Population and behavioural level responses of arable soil earthworms to boardmill sludge application

The response of earthworms to soil application of boardmill waste sludge was quantified in field and laboratory experiments. The influence of one application of 6×10⁴ tonnes ha⁻¹ of unamended sludge was tested against no application on silty-clay arable soil. After 2 years, results in stubble-cultivated soil showed a 1.7 times lower density of earthworms where sludge was added, whereas there was no difference in average earthworm fresh weight. In direct-drilled soil, there was an indication of lower average numbers of Lumbricus terrestris middens when sludge was applied. Negative impacts in the field may have been due to indirect negative effects of the sludge application. In the laboratory, habitat choice tests were undertaken with two common species from the field site using sludge-based mulch and fertiliser products. Aporrectodea caliginosa showed no discernible preference between soil and soil mixed with unamended sludge (mulch), but favoured soil over soil mixed with sludge and chicken manure (fertiliser). Tests with L. terrestris showed a similar pattern. The preference of L. terrestris for the two products was compared with that for chopped barley straw by direct observation of foraging behaviour. Differences in foraging time of L. terrestris for different feeds were not significant, but mass of straw collected was significantly greater compared with either type of sludge treatment.     

Responses of co-occurring populations of Dendrobaena octaedra (Lumbricidae) and Cognettia sphagnetorum (Enchytraeidae) to soil pH, moisture and resource addition

Dendrobaena octaedra (Lumbricidae) and Cognettia sphagnetorum (Enchytraeidae) are the two most dominating soil invertebrates in terms of biomass in boreal coniferous forest soils. A microcosm experiment was set up in order to study the influence of pH, moisture and resource addition on D. octaedra and C. sphagnetorum when both species are simultaneously present. Two kinds of coniferous forest humus were used as substrate, pine stand humus (pH 4.2), and spruce stand humus (pH 4.6); in the third treatment the pine stand humus was adjusted with slaked lime (CaOH₂) to the same initial pH as the spruce stand humus. Each substrate was adjusted to water contents of 25%, 42.5% and 60% of WHC (referred to as ‘dry’, ‘moist’ and ‘wet’). In the second part of the experiment, spruce needle litter and birch leaf litter were separately added into the pine stand humus (‘moist’, unlimed) and compared with a control without litter. The microcosms were plastic jars with 75 g (d.m.) of humus, into which 4 specimens of D. octaedra and 70 specimens of C. sphagnetorum were added. D. octaedra showed the highest biomass and C. sphagnetorum the lowest biomass in the spruce stand humus with higher pH. Moisture did not affect earthworms, while C. sphagnetorum thrived best at the highest moisture. Addition of both kinds of litter increased the numbers and biomass of D. octaedra, while on C. sphagnetorum resource addition had little effect. The results help to explain the abundance of these two species in coniferous forests differing in soil acidity, moisture and fertility.     

Microbial communities of Lumbricus terrestris L. middens: structure, activity, and changes through time in relation to earthworm presence

