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.     

Sublethal effects of imidacloprid on the burrowing behaviour of two earthworm species: Modifications of the 3D burrow systems in artificial cores and consequences on gas diffusion in soil

Earthworms have been termed ‘ecosystem engineers’ (sensu [Jones, C.G., Lawton, J.H., Shachak, M., 1994. Organisms as ecosysem engineers. Oikos 69, 373-386.]) because of the important roles they play in the soil. As a consequence, it is assumed that if earthworms change their behaviour following exposure to pesticides or pollutants this could have a drastic impact on soil functioning. To test this assumption under laboratory conditions, we studied the burrow systems made by two earthworm species (the anecic Aporrectodea nocturna and the endogeic Allolobophora icterica) in artificial soil cores containing imidacloprid, a widely used neonicotinoid insecticide. After 1-month incubation period, the macropores created in the soil core were analyzed by tomography. In order to further characterize transfer properties associated with burrow systems gas diffusion measurements were also carried out. The burrow systems made by the two earthworm species were very different: A. nocturna made more continuous, less branched, more vertical and wider burrows than A. icterica. Some changes to A. nocturna burrow systems were observed after exposure to imidacloprid (they made a smaller burrow system and burrows were more narrow), but only at the highest concentration of imidacloprid used (0.5 mg kg⁻¹). A. icterica worms were more sensitive to imidacloprid and many differences in their burrow systems (length, sinuosity, branching rate and number of burrows) were observed at both concentrations tested (0.1 and 0. 5 mg kg⁻¹). As a consequence, the continuity of the burrow systems made by both species was altered following imidacloprid treatment. Gas diffusion through the A. nocturna soil cores was reduced but no difference in gas diffusion was observed in the A. icterica soil cores.     

Burrowing activity of the earthworms Lumbricus terrestris and Aporrectodea giardi and consequences on C transfers in soil

Two earthworms species, Lumbricus terrestris (epianecic) and Aporrectodea giardi (anecic) were incubated in microcosms with an epigeic ¹³C-labelled litter for 246 d. At the end of the experiment, different soil compartments (surface casts, walls and peripheries of burrows, and surrounding soil) were sampled for ¹³C analysis. Two-dimensional images acquired using X-ray computed tomography allowed to estimate the weight of the ‘burrow wall’ and ‘burrow periphery’ compartments which are required to establish C balance. In the case of L. terrestris, the formed structures were more C litter enriched compared to the other species. The permanent character of the burrow system could lead to a high and constant enrichment of the entire burrow system. As consequence, the percentage of C litter in the ‘burrow wall’ and ‘burrow periphery’ compartments was important in spite of their low volume. The denser system developed by A. giardi resulted in C litter dilution in the whole formed structures. The C litter enrichment decreased with the soil depth, but owing to the intensity of the burrowing activity, the C litter transfers into the ‘burrow walls’ and ‘burrow periphery’ were important and the C litter was homogeneously distributed throughout the whole column.     

Automated observation and analysis of earthworm surface behaviour under experimental habitat quality and availability conditions

Currently, the driving factors of active earthworm dispersal across the soil surface are not sufficiently understood, and distances traversed by individual earthworms have rarely been quantified. Research progress has been hampered by the lack of adequate observation methods as well as fast, objective and quantitative measurements of nocturnal earthworm behaviour.In this work, we report on the potential of a new, automated method using infrared-sensitive webcams and computer image analysis. Nightly surface activities of Lumbricus terrestris L. were monitored quantitatively while manipulating levels of disturbance, burrow availability and congener presence in standard observation units.The automated observation system proved to be simple and inexpensive to build, provided reliable quantitative measures of locomotive behaviour without animal disturbance, and considerably reduced human workload and bias.Waterlogging of the burrow zone stimulated surface activities around and away from the home burrow as compared to habitat disturbance by pesticide application and vibration. However, dispersing earthworms never settled in prefabricated burrows. Surface activity of individuals subjected to waterlogged conditions was influenced by the availability of alternative habitat. Minimal habitat disturbance (vibration) and the presence of conspecific individuals resulted in an increased amount of burrow-anchored, possibly sexually oriented, behaviour, but reciprocal burrow visits and mating were not observed. Pesticide application did not result in dispersal, increased foraging activities or sexual attraction attempts during this short-term study.The presence, extent and mechanisms of the remote assessment of important cues in the surroundings of the home burrow by L. terrestris need further research. Infrared monitoring opens new research avenues of earthworm surface behaviour studies (e.g. density-dependent dispersal, habitat selection, (re)colonization, invasion).     

