Changes in respiration rate and some physicochemical properties of soil during gut transit through Allolobophora molleri (Lumbricidae,Oligochaeta)

Summary Some physicochemical and biological measurements were carried out on the gut content and casts of Allolobophora molleri, an earthworm which lives in humid soils of Northern Spain. In the anterior part of the gut, water (22% of moist weight of soil) and soluble organic matter (27.4%) had been added to the ingested soil and pH had increased from 5.75 to 7.0. The amount of water-soluble compounds decreased sharply in the middle and posterior parts of the gut and were hardly detectable in control soil or casts. The average O₂ consumption, measured at 28° and 21°C, indicated respectively 2.75-to 12-fold increases in microbial respiration in the gut content compared to the non-ingested soil. These results extend the hypothesis of a mutualistic digestion in earthworms previously proposed for tropical endogeic species.     

Mixing of different mineral soil layers by endogeic earthworms affects carbon and nitrogen mineralization

The effect of the endogeic earthworm species Octolasion tyrtaeum (Savigny) on decomposition of uniformly ¹⁴C-labelled lignin (lignocellulose) was studied in microcosms with upper mineral soil (Ah-horizon) from two forests on limestone, representing different stages of succession, a beech- and an ash-tree-dominated forest. Microcosms with and without lower mineral soil (Bw-horizon) were set-up; one O. tyrtaeum was added to half of them. It was hypothesised that endogeic earthworms stabilise lignin and the organic matter of the upper mineral soil by mixing with lower mineral soil of low C content. Cumulative C mineralization was increased by earthworms and by the addition of lower mineral soil. Effects of the lower mineral soil were more pronounced in the beech than in the ash forest. Cumulative mineralization of lignin was strongly increased by earthworms, but only in the beech soil (+24.6%). Earthworms predominantly colonized the upper mineral soil; mixing of the upper and lower mineral soils was low. The presence of lower mineral soil did not reduce the rates of decomposition of organic matter and lignin; however, the earthworm-mediated increase in mineralization was less pronounced in treatments with (+8.6%) than in those without (+14.1%) lower mineral soil. These results indicate that the mixing of organic matter with C-unsaturated lower mineral soil by endogeic earthworms reduced microbial decomposition of organic matter in earthworm casts.     

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.     

Decomposition of lignin in soil microcompartments: A methodical study with three different C-14-labelled lignin substrates

Summary C mineralization of three different C-14-labelled lignin substrates (Klason-lignin, dioxane-lignin, lignocellulose) was investigated in four microcompartments, consisting of white-rotted beechwood (Fagus sylvatica L)., brown-rotted beechwood, earthworm [Octolasion lacteum (Örley)] faecal particles mixed with white-rotted beechwood, and earthworm feacal particles mixed with brown-rotted beechwood, incubated at 10°C for 193 days. Conversion of the labelled substrates to ¹⁴CO₂ was low in both white-rotted and brown-rotted wood without faecal particles. Overall C mineralization followed the order Klason-lignin > dioxane-lignin > lignocellulose, indicating that there were different amounts of labelled contaminants in the lignin substrates. Lignin degradation was more pronounced in earthworm faeces mixed with wood materials, and overall C mineralization ranged between 4.4% and 6.3% of the inital C content. C mineralization of the lignin substrates increased considerably in faecal particles after about 90 days, presumably due to nutrient immobilization and microbial succession. The usefulness of the three lignin substrates in monitoring the time-course and extent of lignin degradation in soil microcompartments is discussed.     

Microbial activity and nutrient dynamics in earthworm casts (Lumbricidae)

Summary Microbial respiration, microbial biomass and nutrient requirements of the microflora (C, N, P) were studied in the food substrate (soil taken from the upper 3 cm of the mineral soil of a beech wood on limestone), the burrow walls and the casts of the earthworm Aporrectodea caliginosa (Savigny). The passage of the soil through the gut caused an increase in soil microbial respiration of about 90% over a 4-week period. Microbial biomass was increased only in freshly deposited casts and decreased in aging faeces to a level about 10% lower than in soil. Microbial respiration of the burrow walls was only increased over a shorter period (about 2 weeks). The microflora of the soil and the burrow walls was limited by P, whereas in earthworm casts, microbial growth was limited by the amount of available C. In aging faeces the P requirement of the microflora increased and approached that of the soil. Immobilization of phosphate in earthworm casts is probably caused by mainly abiotic processes. C mineralization by soil microflora fertilized with glucose and P was limited by N, except in freshly deposited casts. Ammonium, not nitrate, was responsible for this process. N dynamics in earthworm casts are discussed.     