Background, aim, and scope  Earthworms make a major contribution to decomposition in ecosystems where they are present, mainly acting in the drilosphere, that is, galleries, burrows, casts, and middens. Earthworm middens are hot-spots of microbial activity and nutrient dynamics and represent a suitable model for studying earthworm-mediated influences on soil microbial communities by alteration of the patch structure of the microbial environment. We studied the structure and activity of the microbial communities in the soil system formed by middens of Lumbricus terrestris and the soil below and surrounding them and the role of earthworms in maintaining these structures through time. Material and methods  We set up an experiment in which middens were either left (control) or removed from their original place (translocated) and left in a nearby area free of earthworm activity for 2 months. After 1 and 2 months we sampled middens, soil below them, and surrounding soil. We analyzed the phospholipid fatty acid (PLFA) profiles and measured respiratory fluxes of CO₂ and CH₄. Results  Microbial communities of middens clearly differed from those of soil below and surrounding soil samples, showing higher bacterial and fungal PLFAs (p < 0.0001 and p < 0.01, respectively); furthermore, changes in microbial communities were stronger in control middens than in translocated middens. Moreover, gram positive and negative bacterial PLFAs were greater in translocated than control middens (p < 0.0001 and p < 0.001, respectively), as well as total organic carbon (p < 0.001). Microbial activity was higher in middens than in soil below and surrounding soil samples both for CO₂ (p < 0.0001) and CH₄ (p < 0.0001). Discussion  Soil bioturbation by the earthworm L. terrestris was strong in their middens, but there was not any effect on soil below and surrounding soil. Microbial communities of middens maintain their biomass and activity when earthworms were not present, whereas they decreased their biomass and increased their activity when earthworms were present. Conclusions  Earthworms strongly enhanced microbial activity measured as CO₂ production in middens, which indicates that there are hot spots for soil microbial dynamics and increasing habitat heterogeneity for soil microorganisms. Moreover, our data strongly support the fact that the impact of this earthworm species in this soil is restricted to their middens and increasing soil heterogeneity. Recommendations and perspectives  Our data indicate that it is not clear if earthworms enhance or depress microbial communities of middens since the microbial activity increased, but did not modify their biomass and this was not dependent on soil organic C content. These results indicate no competence for C pools between this anecic earthworm and microorganisms, which has been found for other earthworm species, mainly endogeics. Conversely, they suggest some type of facilitation due to the release of additional nutrient pools in middens when earthworms are present, through the digestion of middens' material or the addition of casts produced from other food sources.     

Genetic structure among earthworms (Lumbricus terrestris L.) from different sampling sites in western Germany based on random amplified polymorphic DNA

Earthworms are being used as bio-indicators to assess terrestrial pollution. However, it is often not known whether their populations possess a uniform genetic structure, which would allow comparison of residues or biological properties of earthworms from different sampling locations. In order to investigate this point, random amplified polymorphic DNA (RAPD) variation was surveyed in earthworms (Lumbricus terrestris) from five different sampling sites in Germany. Forty oligonucleotide RAPD primers (10 base pairs in length) were screened, three of which produced high polymorphic band patterns. A total of 61 DNA fragments were detected in 90 individuals of L. terrestris from five sampling sites with 49 (80.3%) RAPD markers being polymorphic. The genetic similarities within (band sharing rates between 0.756 and 0.795) and among the L. terrestris populations (0.635) were similar even at widely separate locations. Inter-population variation in the RAPD pattern for all five earthworm populations accounted for 37.9% of the total variation, while intra-population variation for three adjacent Saarland populations accounted for only 18.0% of the total variation. Principal component analysis (PCA) and the genetic distances of the populations confirm these results. Twenty-four percent of the genetic distance is caused by geographical isolation as shown by a test for isolation by distance. These results show that L. terrestris fulfils the genetic qualifications for a bio-indicator particularly at closely located sampling sites. However, the results also suggest that earthworm studies of widely separated locations should include genetic characterisation of the earthworm samples.     

Zum Einfluß von obstbaulichen Bodenpflegemaßnahmen auf Lumbriciden,insbesondere Lumbricus terrestris L .

Influence of orchard soil management on lumbricids, especially Lumbricus terrestris L. In a long-term soil management experiment (apple orchard treated with cultivation methods such as grass mulch, grass harvest, straw mulch, clean cultivation) 6 plots were selected in order to study the abundance and biomass of earthworm populations. The following results were obtained: In all treatments with the exception of straw mulch L. terrestris represented the majority (60.5% on average) of the lumbricid population. Their share in the biomass amounted to 93.3%. The earthworm biomass was the highest in grass mulch; a close correlation was observed between the litter production (grass and leaves) and the biomass of L. terrestris. Added farm yard-manure on plots with grass mulch had no additional positive effect on earthworms. Although a higher earthworm biomass was found under grass mulch in comparison with straw mulch, the species diversity was higher in the latter. Clean cultivation had a highly detrimental effect on all earthworms, especially on the horizontally burrowing species. The annual nitrogen turnover by L. terrestris in mulched orchards was estimated to approx. 50 kg N/ha.     

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.     

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.     

