The fate of condensed tannins during litter consumption by soil animals

Condensed tannins (CT) can strongly affect litter decomposition, but their fate during the decomposition process, in particular as influenced by detritivore consumption, is not well understood. We tested the hypothesis that litter CT are reduced by the gut passage of two functionally distinct detritivores of Mediterranean forests, the millipede Glomeris marginata, and the land snail Pomatias elegans, as a fixed proportion of initial litter CT, but more so in Pomatias since snails are known to have a more efficient enzymatic capacity. Contrary to our hypothesis, both detritivore species reduced litter CT to near zero in their faecal pellets irrespective of the wide range in initial leaf litter CT concentrations of 9-188 mg g⁻¹ d m among three Mediterranean tree species (Pistacia terebinthus, Quercus ilex, Alnus glutinosa) and different decomposition stages of their litter. The almost complete disappearance of CT even from some litter types highly concentrated in CT, due to either degradation by gut microorganism or complexation of CT into insoluble high molecular weight structures, suggests a high “de-tanning” efficiency across functionally distinct detritivore species. The transformation of CT-rich litter into virtually CT-free faecal pellets by detritivores might be highly relevant for the subsequent decomposition process in ecosystems with a high macrofauna abundance and CT-rich plant species such as Mediterranean forests.     

Effects of heavy metals on the structure and functioning of detritivore communities in a contaminated floodplain area

The aim of this study was to determine the effects of heavy metal pollution on the structure and functioning of detritivore soil communities that consist of isopods, millipedes and earthworms, in 15 heavily polluted flood plain soils, located in the delta area of the rivers Rhine and Meuse, in the Netherlands. The 15 study sites represent a gradient in Zn, Cu and Cd concentrations. The structural attributes of the detritivore community, which were assessed, were the species richness and densities in the field sites. The functioning of the detritivore community was studied by determining organic matter decomposition using litter bags and feeding activity with the bait-lamina method. Concentrations of Cd, Cu and Zn were measured in soil, pore water and 0.01 M CaCl₂ extracts of the soil, in adult earthworms and plant leaves. Results show that metal pollution is not a dominating factor determining the species richness and densities of the selected detritivore groups, although the biomass of the earthworm Lumbricus rubellus was positively and significantly correlated to Zn concentrations in pore water and 0.01 M CaCl₂ extracts. Litter decomposition was significantly and positively correlated to detritivore biomass and 0.01 M CaCl₂ extractable Cd concentrations in soil and negatively to pH-CaCl₂, although the range of pH values was very small. It can be concluded that in spite of high metal levels in the soil, bioavailable concentrations are too low to result in clear negative effects on the structure and functioning of detritivores in the Biesbosch, the Netherlands.     

Effect of passage through the intestinal tract of detritivore earthworms (Lumbricus spp.) on the number of selected Gram-negative and total bacteria

Laboratory experiments were carried out to investigate the fate of bacteria during and after passage through the intestinal tract of detritivore earthworms. Earthworms (Lumbricus spp.) were fed with cattle dung inoculated 7 days previously with one of five different Gram-negative bacteria. Bacterial concentrations were determined 2 days later in dung and soil, and in gut material from different parts of the earthworm intestinal tract. A high percentage (28–82%) of the total bacteria (epifluorescence direct counts) in the earthworm gut content was culturable. The concentration of total heterotrophic aerobic bacteria did not vary significantly among the five different bacterial additions and the non-inoculated control. In earthworm casts the number of total heterotrophs per gram dry matter (2.1×10⁹) was higher than in soil (1.7×10⁸), but lower than in the dung (1.5×10¹⁰). The test-bacteria, however, showed different survival patterns along the earthworm intestinal tract. The concentrations of Escherichia coli BJ 18 and Pseudomonas putida MM 1 and MM 11 in earthworm casts were lower than in the ingested dung, while concentrations of Enterobacter cloacae A 107 and Aeromonas hydrophila DMU 115 in dung and casts were similar. Ent. cloacae, and to aminor extent E. coli, were reduced in numbers by several orders of magnitude in the pharynx and/or crop. In the hind gut, however, the concentration of Ent. cloacae had increased to the same level as in the ingested dung, while the concentration of E. coli remained low. Our observations indicate that the bacterial flora of ingested food materials changes qualitatively and quantitatively during gut transit.     

