Characteristics of earthworms (Eisenia fetida) in PAHs contaminated soil amended with sewage sludge or vermicompost

Earthworms can be used to remove polycyclic aromatic hydrocarbons (PAHs) from soil, but this might affect their survival and they might accumulate the contaminants. Sterilized and unsterilized soil was contaminated with phenanthrene (Phen), anthracene (Anth) and benzo(a)pyrene (BaP), added with or without Eisenia fetida, sewage sludge or vermicompost. Survival, growth, cocoon formation and concentrations of PAHs in the earthworms were monitored for 70 days. Addition of sewage sludge to sterilized or unsterilized soil maintained the number of earthworms and their survival was 94%. The addition of sludge significantly increased the weight of earthworms 1.3 times compared to those kept in the unamended soil or in soil amended with vermicompost. The weight of earthworms was significantly lower in sterilized than in unsterilized soil. Cocoons were only detected when sewage sludge was added to unsterilized soil. A maximum concentration of 62.3 μg Phen kg⁻¹ was found in the earthworms kept in sterilized soil amended with vermicompost after 7 days and 22.3 μg Phen kg⁻¹ when kept in the unamended unsterilized soil after 14 days. Concentrations of Phen in the earthworms decreased thereafter and ≤2 μg kg⁻¹ after 28 days. A maximum Anth concentration of 82.5 μg kg⁻¹ was found in the earthworms kept in sterilized soil amended with vermicompost and 45.8 μg Anth kg⁻¹ when kept in the unamended unsterilized soil after 14 days. A maximum concentration of 316 μg BaP kg⁻¹ was found in the earthworms kept in sterilized soil amended with vermicompost after 56 days and 311 μg BaP kg⁻¹ when kept in the unsterilized soil amended with vermicompost after 28 days. The amount of BaP in the earthworm was generally largest after 28 days, but after 70 days still 60 μg kg⁻¹ was found in E. fetida when kept in the sterilized soil amended with sewage sludge. It was found that E. fetida survived in PAHs contaminated soil and accumulated only small amounts of the contaminants, but sewage sludge was required as food for its survival and cocoon production.     

Maize residue application reduces negative effects of soil salinity on the growth and reproduction of the earthworm Aporrectodea trapezoides,in a soil mesocosm experiment

We studied the effects of maize residue application on some life-cycle parameters of the earthworm Aporrectodea trapezoides in saline agricultural soils with electrical conductivity (EC) ranging from 1.58 to 7.35 dS m⁻¹. This experiment was carried out under controlled laboratory conditions for 150 days. Results showed that soil salinity significantly affected the growth and reproduction of earthworms, decreasing survival, numbers and mean fresh weights of adults, juveniles and cocoons. Maize residue application gave a greater survival of earthworms at all salinity levels, but the differences were only significant at an EC of 7.35 dS m⁻¹, although the mean weight of adult earthworms was significantly increased by maize residue application at all salinity levels. At an EC of 1.58 dS m⁻¹ and 3.35 dS m⁻¹, the application of maize residues gave significantly higher numbers of cocoons and juveniles, but in soils with 5.26 dS m⁻¹ and 7.35 dS m⁻¹ earthworms did not produce any cocoons over the experimental period, irrespective of maize residue application. These results indicated that maize residue application alleviated the negative effects of soil salinity on the growth and reproduction of A. trapezoides up to 3.35 dS m⁻¹, above which maize residues only increased the growth but not on the reproduction of earthworms.     