Reproduction and growth of three deep-burrowing earthworms (Lumbricidae) in laboratory culture in order to assess production for soil restoration

Laboratory experiments were conducted to examine the growth and reproduction of three deep-burrowing lumbricids, Aporrectodea longa, Lumbricus terrestris, and Octolasion cyaneum. The reproductive output was recorded as 18.8, 38.0, and 32.3 cocoons per worm per year for A. longa, L. terrestris, and O. cyaneum, respectively. For the same species, maturity was reached at a mean mass of 3.9, 5.0 and 2.4 g, within 3 months from the hatchling stage by L. terrestris and within 4 months by the other two species. The hatching success of cocoons at 15 and 20°C was within the range of 70–80% for each species, except A. longa at the higher temperature, where a viability of 47% was recorded. Twenty percent of viable O. cyaneum cocoons produced twin hatchlings, compared with only one percent for A. longa and L. terrestris. A combination of these results suggests that a complete life-cycle for each species could be achieved within 6 months (L. terrestris and A. longa) or 7–8 months (O. cyaneum). Each species has particular life-cycle strategies that would aid survival and colonisation, under field conditions, if inoculated into restored soils.     

巨尾桉工业原料林下植物多样性与土壤因子的关系

采用野外调查与室内分析相结合的方法,研究了巨尾桉工业原料林下植物多样性、土壤因子的变化特征以及二者之间的关系。结果表明,物种丰富度指数是草本层最高,而Simpson指数,Shannon—Wiener指数和均匀度指数是灌木层最高。巨尾桉林地土壤水分含量的变化规律为:母质层淀积层腐殖质层,有机质、水解氮、有效磷和速效钾含量的变化规律为:腐殖质层淀积层母质层。腐殖质层、淀积层和母质层的土壤水分含量、有机质含量与灌木层或草本层丰富度指数、多样性指数和均匀度指数无显著相关性;腐殖质层速效钾含量与草本层Simpson指数极显著相关;淀积层速效钾含量与灌木层Simpson指数极显著相关;水解氮与灌木层及草本层以Simpson指数为基础的均匀度指数,以Shannon—Wiener指数为基础的均匀度指数极显著相关;母质层土壤各因子含量与灌木层或草本层多样性指数无显著相关。研究结果为巨尾桉林的可持续经营管理提供理论基础和科学依据。     

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.     

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.     

Spring dynamics of soil carbon, nitrogen, and microbial activity in earthworm middens in a no-till cornfield

Earthworm activity may be an important cause of spatial and temporal heterogeneity of soil properties in agroecosystems. Structures known as “earthworm middens,” formed at the soil surface by the feeding and casting activities of some earthworms, may contribute significantly to this heterogeneity. We compared the temporal dynamics of carbon (C), nitrogen (N), and microbial acitivity in Lumbricus terrestris middens and in surrounding non-midden (bulk) soil during the spring, when seasonal earthworm activity was high. We sampled soil from middens and bulk soil in a no-till cornfield on four dates during May and June 1995. Soil water content and the weight of coarse organic litter (>2mm) were consistently higher in middens than in bulk soil. Total C and N concentrations, C:N ratios, and microbial activity also were greatest in midden soil. Concentrations of ammonium-nitrogen and dissolved organic N were greater in middens than in bulk soil on most dates, suggesting accelerated decomposition and mineralization in middens. However, concentrations of nitrate were usually lower in middens, indicating reduced nitrification or increased leaching and denitrification losses from middens, relative to bulk soil. Fungal activity, as well as total microbial activity, was consistently greater in middens. The contribution of fungae to overall microbial activity differed significantly between middens and bulk soil only on one date when both soils were very dry; the contribution of fungae to microbial activity was lower in the middens on this date. We conclude that the midden-forming activity of L. terrestris can be a major determinant of spatial heterogeneity in some agricultural soils, and that this can potentially affect overall rates of soil processes such as organic matter decomposition, N mineralization, denitrification, and leaching. Received: 4 April 1997     

Effects of endogeic earthworms on soil processes and plant growth in coniferous forest soil