Intermediate tidal stress promotes the detritivore-mediated decomposition of Spartina litter

Saltmarshes, functionally important habitats in the marine–terrestrial ecotone that are regularly affected by tidal inundation, are mainly detritus-based in terms of fluxes of nutrients and energy. With respect to the mediating influence of saltmarsh detritivores on microbial colonisation of detritus and on decomposition processes, we tested whether the “intermediate disturbance hypothesis” (IDH) is also applicable to the effects of stress in this stressful environment. Decomposition experiments with litter of the cordgrass, Spartina anglica, and with terrestrial [Porcellio scaber (Isopoda)] and marine/semi-terrestrial [Orchestia gammarellus (Amphipoda)] detritivores as well as animal-free controls were carried out in an artificial saltmarsh system. Different daily flooding regimes served as experimental levels of stress. Both litter mass loss and microbial respiration were mostly higher under aquatic than under terrestrial conditions, no matter whether detritivores were present or not. Considering the intertidal zone, low to intermediate daily inundation rates resulted in increased microbial respiration and an increased influence of detritivores on litter mass loss in early stages of cordgrass decomposition with high rates of detritus mass loss, and intermediate tidal stress led to higher microbial cell counts throughout the entire experiment. Summarised over 3 months, regression analyses suggested that microbial activity and detritus mass loss show a trend towards highest values at low inundation rates and under permanent inundation, although microbial density was higher under longer daily inundation. Access to detritus by detritivores enhanced both litter mass loss and microbial respiration, especially in later decomposition stages, whereas microbial density was reduced by detritivores. In conclusion, we predict that overall the decomposition of cordgrass detritus in saltmarshes is promoted in the intermediate to high area of the intertidal zone with daily inundation of ca. 4–10 hours where both marine and terrestrial detritivores have access to promote decomposition processes through feeding and mediating microbial activity.     

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.     

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 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 microarthropod abundance and climatic factors on weight loss and mineral nutrient contents of Eucalyptus leaf litter during decomposition

Summary Densities of the different taxa of microarthropods per gram of litter in litter bags varied widely from the rainy to the dry season. Collembola and Acarina constituted more than 86% of the total microarthropods, and occurred in significantly greater numbers in the coarse-mesh bags than the fine-mesh bags. There were no fauna in the litter of suspended bags. The average weight loss was greater in the coarse-mesh bags (50.4%) than in the fine-mesh bags (44.5%), and the suspended bags (7.4%). Similarly, the concentrations of N and Ca were greater in the litter of coarse-mesh bags compared to that of the fine-mesh bags. In contrast, the concentrations of P and K were comparatively lower in coarse-mesh bags. The mass loss of litter showed a negative linear correlation with the total Collembola and with litter temperature. The N concentrations in the litter in both the mesh bags showed negative correlations with the abundance of total Collembola, and with that of Lepidocyrtus sp. and Sminthuridae, and rainfall. The N concentration in the litter in the coarse-mesh bags was positively correlated with the total number of arthropods but, surprisingly, was negatively correlated with the total number of Acarina. The concentration of Ca showed negative correlations with rainfall and litter moisture only. The P concentration showed positive correlations with total Collembola, with Lepidocyrtus sp. and Sminthuridae in both the mesh bags, with rainfall in the fine-mesh bags, and with total microarthropods in the coarse-mesh bags.     

Growth of Lumbricus terrestris and Aporrectodea caliginosa in an acid forest soil, and their effects on enchytraeid populations and soil properties

A laboratory experiment was carried out to test the hypothesis that the earthworms Lumbricus terrestris and Aporrectodea caliginosa are able to maintain their populations and reproduce in the acid forest soil of a deciduous forest where no lumbricids were found in the field. The experiment was conducted in 45-l containers in which layers of mineral subsoil, humus and organic topsoil collected from the site were established. Both species survived and at least L. terrestris reproduced during the 60 weeks’ incubation. Burrows and middens of L. terrestris were recorded and quantities of litter were consumed. The presence of lumbricids increased the organic matter content of humus, reduced the acidity of the topsoil and humus layers, and suppressed the population of the enchytraeid Cognettia sphagnetorum. A dense population of Enchytraeus albidus was found in L. terrestris middens. It is concluded that edaphic factors do not explain the absence of earthworms, but isolation from cultural landscapes and lack of opportunity to colonize the site from the surroundings is the decisive factor.     