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.     

The drilosphere concept: Fine-scale incorporation of surface residue-derived N and C around natural Lumbricus terrestris burrows

Anecic (deep-burrowing) earthworms are important for soil biogeochemical functioning, but the fine-scale spatial range at which they incorporate C and N around their burrows (the drilosphere sensu stricto) needs to be investigated under realistic conditions. We conducted a field experiment to delimit spatially the extent to which soil around natural Lumbricus terrestris burrows is influenced biochemically. We placed plant litter dual-labelled with ¹³C and ¹⁵N stable isotope tracers on L. terrestris burrow openings and we measured residue-derived ¹³C and ¹⁵N in thin concentric layers (0–2, 2–4, 4–8 mm) around burrows with or without a resident earthworm. After 45 days, earthworms were significantly enriched in ¹³C and ¹⁵N as a result of feeding on the plant litter. At 0–5 cm soil depth, soil ¹⁵N concentrations were significantly higher around occupied than unoccupied burrows, and they were significantly higher in all burrow layers (including 4–8 mm) than in bulk soil (50–75 mm from burrow). This suggests that biochemical drilosphere effects of anecic earthworms, at least in the uppermost portion of the burrow, extend farther than the 2 mm layer assumed traditionally.     

Assessment of earthworm contribution to soil hydrology: a laboratory method to measure water diffusion through burrow walls

The capacity for water diffusion in burrow walls (i.e. the coefficient of sorptivity) either burrowed by Lumbricus terrestris (T-Worm) or artificially created (T-Artificial) was studied through an experimental design in a 2D terrarium. In addition, the soil density of earthworm casts, burrow walls (0–3 mm around the burrow) and the surrounding soil (>3 mm) were measured using the method of petroleum immersion. This study demonstrated that the quantity of water which transits through burrows of L. terrestris in the soil matrix was lower than that transited through soil fractures, due to a reduction of soil porosity in burrow walls (compaction: cast > worms burrow walls > surrounding soil > artificial burrow walls). Earthworm behaviour, in particular burrow reuse with associated cast pressing on walls, could explain the larger burrow wall compaction in earthworm burrows. If water diffusion was lower through the compacted burrows, burrow reuse by the worms makes them more stable (worms would maintain the structure over years) than unused burrows. The present experimental design could be used to test and measure the specific differences between earthworm species in their contributions to water diffusion. Probably, these contributions depend on the presumed related-species behaviours which would determine the degree of burrow wall compaction.     

Characterization of the burrow system of the earthworms Lumbricus terrestris and Aporrectodea giardi using X-ray computed tomography and image analysis

 The burrow systems of two earthworm species (Lumbricus terrestris and Aporrectodea giardi) were studied in artificially packed soil columns placed in controlled conditions in the laboratory. At the end of the incubation, which lasted 246 days, the burrow systems were characterized on the undisturbed columns using X-ray computed tomography. This method provided a set of digitized images corresponding to horizontal 3-mm-thick sections. The following parameters were measured using image analysis on each section: number of biopores, their volume, and their individual orientation calculated using an elliptical model of the earthworm channel. The profiles of these parameters through the columns showed that the burrow systems of L. terrestris and A. giardi, which are both anecic species, were very different in terms of total volume, number of burrows, burrow orientation and extension with soil depth. These results led us to conclude that the burrow system of L. terrestris can be considered as a permanent structure whereas that of A. giardi is closer to the burrow system of endogeic species. Received: 12 June 1998     

Impact of soil compaction on earthworm burrow systems using X-ray computed tomography: preliminary study