Summary The effects of the endogeic earthworm, Aporrectodea caliginosa tuberculata (Eisen) on decomposition processes in moist coniferous forest soil were studied in the laboratory. The pH preference of this species and its effects on microbial activity, N and P mineralization, and the growth of birch seedlings were determined in separate pot experiments. Homogenized humus from a spruce stand was shown to be too acid for A. c. tuberculata. After liming, the earthworms thrived in the humus and their biomass increased (at pH above 4.8). In later experiments in which the humus was limed, the earthworms positively influenced the biological activity in humus and also increased the rate of N mineralization. A. c. tuberculata increased the growth of birch seedlings, with increases observed in stems, leaves, and roots. Neither NH ₄ ⁺ -N fertilizer nor mechanical mixing with artificial worms affected seedling growth. No plant-growth-affecting compounds (e.g., hormone-like compounds) due to the earthworms were present in the humus. The shoot: root ratio in the birch seedlings was not affected by either the earthworms or the fertilizer. The experiments revealed the impact of earthworm activity on soil processes and plant growth.     

Effects of lumbricids and enchytraeids on nematodes in limed and unlimed coniferous mor humus

In a factorial laboratory experiment, specimens of Dendrobaena octaedra (Lumbricidae) and Cognettia sphagnetorum (Enchytraeidae) were added to microcosms with unlimed (pH 4.5) and limed (pH 5.5) coniferous mor humus containing bacteria, fungi, protozoans, and nematodes. Effects on the nematodes were assessed after an incubation period of 207 days at 15°C and a soil moisture content of 60% water-holding capacity. When D. octaedra was absent, nematodes were significantly more abundant in the limed humus than in the unlimed humus. The presence of D. octaedra markedly reduced the number of nematodes in the limed humus but not in the unlimed one, where D. octaedra lost weight and probably did not feed. Most nematodes (92–97%) were bacterial-feeders. The presence of D. octaedra did not decrease the number or biomass of bacteria, indicating that the reduction in nematode numbers was not the result of competition for bacteria between D. octaedra and the nematodes. The presence of C. sphagnetorum had no effect on the nematodes in either of the treatments. We suggest that the reason why D. octaedra, but not C. sphagnetorum, reduced nematode numbers is that the former was more likely to inadvertently ingest the nematodes because of its much greater size. The results provide a possible explanation for the observation that liming sometimes enhances nematode populations, when lumbricids do not respond to the treatment, and sometimes causes decreases, when lumbricids increase in number.     

Monitoring the effects of pesticide pellets to address farmers' concerns on soil fauna,specifically earthworms

Slugs are a major pest in arable agriculture that are typically controlled by using pesticide-impregnated pellets (molluscicides). We received multiple requests from farmers in 2018 for research into the impact of slug pellets on earthworms because they had been told that ferric-based pellets were ‘worm-friendly’. We measured soil fauna feeding activities within Lumbricus terrestris middens on a replicated field plot experiment comparing the recommended product rates of a metaldehyde (4%) based or ferric phosphate EDDS (3%) based slug pellets. There was a significant decrease in feeding activity in the presence of either type of slug pellet, particularly in the top 25 mm of the soil indicating movement away from the pesticide. This could be explained by the significantly lower abundance (numbers and biomass) of earthworms in the topsoil. Four months after ferric phosphate-based slug pellet application, there was no significant difference between the control and treated plots in earthworm abundance, indicating the effects were temporary. In the laboratory, individual Lumbricus terrestris earthworms consumed up to 4 slug pellets over 14 days and no significant differences in biomass were measured. A burrowing fitness assay indicated that earthworms exposed to ferric phosphate-based slug pellets burrowed to a greater depth. This suggests an avoidance behaviour response, which could also explain the field results. A consultation with farmers (n = 68) rated this information service as useful and indicated widespread interest in minimizing the effects of pesticides on non-target soil fauna.     

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.     

Effect of management practices and cropping systems on earthworm abundance and microbial activity in Sanborn Field: a 115-year-old agricultural field