Effect of Hormogaster elisae (Oligochaeta; Hormogastridae) on the stability of soil aggregates

The production and stability of soil aggregates produced by laboratory cultures of the endogeic earthworm Hormogaster elisae was studied using three different techniques: the determination of the soil mean weight diameter (MWD), the aggregate tensile strength, and by the Blanchart method, which involves three different tests. The MWD index of soils was higher in microcosms containing H. elisae. Tensile strength was significantly higher in earthworm casts than in naturally formed aggregates. The Blanchart method also showed aggregates produced by earthworms to be more stable. The results of all three methods concur in that aggregates produced by H. elisae are larger and more stable than those produced in control microcosms without earthworms.     

Nitrogen mineralization and reorganization in casts of the geophagous tropical earthworm Pontoscolex corethrurus (Glossoscolecidae)

Summary Mineral N concentrations ranged from 133.1 to 167.8 g g^-1 dry soil in fresh casts of the endogeic earthworm Pontoscolex corethrurus fed on an Amazonian Ultisol; this was approximately five times the concentration in non-ingested soil. Most of this N was in the form of NH _inf4 ^sup+ . N also accumulated in microbial biomass, which increased from a control value of 10.5–11.3 to 67.5–74.1 g g^-1 in fresh casts. During a 16-day incubation, part of the NH _inf4 ^sup+ -N was nitrified and/or transferred to the microbial biomass. Total labile N (i.e., mineral+biomas N) decreased sharply at first (ca. 50% in the first 12 h), and then more slowly. The exact fate of this N (microbial metabolites, denitrification, or volatilization) is not known. After 16 days, the overall N content of the casts was still 28% higher than that of the control soil. Incubation of the soil before ingestion by the earthworms significantly increased the production of NH _inf4 ^sup+ in casts. We calculate that in a humid tropical pasture, 50–100 kg mineral N may be produced annually in earthworm casts. Part of this N may be conserved in the compact structure of the cast where the cast is not in close contact with plant roots.     

Abundance and trophic structure of macro-decomposers on alpine pastureland (Central Alps, Tyrol): effects of abandonment of pasturing

On four differently managed and abandoned alpine meadows and pastures densities, biomasses and ¹⁵N signatures of the macrofauna were assessed to evaluate the structural and functional changes of the decomposer food webs. The composition of the macrofauna decomposer community changes remarkably after the abandonment of alpine meadows and pastures. Lumbricus rubellus functions as key primary decomposer on alpine meadows and pastures whereas on abandoned sites other primary decomposers including Dendrobaena octaedra, Cylindroiulus meinerti, C. fulviceps and diptera larvae become more important. Decomposer species, such as Enantiulus nanus, presumably function as both primary and secondary decomposers and endogeic earthworms, such as Octolasion lacteum and Aporrectodea rosea, uniformly function as secondary decomposers. Abandonment of pasturing causes a shift in the composition of the macrofauna and the newly established fauna is unable to process and translocate the litter materials produced by the plants of the abondoned sites.     

Earthworm effects on the use of C sources by microorganisms: Non-linear response to temperature alteration

A microcosm was used to study the effect of the endogeic earthworm Aporrectodea caliginosa (Savigny) on the use of C by microorganisms in a calcareous beech forest soil and its dependence on temperature (5–25%C). Inclusion of ¹⁴C-labelled beech leaf litter made it possible to differentiate between C use by litter-colonizing microflora and by autochthonous soil microflora. The effect of temperature on the soil microbial biomass ¹²C was confined to a significant increase at 15 and 20°C. The size of the ¹⁴C-labelled microbial biomass, in contrast, was positively correlated with temperature. The ¹²C mineralization increased exponentially with temperature. The relationship between ¹⁴C mineralization and temperature, in contrast, followed a logistic curve. Significant main effects of A. caliginosa were confined to ¹²C mineralization, reflecting an increase in ¹²CO₂–C production in the earthworm treatments. The earthworm effects on ¹²CO₂–C production and on ¹⁴C incorporation of the microflora were not linear. The effect of A. caliginosa on ¹²CO₂–C production was most pronouned at intermediate temperatures. It is concluded that temperature alterations affect the microbial use of different C sources in different ways and that the temperature effects can be significantly modified by endogeic earthworms.     