This study is a first approach of the impact of soil compaction due to trafficking by machinery on earthworm burrow systems. To this end, two experiments were established. In the first one, microcosms were incubated in the laboratory for 70 d with Lumbricus terrestris or Aporrectodea giardi. In the second experiment, soil cores were excavated from a sugar beet field mainly colonised by L. terrestris and Aporrectodea caliginosa. The cores were then artificially compacted at 0.12 MPa or 0.25 MPa (which corresponds to the compaction due to trafficking by machinery in the field) or remained non-compacted. The whole cores were submitted to an X-ray computed tomography scan. This method allowed to compare the characteristics of the entire burrow system (total and mean burrowed length, mean length and number of burrows) and of the burrows themselves (number, area and roundness of pores constituting the burrows) in the compacted and non-compacted cores. The results showed that soil compaction contributes to close numerous pores, reduce mean length of burrows and increase the number of fragmented burrows. We concluded that soil compaction affects to a large extent the functionality of burrow systems by fragmenting them and affecting their continuity. This impact increases with the intensity of compaction. The applied soil compaction mainly affected the burrows located in the upper part of the soil. From a general point of view and especially for the upper part of the cores, horizontal burrows seemed to be more affected by compaction than vertical or oblique ones. Globally, the effects of soil compaction were especially observed at the greatest applied pressure (0.25 MPa). At 0.12 MPa, the damage to the burrows was less obvious.     

Distinct microbial and faunal communities and translocated carbon in Lumbricus terrestris drilospheres

Lumbricus terrestris is a deep-burrowing anecic earthworm that builds permanent, vertical burrows with linings (e.g., drilosphere) that are stable and long-lived microhabitats for bacteria, fungi, micro- and mesofauna. We conducted the first non-culture based field study to assess simultaneously the drilosphere (here sampled as 0–2 mm burrow lining) composition of microbial and micro/mesofaunal communities relative to bulk soil. Our study also included a treatment of surface-applied ¹³C- and ¹⁵N-labeled plant residue to trace the short-term (40 d) translocation of residue C and N into the drilosphere, and potentially the assimilation of residue C into drilosphere microbial phospholipid fatty acids (PLFAs). Total C concentration was 23%, microbial PLFA biomass was 58%, and PLFAs associated with protozoa, nematodes, Collembola and other fauna were 200-to-300% greater in the drilosphere than in nearby bulk soil. Principal components analysis of community PLFAs revealed that distributions of Gram-negative bacteria and actinomycetes and other Gram-positive bacteria were highly variable among drilosphere samples, and that drilosphere communities were distinct from bulk soil communities due to the atypical distribution of PLFA biomarkers for micro- and mesofauna. The degree of microbial PLFA ¹³C enrichment in drilosphere soils receiving ¹³C-labeled residue was highly variable, and only one PLFA, 18:1ω9c, was significantly enriched. In contrast, 11 PLFAs from diverse microbial groups where enriched in response to residue amendment in bulk soil 0–5 cm deep. Among control soils, however, a significant δ¹³C shift between drilosphere and bulk soil at the same depth (5–15 cm) revealed the importance of L. terrestris for translocating perennial ryegrass-derived C into the soil at depth, where we estimated the contribution of the recent grass C (8 years) to be at least 26% of the drilosphere soil C. We conclude that L. terrestris facilitates the translocation of plant C into soil at depth and promotes the maintenance of distinct soil microbial and faunal communities that are unlike those found in the bulk soil.     

Earthworms promote the reduction of Fusarium biomass and deoxynivalenol content in wheat straw under field conditions