Deep-burrowing earthworm additions changed the distribution of soil organic carbon in a chisel-tilled soil

We investigated the influence of earthworms on the three-dimensional distribution of soil organic carbon (SOC) in a chisel-tilled soil. By burrowing, foraging, and casting at the surface and throughout the soil, anecic earthworms such as Lumbricus terrestris L. may play a major role in regulating the spatial distribution of organic matter resources both at the surface and within the soil. In the fall of 1994, we manipulated ambient earthworm communities, which were without deep burrowing species, by adding 100 earthworm individuals m⁻² in spring and fall for 3 years. Overall, the biomass of L. terrestris was increased with earthworm additions and total earthworm biomass declined compared with ambient control treatments. To investigate the spatial variability in soil organic carbon due to this shift in earthworm community structure, we sampled soil on a 28×24 cm grid from the surface to 40 cm in four layers, 10 cm deep. Samples were analyzed for total carbon. We found that additions of anecic earthworms significantly increased average soil organic carbon content from 16.1 to 17.9 g C kg⁻¹ for the 0–10 cm soil, and from 12.4 to 14.7 g kg⁻¹ at 10–20-cm depth, and also changed the spatial distribution of soil organic carbon from uniform to patchy, compared with the ambient treatment.     

Toxicological effects of TiO2 and ZnO nanoparticles in soil on earthworm Eisenia fetida

Nanoparticles (NPs) of TiO₂ and ZnO are receiving increasing attention due to their widespread applications. To evaluate their toxicities to the earthworm Eisenia fetida (Savigny, 1826) in soil, artificial soil systems containing distilled water, 0.1, 0.5, 1.0 or 5.0 g kg⁻¹ of NPs were prepared and earthworms were exposed for 7 days. Contents of Zn and Ti in earthworm, activities of antioxidant enzymes, DNA damage to earthworm, activity of cellulase and damage to mitochondria of gut cells were investigated after acute toxicity test. The results from response of the antioxidant system combined with DNA damage endpoint (comet assay) indicated that TiO₂ and ZnO NPs could induce significant damage to earthworms when doses were greater than 1.0 g kg⁻¹. We found that Ti and Zn, especially Zn, were bioaccumulated, and that mitochondria were damaged at the highest dose in soil (5.0 g kg⁻¹). The activity of cellulase was significantly inhibited when organisms were exposed to 5.0 g kg⁻¹ of ZnO NPs. Our study demonstrates that both TiO₂ and ZnO NPs exert harmful effects to E. fetida when their levels are higher than 1.0 g kg⁻¹ in soil and that toxicity of ZnO NPs was higher than TiO₂.     

Interactions between residue placement and earthworm ecological strategy affect aggregate turnover and N2O dynamics in agricultural soil

Previous laboratory studies using epigeic and anecic earthworms have shown that earthworm activity can considerably increase nitrous oxide (N₂O) emissions from crop residues in soils. However, the universality of this effect across earthworm functional groups and its underlying mechanisms remain unclear. The aims of this study were (i) to determine whether earthworms with an endogeic strategy also affect N₂O emissions; (ii) to quantify possible interactions with epigeic earthworms; and (iii) to link these effects to earthworm-induced differences in selected soil properties. We initiated a 90-day ¹⁵N-tracer mesocosm study with the endogeic earthworm species Aporrectodea caliginosa (Savigny) and the epigeic species Lumbricus rubellus (Hoffmeister). ¹⁵N-labeled radish (Raphanus sativus cv. Adagio L.) residue was placed on top or incorporated into the loamy (Fluvaquent) soil. When residue was incorporated, only A. caliginosa significantly (p < 0.01) increased cumulative N₂O emissions from 1350 to 2223 μg N₂O-N kg⁻¹ soil, with a corresponding increase in the turnover rate of macroaggregates. When residue was applied on top, L. rubellus significantly (p < 0.001) increased emissions from 524 to 929 μg N₂O-N kg⁻¹, and a significant (p < 0.05) interaction between the two earthworm species increased emissions to 1397 μg N₂O-N kg⁻¹. These effects coincided with an 84% increase in incorporation of residue ¹⁵N into the microaggregate fraction by A. caliginosa (p = 0.003) and an 85% increase in incorporation into the macroaggregate fraction by L. rubellus (p = 0.018). Cumulative CO₂ fluxes were only significantly increased by earthworm activity (from 473.9 to 593.6 mg CO₂-C kg⁻¹ soil; p = 0.037) in the presence of L. rubellus when residue was applied on top. We conclude that earthworm-induced N₂O emissions reflect earthworm feeding strategies: epigeic earthworms can increase N₂O emissions when residue is applied on top; endogeic earthworms when residue is incorporated into the soil by humans (tillage) or by other earthworm species. The effects of residue placement and earthworm addition are accompanied by changes in aggregate and SOM turnover, possibly controlling carbon, nitrogen and oxygen availability and therefore denitrification. Our results contribute to understanding the important but intricate relations between (functional) soil biodiversity and the soil greenhouse gas balance. Further research should focus on elucidating the links between the observed changes in soil aggregation and controls on denitrification, including the microbial community.     