The earthworm population of a winter cereal field and its effects on soil and nitrogen turnover

The earthworm population in a winter cereal field in Ireland was studied over a 3-year-period and its effects on soil and N turnover were assessed. The mean annual population density was 346–471 individuals m^-2 and the mean biomass was 56.9–61.2 g m^-2. Twelve species were recorded, the most abundant being Allolobophora chlorotica followed by Aporrectodea caliginosa, and 242 mg at 5°C to 713 mg at 10°C in the case of juvenile Lumbricus terrestris. Gut contents (dry mass of soil) comprised 6.7–15.5% of the A. caliginosa live mass, and 9.7–14.7% of the Lumbricus terrestris mass. Annual soil egestion by the field population was estimated as 18–22 kg m^-2. Tissue production ranged from 81.7 to 218.5 g m^-2, while N turnover resulting from mortality was calculated as 1.5–3.9 g m^-2 depending on the year and the method of calculation. Earthworms were estimated to contribute an additional 3.4–4.1 g mineral N to the soil through excretion, mucus production, and soil ingestion. Independent estimates of N output via mucus and excretion derived from ¹⁵N laboratory studies with Lumbricus terrestris were 2.9–3.6 g m^-2 year^-1.     

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.     

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.     

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 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.     

Inter- and intra-specific interactions of Lumbricus rubellus (Hoffmeister, 1843) and Octolasion lacteum (Örley, 1881) (Lumbricidae) and the implication for C cycling

Earthworms are important engineering species of many terrestrial ecosystems as they play a significant role in regulating C turnover. The effects of earthworms on moderating C decomposition processes differ across species and with interactions between species, which is not fully understood. We carried out an experiment to study the interactions of Lumbricus rubellus and Octolasion lacteum, and their effects on soil respiration. Laboratory mesocosms were set up using tulip poplar (Liriodendron tulipifera) leaf litter and varying densities of earthworms in single and combined species treatments. CO₂ efflux rate was used as an indicator of C decomposition rates, and measured with CO₂ sensors every five days over one month. L. rubellus induced higher leaf consumption rate and higher CO₂ efflux than O. lacteum; meanwhile O. lacteum grew more than L. rubellus. Both litter consumption rate and growth rate of earthworms decreased with increasing earthworm density. Soil CO₂ efflux increased with increasing earthworm density (from ∼1-2 μg CO₂ g⁻¹ hr⁻¹ with no earthworms to ∼ 4 μg CO₂ g⁻¹ hr⁻¹ with 8 earthworms). Combining the two species had a synergistic effect on leaf litter consumption, and neutralizing effects on soil respiration. The data suggest that the strength of intra- and inter-specific interactions among earthworm ecological groups varies at different absolute and relative densities, leading to altered leaf litter decomposition and C cycling.     

Short-term effects of earthworm activity and straw amendment on the microbial C and N turnover in a remoistened arable soil after summer drought

Short-term effects of actively burrowing Octolasion lacteum (Örl.) (Lumbricidae) on the microbial C and N turnover in an arable soil with a high clay content were studied in a microcosm experiment throughout a 16 day incubation. Treatments with or without amendment of winter wheat straw were compared under conditions of a moistening period after summer drought. The use of ¹⁴C labeled straw allowed for analyzing the microbial use of different C components. Microbial biomass C, biomass N and ergosterol were only slightly affected by rewetting and not by O. lacteum in both cases. Increased values of soil microbial biomass were determined in the straw treatments even after 24 h of incubation. This extra biomass corresponded to the initial microbial colonization of the added straw. O. lacteum significantly increased CO₂ production from soil organic matter and from the ¹⁴C-labeled straw. Higher release rates of ¹⁴C-CO₂ were recorded shortly after insertion of earthworms. This effect remained until the end of the experiment. O. lacteum enhanced N mineralization. Earthworms significantly increased both mineral N content of soil and N leaching in the treatments without straw addition. Moreover, earthworms slightly reduced N immobilization in the treatments with straw addition. The immediate increase in microbial activity suggests that perturbation of soil is more important than substrate consumption for the effect of earthworms on C and N turnover in moistening periods after drought.     