A field experiment was conducted to elucidate ecosystem services provided by earthworms on the repression of phytopathogenic and toxinogenic fungi. The study focussed on decomposing Fusarium culmorum-infected and deoxynivalenol (DON)-contaminated wheat straw remaining on the soil surface as part in conservation tillage. Mesocosms were established in the topsoil of a winter wheat field located in Northern Germany, where conservation tillage has been practised for 20 years. Besides a non-earthworm treatment, two earthworm species were inoculated in the mesocosms either separately or combined: Lumbricus terrestris (anecic, detritivorous) and Aporrectodea caliginosa (endogeic, geophagous). The earthworms were exposed either to artificially Fusarium-infected wheat straw highly contaminated with DON or to non-infected straw serving as a control. The experiment was conducted during an eight week period after harvest from mid August to mid October. For both species, the artificially Fusarium-infected and DON-contaminated wheat straw was a more attractive food source than the non-infected control. In contrast to A. caliginosa, L. terrestris incorporated infected straw faster into the soil compared to control straw. Furthermore, the reduction of Fusarium biomass and DON concentration in wheat straw was significantly higher in the presence of L. terrestris than in treatments with A. caliginosa and without earthworms. Here, no significant differences could be measured between the Fusarium biomass and DON concentration in wheat straw. A. caliginosa seems not to be relevant for the reduction of Fusarium biomass and DON concentration. We concluded that amongst earthworms, anecic detritivorous species are the drivers to compensate possible negative consequences (like crop infection) of conservation tillage. They take an important role in the control of phytopathogenic and toxinogenic fungi surviving on plant residues and in the degradation of their mycotoxins.     

Earthworms bring compacted and loose soil to a similar mechanical state

We tested the hypothesis that earthworms stabilise loose soil and loosen compacted soil to a similar mechanical state. Casts collected from initially loose soil (980 kg m⁻³) had 10-fold greater viscosity (31 kPa s) and 5-fold greater yield stress (200 Pa) than a control soil without worms. Lumbricus terrestris; Dendrobaena sp. and Aporrectodea longa were all investigated, with no difference found between species. In compacted soils (1300 kg m⁻³), A. longa produced casts with similar mechanical properties to loose soil, with viscosity and yield stress decreased by 45% compared to the control without worms. Earthworms were shown to bring initially loose and compacted soil to an intermediate mechanical state that is more favourable for structural stability and root growth.     

Assessment of earthworm burrowing efficiency in compacted soil with a combination of morphological and soil physical measurements

Summary Column experiments were carried out to quantify the effect of earthworms on compacted soil. The earthworms (Lumbricus terrestris) were able to burrow into soil which was artificially compacted to a pore volume as low as 40%; they may also penetrate an artificial plough pan deep in the soil. The effect of the burrowing activity of Lumbricus terrestris was quantified by measuring hydraulic conductivities and infiltration rates through the whole soil column (19 cm wide, 40 cm long). Morphological parameters, mainly the vertically projected burrow depth, were correlated with the saturated hydraulic conductivity. The amount of casts deposited by Lumbricus terrestris on the soil surface increased with the degree of soil compaction. The bulk density of casts was always less than that of the original soil.     

Worms from the cold: Lumbricid life stages in boreal clay during frost

The vertical distribution and activity of earthworm life stages were studied in an arable field during 0.5 m deep frost. The anecic Lumbricus terrestris L. were below the frost at the bottom of their home burrows (max. depth 1.0 m) and remained there apparently active. Their burrows were open, free of ice and water. The endogeic Aporrectodea caliginosa Sav., mainly small juveniles, were aestivating in the frost layer, which confirms freeze-tolerance in this species. Large A. caliginosa individuals were actively burrowing below the frost down to 1 m depth at soil temperatures close to +1 °C, frost evidently triggering much deeper burrowing than summer droughts. Demonstrating cold-hardiness, viable cocoons of both A. caliginosa and L. terrestris were obtained within a 0-0.25 m layer, frozen for ca. one month prior to sampling. These two common earthworms of boreal soils seem to over-winter in all life stages and remain active below the frost, potentially contributing to the maintenance of subsoil processes during the winter months.     

Is the soil microbial community related to the basal area of trees in a Scots pine stand?

The main energy sources of soil microorganisms are litter fall, root litter and exudation. The amount on these carbon inputs vary according to basal area of the forest stand. We hypothesized that soil microbes utilizing these soil carbon sources relate to the basal area of trees. We measured the amount of soil microbial biomass, soil respiration and microbial community structure as determined by phospholipid fatty acid (PLFA) profiles in the humus layer (FH) of an even-aged stand of Scots pine (Pinus sylvestris L.) with four different basal area levels ranging from 19.9 m² ha⁻¹ in the study plot Kasper 1 to 35.7 m² ha⁻¹ in Kasper 4. Increasing trend in basal respiration, total PLFAs and fungal-to-bacterial ratio was observed from Kasper 1 to Kasper 3 (basal area 29.2 m² ha⁻¹). The soil microbial community structure in Kasper 3 differed from that of the other study plots.     