Nutrient release from decomposing leaf litter of temperate deciduous forest trees along a gradient of increasing tree species diversity

In the litter of six deciduous tree species (Fagus sylvatica, Tilia spp., Fraxinus excelsior, Carpinus betulus, Acer pseudoplatanus and Acer platanoides) and in stand-specific litter mixtures, we compared mass loss and nutrient release across diversity levels along a gradient of decreasing proportion of Fagus in stands with similar environmental and physical soil conditions. The litterbag studies ran over 22 months. The decomposition rate constants (k) of the temperate forest species ranged from k = 0.5 for Fagus to k = 1.5-2 for all other tree species. In Fagus, k was closely negatively correlated with the thickness of the litter layer and positively correlated with soil pH and isopod abundance. k was significantly higher in the mixed species stands (except for Carpinus and Fraxinus) and was positively correlated with earthworm abundance. Over the whole incubation time, nutrient amount and release rates of N, P, K, Ca and Mg as well as C-related ratios showed significant differences between tree species but no consistent differences among the diversity levels. Initial C-related nutrient ratios of the leaf litter and abundance of mesofauna and macrofauna were correlated with the length of time lag before nutrient release. We conclude that the mere number of tree species is not the main driver of nutrient release and decomposition processes, but that key groups of the decomposer fauna as well as the characteristic traits of the individual tree species are decisive.     

Effect of earthworms on decomposition processes in raw humus forest soil: A microcosm study

Summary The earthworms Lumbricus rubellus (Hoffmeister) and Dendrobaena octaedra (Savigny) were studied in the laboratory to determine their effects on decomposition and nutrient cycling in coniferous forest soil. CO₂ evolution was monitored, and pH, PO ₄ ^3– –P, NH ₄ ⁺ –N, NO ₃ ^– –N, total N, and total C in the leaching waters were measured. After three destructive samplings, numbers of animals, mass loss, pH, and KCl-extractable nutrients were analysed.The earthworms clearly enhanced the mass loss of the substrate, especially that of litter. L. rubellus stimulated microbial respiration by 15–18%, whereas D. octaedra stimulated it only slightly. The worms significantly raised the pH of the leaching waters and the humus; L. rubellus raised the value by 0.2–0.6 pH units and D. octaedra by 0.1–0.4 units. Both worms increased N mineralization. Although the biomass of both worms decreased during the experiment, the N released from decomposing tissues did not explain the increase in N leached in the presence of earthworms. The worms influenced the level of PO ₄ ^3– –P only slightly.     

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.     

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.     

Effects of litter quality and microarthropods on N dynamics and retention of exogenous 15N in decomposing litter