The combined effects of earthworms and arbuscular mycorrhizal fungi on microbial biomass and enzyme activities in a calcareous soil spiked with cadmium

Earthworms and arbuscular mycorrhizal fungi (AMF) are known to independently affect soil microbial and biochemical properties, in particular soil microbial biomass (SMB) and enzymes. However, less information is available about their interactive effects, particularly in soils contaminated with heavy metals such as cadmium (Cd). The amount of soil microbial biomass C (MBC), the rate of soil respiration (SRR) and the activities of urease and alkaline phosphatase (ALP) were measured in a calcareous soil artificially spiked with Cd (10 and 20 mg Cd kg⁻¹), inoculated with earthworm (Lumbricus rubellus L.), and AMF (Glomus intraradices and Glomus mosseae species) under maize (Zea mays L.) crop for 60 days. Results showed that the quantity of MBC, SRR and enzyme activities decreased with increasing Cd levels as a result of the elevated exchangeable Cd concentration. Earthworm addition increased soil exchangeable Cd levels, while AMF and their interaction with earthworms had no influence on this fraction of Cd. Earthworm activity resulted in no change in soil MBC, while inoculation with both AMF species significantly enhanced soil MBC contents. However, the presence of earthworms lowered soil MBC when inoculated with G. mosseae fungi, showing an interaction between the two organisms. Soil enzyme activities and SRR values tended to increase considerably with the inoculation of both earthworms and AMF. Nevertheless, earthworm activity did not affect ALP activity when inoculated with G. mosseae fungi, while the presence of earthworm enhanced urease activity only with G. intraradices species. The increases in enzyme activities and SRR were better ascribed to changes in soil organic carbon (OC), MBC and dissolved organic carbon (DOC) contents. In summary, results demonstrated that the influence of earthworms alone on Cd availability is more important than that of AMF in Cd-polluted soils; and that the interaction effects between these organisms on soil microorganism are much more important than on Cd availability. Thus, the presence of both earthworms and AMF could alleviate Cd effects on soil microbial life.     

Organic nitrogen stimulates the heterotrophic nitrification rate in an acidic forest soil

Many previous studies have demonstrated that heterotrophic nitrification processes play an important role in the production of NO₃⁻ in acidic soils. However, it is not clear whether a low concentration of nitrogenous organic compounds support heterotrophic nitrification processes in natural soils. In this study, we performed an ¹⁵N tracer experiment with a glycine concentration gradient (20, 40, 80, and 160 mg N kg⁻¹) to investigate the effect of the organic nitrogen concentration on the heterotrophic nitrification rate and its relative contribution to the total nitrification of the studied acidic forest soil. This experiment demonstrated that ¹⁵N–NO₃⁻ accumulated over time with all nitrogen treatments in the presence of acetylene, confirming that heterotrophic nitrification occurred even at a low organic nitrogen concentration (20 mg kg⁻¹) in the studied acidic forest soil. In the presence of acetylene, the ¹⁵N–NO₃⁻ concentration in the 20 and 40 mg kg⁻¹ glycine-N treatments was significantly lower than in the 80 and 160 mg kg⁻¹ glycine-N treatments (p < 0.05), indicating that a high organic nitrogen concentration stimulated the heterotrophic nitrification rate. There was no significant difference in the average contribution of heterotrophic nitrification to total nitrification among the different nitrogen treatments, suggesting that the organic nitrogen concentration did not affect the relative contribution of heterotrophic nitrification to total nitrification in the studied acidic soil. Our results confirmed that a low concentration of organic N (20 mg kg⁻¹) supported heterotrophic nitrification in the studied soil. The organic nitrogen concentration stimulates the heterotrophic nitrification rate, but does not affect the relative contribution of heterotrophic nitrification to total nitrification in the studied acidic soil.     