Earthworm activity as a determinant for N2O emission from crop residue

Earthworm activity may have an effect on nitrous oxide (N₂O) emissions from crop residue. However, the importance of this effect and its main controlling variables are largely unknown. The main objective of this study was to determine under which conditions and to what extent earthworm activity impacts N₂O emissions from grass residue. For this purpose we initiated a 90-day (experiment I) and a 50-day (experiment II) laboratory mesocosm experiment using a Typic Fluvaquent pasture soil with silt loam texture. In all treatments, residue was applied, and emissions of N₂O and carbon dioxide (CO₂) were measured. In experiment I the residue was applied on top of the soil surface and we tested (a) the effects of the anecic earthworm species Aporrectodea longa (Ude) vs. the epigeic species Lumbricus rubellus (Hoffmeister) and (b) interactions between earthworm activity and bulk density (1.06 vs. 1.61 g cm⁻³). In experiment II we tested the effect of L. rubellus after residue was artificially incorporated in the soil. In experiment I, N₂O emissions in the presence of earthworms significantly increased from 55.7 to 789.1 μg N₂O-N kg⁻¹ soil (L. rubellus; p<0.001) or to 227.2 μg N₂O-N kg⁻¹ soil (A. longa; p<0.05). This effect was not dependent on bulk density. However, if the residue was incorporated into the soil (experiment II) the earthworm effect disappeared and emissions were higher (1064.2 μg N₂O-N kg⁻¹ soil). At the end of the experiment and after removal of earthworms, a drying/wetting and freezing/thawing cycle resulted in significantly higher emissions of N₂O and CO₂ from soil with prior presence of L. rubellus. Soil with prior presence of L. rubellus also had higher potential denitrification. We conclude that the main effect of earthworm activity on N₂O emissions is through mixing residue into the soil, switching residue decomposition from an aerobic and low denitrification pathway to one with significant denitrification and N₂O production. Furthermore, A. longa activity resulted in more stable soil organic matter than L. rubellus.     

Role of soil water content in the carbon and nitrogen dynamics of Lumbricus terrestris L. burrow soil

We evaluated the role of soil water content in controlling C and N dynamics within the drilosphere created by the anecic earthworm Lumbricus terrestris (L.). Mesocosms (volume = 3.1 l) were each amended with corn litter and three earthworms. Control treatments received no earthworms and no other earthworm species were present in the soil. WET and DRY treatments received a total of 9.25 cm and 3.25 cm of water, respectively. Water was added on weeks 1, 3, 7, and 10 at a rate of 2.0 cm per mesocosm for WET treatments and 0.5 cm per mesocosm for DRY treatments. Mesocosms were sampled destructively after incubation at 18–20 °C for 0, 3, 7, and 13 weeks. The water content of WET burrow soil ranged from 0.12 g g⁻¹ to 0.18 g g⁻¹ and was significantly higher than in the DRY treatment throughout the incubation period. The live weight of earthworms was significantly higher in the WET treatment only on week 13, whereas litter consumption was significantly lower in the DRY treatment for week 13. Carbon mineralization, measured as CO₂ evolved after a 24-h incubation, was consistently higher in WET than in DRY burrow soil. Effects of differences in soil water content were also apparent for biomass C and metabolic quotient. Soil water content did no affect the total C concentration of burrow soil. DRY burrow soil had consistently lower levels of nitrate than WET soil throughout the experiment. Lower levels of ammonium and inorganic N were observed for WET burrow soil on weeks 3 and 7. Water content did not have a significant effect on burrow soil total N. We concluded that the water content of the drilosphere affects both C and N dynamics and can affect the speciation of inorganic N; yet, the effects of soil water content do not appear to result from differences in the feeding activities of anecic earthworms.     