Sanborn Field, an agricultural demonstration field at the University of Missouri-Columbia, has over 100 years of cropping histories and management practices implemented at this site. We examined the effect of these cropping systems and management practices on earthworm populations and microbial activity. A field experiment was conducted to characterize and quantify earthworms and to determine the microbial activity in the same selected plots. We sampled 14 plots at the site, six of which were the original plots established in 1888 with the same cropping histories and management, and eight of which had been modified over the years to accommodate changing farmer needs and scientific questions. Earthworms and soil samples were collected in the spring and fall of 1999. Aporrectodea trapezoides, Aporrectodea caliginosa, and Lumbricus terrestris were the dominant species found in this field. Lumbricus terrestris was commonly associated with the no-tillage plot at Sanborn Field. Due partly to soil moisture conditions, most of the species were found and identified during the spring. Microbial activity, as expected, was greatest during the spring. Manure, no-tillage and crop rotations that include legumes had the greatest earthworm abundance and microbial activity. In general, where no fertility treatments were added to soil or where food sources were lacking, earthworm abundance decreased significantly. The abundance of earthworms found in this field reflected the cropping history and management practices used recently and within the last 100 years. In most cases, microbial activity showed similar trends as earthworm density. However, some dynamic microbial transformations, like nitrification, do not always follow the same trend as potential soil quality biotic indicators. Earthworms, along with microbial activity, may serve as useful soil quality indicators in sustained and short-term field trials.     

Contribution of anecic earthworms to biopore formation during cultivation of perennial ley crops

Large sized biopores (diameter >2 mm) in the subsoil can be created by tap roots, which leave voids after their decay, or by the burrowing activity of anecic earthworms which may benefit from the temporary lack in tillage in perennial cropping systems. However, the interactions between root growth and earthworm activity in the process of biopore formation during perennial ley cropping are not well understood. The aim of this field study was to quantify the development of the abundance of the anecic earthworm Lumbricus terrestris and the biopore density during the cultivation of lucerne (Medicago sativa L.), chicory (Cichorium intybus L.) and tall fescue (Festuca arundinacea Schreb.) grown for either one, two or three years. An increased abundance of L. terrestris was already recorded after two years of continuous ley when compared with one year cultivation. The ley crop species had only minor influence on the abundance of L. terrestris. Biopore densities of both diameter classes under study (2–5 mm and >5 mm) were not significantly affected by the duration of ley cropping. In contrast, biopore densities were influenced by ley crop species. More biopores of diameter class 2–5 mm were recorded after chicory than after fescue. Lucerne cropping resulted in intermediate biopore density. Additionally, in an incubation experiment under field conditions, we quantified whether L. terrestris preferentially created new biopores or colonized abandoned, previously existing ones. After three weeks of incubation, one third of the adult individuals incubated in the experimental area created new biopores at 0.4 m soil depth. A similar percentage of individuals colonized previously existing biopores, partially widening the lumen of smaller sized biopores. The remaining individuals remained in the topsoil. Sub-adult individuals rarely formed new pores. Half of the introduced sub-adults remained in the topsoil. We conclude that in crop rotations new biopores can be generated during perennial ley cropping by taproot systems of ley crops, but that a two to three- year period without tillage is not sufficient for populations of anecic earthworms to make a marked contribution to biopore density in the subsoil. The relevance of anecic earthworms for altering physical and chemical properties of biopores during ley cropping still needs further investigation.     

The effect of feeding behavior on Hg accumulation in the ecophysiologically different earthworms Lumbricus terrestris and Octolaseon cyaneum: A microcosm experiment

 A soil microcosm experiment was performed to assess the uptake of Hg from various Hg-spiked food sources (soil, leaf litter and root litter of Trifolium alexandrinum) by two earthworm species, Lumbricus terrestris (anecic) and Octolaseon cyaneum (endogeic). Treatments were applied in which one of the three food sources was Hg spiked and the other two were not. Additional treatments in which all or none of the food sources were Hg spiked were used as controls. Uptake of Hg from soil into tissues of both earthworm species was significantly higher than uptake of Hg from leaf litter or root litter, indicating that soil may be the most important pool for the uptake of Hg into earthworms. In addition, the anecic L. terrestris significantly accumulated Hg from all Hg-spiked food sources (leaf litter, root litter and soil), whereas the endogeic O. cyaneum took up Hg mainly from soil particles. Interestingly, there was no further increase in Hg in L. terrestris when all food sources were Hg spiked compared to the single Hg-spiked sources. This may be attributed to the relatively high Hg content in the soil, which may have influenced the feeding behavior of the earthworms, although their biomass did not significantly decline. We suggest that, in addition to the physiological differences, feeding behavior may also play a role in the contrasting uptake of Hg by the two earthworm species.     

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.