Summary Surface additions of (¹⁵NH₄)₂SO₄ were used to measure the immobilization and subsequent movement of exogenous N added to two litter types of contrasting quality (Cornus florida and Quercus prinus). Litterbaskets were used to measure the litter mass loss and N dynamics and to follow the movement of the ¹⁵N label through litter, F layer, and soil pools. Half of the litterbaskets of each species were treated with naphthalene to reduce microarthropod densities. The faster decomposing C. florida litter maintained a higher excess atom % ¹⁵N, and a greater relative concentration of the labeled input (g ¹⁵N g^–1) than did Q. prinus litter. In both litter types the excess atom % ¹⁵N, relative concentration (g ¹⁵N g^–1), and absolute amount of label recovered in the litter declined over time. This occurred during a period of net accumulation of total litter N, implying simultaneous release of the initial input and immobilization of N from other sources. The concentration of ¹⁵N in the soil increased over time, while the F layer apparently acted as an intermediary in the transfer of ¹⁵N from litter to soil. Naphthalene effectively reduced microarthropod numbers in all horizons of the litterbaskets and significantly reduced the decay rates of Q. prinus, but not C. florida litter. Naphthalene did not appear to affect total N dynamics in the litter. However, with all horizons taken together, the naphthalene-treated litterbaskets retained more total ¹⁵N than the control litterbaskets. Naphthalene also changed the vertical distribution of ¹⁵N within litterbaskets, so that the litter retained less of the ¹⁵N-labeled input and the F layer and soil horizons retained more of the labeled input than in control litterbaskets. Our major conclusions are: (1) the N pool of decomposing litter is dynamic, with simultaneous N release and immobilization activating N turnover even during the net accumulation phase; (2) litter quality is an important determinant of immobilization and retention of exogenous N inputs and, therefore, turnover of the litter N pool; and (3) microarthropod activity can significantly affect the incorporation and retention of exogenous N inputs in decomposing litter, although these changes are apparently not reflected in net N accumulation or release during the 1st year of decomposition. However, the naphthalene may have affected microbially mediated N dynamics and this possibility needs to be considered in interpreting the results.     

Enhanced root nodulation of subterranean clover (Trifolium subterraneum) byRhizobium leguminosarium biovartrifolii in the presence of the earthwormAporrectodea trapezoides (Lumbricidae)

In a greenhouse study, the effect of the earthwormAporrectodea trapezoides on root nodulation in seedlings of subterranean clover (Trifolium subterraneum) was examined in the presence and absence of addedRhizobium leguminosarium biovartrifolii (strain NA 30). WhenR. trifolii NA 30 was inoculated into dung and placed on the soil surface, the total number of root nodules was five times greater (P<0.001) in the presence of earthworms than without earthworms and the number of nodules on the primary root of the plants 2–8 cm below the soil surface was 4 to 6 times greater (P<0.001) in the presence of earthworms. The additional nodulation did not affect plant growth or foliar N. When NA30 was dispersed through the soil at the beginning of the experiment, the presence of earthworms did not influence the level of root nodulation. The presence of earthworms increased root dry weight by 20–30%, plant top weight by up to 125% (P<0.001), and foliar N by 5–25% (P<0.001). Surface-applied dung increased the dry weight of plant tops (2-to 3-fold,P<0.001) but did not affect the concentration of foliar N (P<0.005).     

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.     

Decomposer animals (Lumbricidae, Collembola) and organic matter distribution affect the performance of Lolium perenne (Poaceae) and Trifolium repens (Fabaceae)

Decomposer animals stimulate plant growth by indirect effects such as increasing nutrient availability or by modifying microbial communities in the rhizosphere. In grasslands, the spatial distribution of organic matter (OM) rich in nutrients depends on agricultural practice and the bioturbation activities of large detritivores, such as earthworms. We hypothesized that plants of different functional groups with contrasting nutrient uptake and resource allocation strategies differentially benefit from sites in soil with OM accumulation and the presence of decomposer animals. In a greenhouse experiment we investigated effects of spatial distribution of ¹⁵N-labelled grass litter, earthworms and collembola on a simple grassland community consisting of Lolium perenne (grass) and Trifolium repens (legume). Litter aggregates (compared to homogeneous litter distribution) increased total shoot biomass, root biomass and ¹⁵N uptake by the plants. Earthworms and collembola did not affect total N uptake of T. repens; however, the presence of both increased ¹⁵N uptake by T. repens and L. perenne. Earthworms increased shoot biomass of T. repens 1.11-fold and that of L. perenne 2.50 fold. Biomass of L. perenne was at a maximum in the presence of earthworms, collembola and with litter concentrated in a single aggregate. Shoot biomass of T. repens increased in the presence of collembola, with L. perenne generally responding opposingly. The results indicate that the composition of the decomposer community and the distribution of OM in soil affect plant competition and therefore plant community composition.     