Interaction between earthworms and arbuscular mycorrhizal fungi on the degradation of oxytetracycline in soils

The interactive impact of earthworms (Eisenia fetida) and arbuscular mycorrhizal fungi (Rhizophagus intraradices, AM fungi) on the degradation of oxytetracycline (OTC) in soils was studied under greenhouse conditions. Treatments included maize plants inoculated vs. not inoculated with AM fungi and treated with or without earthworms at low (1 mg kg⁻¹ soil DM) or high (100 mg kg⁻¹ soil DM) OTC rates. The root colonization rate, the hyphal density of mycorrhizae, the residual OTC concentration in soils, catalase, dehydrogenase, urease, soil microbial biomass C, Shannon–Wiener index (H) for microbial communities from T-RFLP profiles were measured at harvest. The results indicated that earthworms and AM fungi would individually or interactively enhance OTC decomposition and significantly decreased the residual OTC concentration at both high and low OTC rates. Both earthworms and AM fungi could promote the degradation of OTC by increasing soil microbial biomass C at both high and low OTC rates. The effect of soil enzyme activity and soil microbial diversity on OTC decomposition was different between high and low OTC rates. Hyphomicrobium and Bacillus cereus were dominant bacteria, and Thielavia and Chaetomium were dominant phyla of fungi at all occasions. Earthworm activity stimulated the growth of Hyphomicrobium and Thielavia, while AM fungi may stimulate B. cereus, Thielavia and Chaetomium, resulting in greater OTC decomposition. The interaction between earthworms and AM fungi in affecting the degradation of OTC may be attributed to different mechanisms, depending on soil microbial biomass, function (enzyme activity) and communities (the abundance of Hyphomicrobium, B. cereus, Thielavia and Chaetomium) in the soil.     

Trace metals and metalloids in forest soils and exotic earthworms in northern New England,USA

Trace metals and metalloids (TMM) in forest soils and invasive earthworms were studied at 9 sites in northern New England, USA. Essential (Cu, Mo, Ni, Zn, Se) and toxic (As, Cd, Pb, Hg, U) TMM concentrations (mg kg⁻¹) and pools (mg m⁻²) were quantified for organic horizons (forest floor), mineral soils and earthworm tissues. Essential TMM tissue concentrations were greatest for mineral soil-feeding earthworm Octolasion cyaneum. Toxic TMM tissue concentrations were highest for organic horizon-feeding earthworms Dendobaena octaedra, Aporrectodea rosea and Amynthas agrestis. Most earthworm species had attained tissue concentrations of Pb, Hg and Se potentially hazardous to predators. Bioaccumulation factors were Cd > Se > Hg > Zn > Pb > U > 1.0 > Cu > As > Mo > Ni. Only Cd, Se, Hg and Zn were considered strongly bioaccumulated by earthworms because their average bioaccumulation factors were significantly greater than 1.0. Differences in bioaccumulation did not appear to be caused by soil concentrations as earthworm TMM tissue concentrations were poorly correlated with TMM soil concentrations. Instead, TMM bioaccumulation appears to be species and site dependent. The invasive A. agrestis had the greatest tissue TMM pools, due to its large body mass and high abundance at our stands. We observed that TMM tissue pools in earthworms were comparable or exceeded organic horizon TMM pools; earthworm tissue pools of Cd were up 12 times greater than in the organic horizon. Thus, exotic earthworms may represent an unaccounted portion and flux of TMM in forests of the northeastern US. Our results highlight the importance of earthworms in TMM cycling in northern forests and warrant more research into their impact across the region.     