Earthworms change the distribution and availability of phosphorous in organic substrates

In laboratory controlled soil microcosms, the distribution and availability of phosphorous (P) were determined in the surface-casts and the burrows-linings of the anecic earthworm L. terrestris and were compared with non-ingested soil. To simulate more realistic earthworm community conditions, a combination of L. terrestris plus the endogeic A. caliginosa was tested. For a 2-month period, the earthworms were given two organic food substrates: rye-grass littered onto the soil surface and sewage sludge mixed with soil. The following treatments were designed: (i) soil alone (S), (ii) soil and sewage sludge (SS), soil and rye-grass litter (SL), and (iv) soil, litter and sludge (SSL). Analyses were performed for P contents (total, available and organic), organic matter content (organic carbon, C_org and total nitrogen, N_tot) and the two acid and alkaline phosphatase activities (AcPA and AkPA). Earthworms enhanced AcPA and were also responsible for additional AkPA in soil. The two AcPA and AkPA increased not only in surface-casts but also in burrows-linings that paralleled with the decrease of organic P in SL and SSL treatments. The stimulation of AcPA began quickly and declined rapidly in casts (from 19 to 8 μmol phenol g⁻¹ dry wt h⁻¹, respectively at week 2 and 8 in the SL treatment) but it was initiated later and maintained at a high level for longer in burrows (more than 10 μmol phenol g⁻¹ dry wt h⁻¹ at week 8 in the SL treatment). Significant positive correlations were found between the AkPA activities and N_tot contents (r=0.95, p=0.001) and to a lesser extend with C_org contents (r=0.76, p=0.05) in casts from the SL treatment, while AcPA significantly correlated with N_tot (r=0.91, p=0.004) but not with C_org (r=0.72, p=0.06). P availability was always highest in casts. However, the available P contents decreased sharply over time in casts and were still low in burrow-linings, suggesting that a large part of inorganic P produced was rapidly immobilized for the microbial growth. Total P content was unchanged except in the SL treatment in which it increased in casts and burrows (ca. 725 μg g⁻¹, at week 4). Organic P was first the highest in casts and then decreased over time (from 168 at week 1 to 140 μg g⁻¹ at week 8 in the SL treatment). This study illustrates that earthworms facilitate P transfer downward increasing a P patchy distribution in the soil, and significantly change the biogeochemical status of P (availability, organic phosphorous pool, AcPA activities) in certain hot spots such as casts and burrow-linings.     

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.     

The effect of two endogeic earthworm species on zinc distribution and availability in artificial soil columns

The objective of this study was to determine the impact of earthworm bioturbation on the distribution and availability of zinc in the soil profile.Experiments were carried out with Allolobophora chlorotica and Aporrectodea caliginosa in 24 perspex columns (∅ 10 cm), filled with 20-23 cm non-polluted soil (OM 2%, clay 2.9%, pH 0.01 M CaCl₂ 6.4), that was covered by a 3-5 cm layer of aged zinc spiked soil (500 mg Zn/kg dry soil) and another 2 cm non-polluted soil on top. After 80 and 175 days, columns were sacrificed and each cm from the top down to a depth of 15 cm was sampled. Earthworm casts, placed on top of the soil, were collected. Each sample was analyzed for total and CaCl₂-exchangeable zinc concentrations.Effects of earthworm bioturbation were most pronounced after 175 days. For A. chlorotica, total and CaCl₂-exchangeable zinc concentrations in the polluted layers were lower with than without earthworms. Total zinc concentrations in the non-polluted layers were higher in columns with earthworms. Casts of A. chlorotica collected on the soil surface showed slightly higher total zinc concentrations than non-polluted soil. Casts were found throughout the whole column. For A. caliginosa there were no differences in total zinc concentration between columns with and without earthworms. CaCl₂-exchangeable zinc concentrations in the polluted layers were lower for columns with earthworms. Casts were mainly placed on top of the soil and contained total zinc concentrations intermediate between those in non-polluted and polluted soil layers.This study shows that different endogeic earthworm species have different effects on zinc distribution and availability in soils. A. chlorotica transfers soil throughout the whole column, effectively mixing it, while A. caliginosa decreases metal availability and transfers polluted soil to the soil surface.