Combined effect of bioaugmentation and bioturbation on atrazine degradation in soil

Earthworms, because they change soil physical and chemical properties, are efficient engineers that act on soil microbial community and activity. Thus they may drive pollutant biodegradation in soil such as atrazine mineralization. We hypothesized that earthworms modify the abundance of indigenous soil bacteria and the fate and activity of atrazine-degraders in the soil they engineer by bioturbation. Two bacterial strains were used as bioaugmentation agents: Pseudomonas sp. ADP and Chelatobacter heintzii, which have acquired the capacity to metabolize atrazine by carrying plasmidic atz A, B, C, D, E, F and atzA, B, C, trzD genes, respectively. We analyzed the interactions between earthworms (Lumbricus terrestris) and the indigenous and atrazine-degrading (indigenous and inoculated) bacterial communities by quantifying the 16S rRNA and the atzA gene sequence copies numbers, respectively, in different earthworm microsites. The kinetics of atrazine mineralization were measured to link the bacterial community changes with the degradation function. Digestion by earthworms significantly impacted the number of indigenous bacteria and atrazine mineralization in bioaugmented soils. Regarding the fate of the two atrazine-degraders tested, Pseudomonas sp. strain ADP survived better within the 10 days of experiment than C. heintzii in the bulk soil but the surviving fraction of C. heintzii was still metabolically active and able to mineralize atrazine. A positive “burrow-lining” effect on the atzA sequence copies number was observed in soil whether bioaugmented with C. heintzii or not (i.e. native indigenous atzA) thereby indicating that burrow-linings form a specific ‘hot spot’ for atrazine-degraders. The present study is the first to report the role of earthworms in selecting native catabolic key-genes in soil (indigenous atzA). This catabolic gene selection through earthworm soil bioturbation could be important in sustaining the degradation (detoxification) function of soil.     

Earthworms disseminate a soil-borne plant pathogen,Fusarium oxysporum f. sp. raphani

Radish plants infested with a soil-borne plant pathogen, Fusarium oxysporum f. sp. raphani PEG-4, which is resistant to hygromycin B, were placed on the surface of a soil microcosm containing earthworms (Pheretima sp.). The earthworms ate the radish plants and scattered individual casts everywhere in the burrows. The fungal propagules were detected in the gut of the earthworms and in 26 out of 28 casts. These results suggested that the pathogen could survive passage through the alimentary canal of the earthworms and be disseminated anywhere earthworms wandered. The collected casts were incubated in a moist chamber at 28°C for 14 days, which resulted in a decline in the number of the propagules. The effects of the earthworms on the population dynamics of the pathogen were estimated. The earthworms seemed to cause a decline in total propagules of the pathogen in soil, although they expanded its distribution in soil.     

Decomposition of willow-leaf litter in a short-rotation forest in relation to fungal colonization and palatability for earthworms

Summary The influence of leaf litter from three Salix spp. on fungal growth and microbial decomposition was studied using 1-mm-mesh litter-bags, and the effect on additional soil macrofaunal activity was studied by measuring litter disappearance from 4-mm-mesh bags and under 4-mm-mesh nets. Mineral macro-elements, water-and ethanol-extractable substances, lignin, and protein-precipitating substances (astringency) in the litter were determined, taking contaminating of the litter with soil particles into account. As expected, the litter disappeared more quickly from the large-mesh bags than from the small-mesh bags, which was attributed to earthworm activity. During the 1st year, the rate of leaf disappearance from both types of bags and under the nets was much higher for S. daphnoides than for S. viminalis and S. fragilis. The lower initial astringency, related to the tannin content, of the S. daphnoides litter might account for this difference. Tannin metabolites probably hampered both microbial decomposition and earthworm acceptability for some time also after the astringency was lost. Neither the content of macronutrients nor that of the other organic fractions studied can be assumed to have had any effect on weight losses due to microbial decomposition. Although, the S. daphnoides leaves initially contained the least amount of fungal mycelium (m g^-1 dry weight), the increase after contact with soil was most pronounced in this litter. The species composition of Fungi Imperfecti in the leaves of S. viminalis and S. daphnoides differed only for fresh litter, whereas the number of isolates was somewhat higher for S. daphnoides throughout the study. Similar seasonal variations in fungal composition occurred in both the S. viminalis and the S. daphnoides litter.