and arbuscular mycorrhizal fungi (Glomus intraradices) on plant performance are not interrelated

Earthworms and arbuscular mycorrhizal fungi (AMF) might interactively impact plant productivity; however, previous studies reported inconsistent results. We set up a three-factorial greenhouse experiment to study the effects of earthworms (Aporrectodea caliginosa Savigny and Lumbricus terrestris L.) and AMF (Glomus intraradices N.C. Schenck & G.S. Sm.) on the performance (productivity and shoot nutrient content) of plant species (Lolium perenne L., Trifolium pratense L. and Plantago lanceolata L.) belonging to the three functional groups grasses, legumes and herbs, respectively. Further, we investigated earthworm performance and plant root mycorrhization as affected by the treatments. Our results accentuate the importance of root derived resources for earthworm performance since earthworm weight (A. caliginosa and L. terrestris) and survival (L. terrestris) were significantly lower in microcosms containing P. lanceolata than in those containing T. pratense. However, earthworm performance was not affected by AMF, and plant root mycorrhization was not modified by earthworms. Although AMF effectively competed with T. pratense for soil N (as indicated by δ¹⁵N analysis), AMF enhanced the productivity of T. pratense considerably by improving P availability. Remarkably, we found no evidence for interactive effects of earthworms and AMF on the performance of the plant species studied. This suggests that interactions between earthworms and AMF likely are of minor importance.     

Effects of heavy metals on the litter consumption by the earthworm Lumbricus rubellus in field soils

Aim of this study was to determine effects of heavy metals on litter consumption by the earthworm Lumbricus rubellus in National Park the “Brabantsche Biesbosch”, the Netherlands. Adult L. rubellus were collected from 12 polluted and from one unpolluted field site. Earthworms collected at the unpolluted site were kept in their native soil and in soil from each of the 12 Biesbosch sites. Earthworms collected in the Biesbosch were kept in their native soils. Non-polluted poplar (Populus sp.) litter was offered as a food source and litter consumption and earthworm biomass were determined after 54 days. Cd, Cu and Zn concentrations were determined in soil, pore water and 0.01 M CaCl₂ extracts of the soil and in earthworms. In spite of low available metal concentrations in the polluted soils, Cd, Cu and Zn concentrations in L. rubellus were increased. The litter consumption rate per biomass was positively related to internal Cd and Zn concentrations of earthworms collected from the Biesbosch and kept in native soil. A possible explanation is an increased demand for energy, needed for the regulation and detoxification of heavy metals. Litter consumption per biomass of earthworms from the reference site and kept in the polluted Biesbosch soils, was not related to any of the determined soil characteristics and metal concentrations.     

Fluorine contamination of soils and earthworms (Lumbricus spp.) near a site of long-term industrial emission in southern Germany

Summary The F contamination of soils and Lumbricus spp. around a site of long-term industrial emission in southern Germany was examined. Among total, water extractable, and HCl-soluble fractions, the latter most appropriately characterized anthropogenic F accumulation. Based on the HCl-soluble fractions from 88 sampling sites, a contamination map consisting of three zones was established. F accumulation in the calcareous soils of the area was restricted to the top 40–50 cm and can be explained by precipitation as CaF₂. Earthworms (Lumbricus spp.) collected from the different zones reflected the F contamination well in the significant correlations found between total F in earthworms with and without gut and the corresponding soils. The bioaccumulation of F in earthworms is obvious, and may become hazardous for the earthworms themselves and for other animals feeding on contaminated soil and/or its fauna. A significantly higher F value was recorded in the linings of earthworm tubes than in the corresponding soil. F translocation by earthworm burrowing may be a mechanism of subsoil contamination.