Effect of diet on cast production by the megascolecid earthworm Amynthas alexandri in laboratory culture

Feeding and casting activity of Amynthas alexandri fed on corn, wheat leaves, and mixed grasses were monitored in laboratory cultures. Casts were produced on the surface and sides of the containers. Food consumption varied from 36.5 to 69 mg g^–1 live worm day^–1. Cast production ranged from 3.95 to 5.9 mg g^–1 live worm day^–1. The C:N ratio in casts in laboratory cultures (11.17) and in field samples (8.84) was consistently lower than the corresponding ratio in the parent soil (13.19 and 10.54, respectively). This was probably due to mineralization of plant-derived organic material during passage through earthworms with consequent low C:N ratios.     

The influence of mineral and organic fertilisers on the growth of the endogeic earthworm Octolasion tyrtaeum (Savigny)

Endogeic earthworms play an important role in mobilisation and stabilisation of carbon and nitrogen in forest and arable soils. Soil organic matter is the major food resource for endogeic earthworms, but little is known about the size and origin of the organic matter pool on which the earthworms actually live. We measured changes in body mass of juvenile endogeic earthworms, Octolasion tyrtaeum (Savigny), in soils with different C and N contents resulting from different fertiliser treatments. The soil was taken from a long-term experiment (Statischer Düngungsversuch, Bad Lauchstädt, Germany). The treatments included (1) non-fertilised soil, (2) NPK fertilised soil, (3) farmyard manure fertilised soil and (4) NPK + farmyard manure fertilised soil. The soil was incubated in microcosms with and without one juvenile O. tyrtaeum for 80 days.Earthworm biomass decreased in non-fertilised soil by 48.6%, in NPK soil by 9.4%, but increased in farmyard manure soil by 19.7% and 42.8% (soil with additional NPK application). In farmyard manure treatments the biomass of bigger individuals decreased, but in smaller individuals it increased. In NPK fertilised soil without farmyard manure only small O. tyrtaeum increased in body mass, whereas in the non-fertilised soil all individuals decreased in body mass. Generally, soil respiration correlated positively with soil carbon content. Earthworms significantly increased soil respiration and nitrogen leaching and this was most pronounced in farmyard manure treatments. Microbial activity was generally higher in farmyard manure soil indicating that farmyard manure increases labile organic matter pools in soil. Also, biomass of earthworms and microorganisms was increased in farmyard manure soil. The presence of earthworms reduced microbial biomass, suggesting that earthworms feed on microorganisms or/and that earthworms and soil microorganisms competed for similar organic matter pools in soil. The results demonstrate that NPK fertilisation only is insufficient to sustain O. tyrtaeum, whereas long-term fertilisation with farmyard manure enables survival of endogeic species due to an increased pool of utilisable soil organic matter in arable soil.     

C and N turnover of fermented residues from biogas plants in soil in the presence of three different earthworm species (Lumbricus terrestris,Aporrectodea longa,Aporrectodea caliginosa)

A soil microcosm experiment was performed to assess (1) the C- and N- turnover of residues from biogas plants in soils in the presence of three earthworm species (Lumbricus terrestris, Aporrectodea longa and Aporrectodea caliginosa) and (2) the resulting changes in soil chemical and microbiological properties when using these residues as fertilizer in comparison to conventional slurry. Earthworms were exposed in soils, fertilized with an equivalent amount of 120 kg of NH₄-N ha^?1 from: (1) conventional cattle slurry and (2) a fermented residue derived from cattle slurry, grass (silage) and maize. Additional treatments without slurry and earthworms were used as controls.There was considerable evidence that soils fertilized by fermented slurry comprised fewer amounts of readily available nutrients for microbial C and N turnover. We observed significant stimulation of microbial biomass, basal respiration and nitrification in treatments with conventional slurry, especially in the presence of earthworms. However, the stimulation of microbial activity by manure and earthworms were significantly lower in treatments with fermented slurry. Moreover, the results showed clear interactions between different earthworm species and manures. While the biomass of the anecic species (L. terrestris and A. longa) increased in both slurry treatments, the biomass of A. caliginosa (endogeic) decreased, with a significantly stronger biomass decline in treatments with fermented slurry. The metabolic quotients revealed microbial stress metabolism in fermented slurry treatments, predominantly in treatments with A. caliginosa. We conclude that particularly A. caliginosa and soil microorganisms competed for labile C sources in treatments with fermented slurry. An application of these residues as fertilizer might result in a reduction of microbial activity in agricultural soils and in a decline of endogeic earthworms.