Role of earthworms in regenerating soil structure after compaction in reduced tillage systems

New non-tillage or reduced tillage agricultural practises are being increasingly adopted but generally result in higher soil compaction. Due to their recognised physical influence mainly through burrow creation, it is often claimed that earthworm activity could alleviate soil compaction in these systems. To put this assumption to the test, an experimental compaction event was carried out on one plot of arable land. The abundance and biomass of earthworms were evaluated in compacted (under wheel tracks) and non-compacted (between wheel tracks) zones, seven times over a two-year period. In addition, the functional consequences of earthworm activity, defined by burrow abundance assessed in 2D and 3D and water infiltration, were measured three times over the same period. The short-term (less than three months) effects of the compaction were clear: soil bulk density increased from 1.46 to 1.57 g cm⁻³, the abundance and biomass of earthworms were greatly reduced (−40% and −70% respectively) and the number and continuity of macroporosity were lower under wheel tracks at least until a depth of 30 cm. After these initial detrimental effects, we observed a rapid recovery of earthworm populations with no statistical difference between compacted and control zones more than three months after the compaction. However, the recovery of soil functional properties linked to earthworm activity, macroporosity and water infiltration, was much slower and took between 12 and 24 months. Despite these modifications, there were no significant changes in soil bulk density with time during the two-year period. This study demonstrates that earthworms are important actors in the regeneration of compacted soil. Although the complete regeneration of compacted soil by earthworms is a slow process, agricultural practises that promote earthworm density and activity should be encouraged in reduced or minimum tillage systems.     

Effect of tillage on earthworms over short- and medium-term in conventional and organic farming

Earthworms play an important role in many soil functions and are affected by soil tillage in agricultural soils. However, effects of tillage on earthworms are often studied without considering species and their interactions with soil properties. Furthermore, many field studies are based on one-time samplings that do not allow for characterisation of temporal variation. The current study monitored the short (up to 53 days) and medium term (up to 4 years) effects of soil tillage on earthworms in conventional and organic farming. Earthworm abundances decreased one and three weeks after mouldboard ploughing in both conventional and organic farming, suggesting direct and indirect mechanisms. However, the medium-term study revealed that earthworm populations in mouldboard ploughing systems recovered by spring. The endogeic species Aporrectodea caliginosa strongly dominated the earthworm community (76%), whereas anecic species remained <1% of all earthworms in all tillage and farming systems over the entire study. In conventional farming, mean total earthworm abundance was not significantly different in reduced tillage (153 m⁻²) than mouldboard ploughing (MP; 130 m⁻²). However, reduced tillage in conventional farming significantly increased the epigeic species Lumbricus rubellus from 0.1 m⁻² in mouldboard ploughing to 9 m⁻² averaged over 4 years. Contrastingly, in organic farming mean total earthworm abundance was 45% lower in reduced tillage (297 m⁻²) than MP (430 m⁻²), across all sampling dates over the medium-term study (significant at 3 of 6 sampling dates). Reduced tillage in organic farming decreased A. caliginosa from 304 m⁻² in mouldboard ploughing to 169 m⁻² averaged over 4 years (significant at all sampling dates). Multivariate analysis revealed clear separation between farming and tillage systems. Earthworm species abundances, soil moisture, and soil organic matter were positively correlated, whereas earthworm abundances and penetration resistance where negatively correlated. Variability demonstrated between sampling dates highlights the importance of multiple samplings in time to ascertain management effects on earthworms. Findings indicate that a reduction in tillage intensity in conventional farming affects earthworms differently than in organic farming. Differing earthworm species or ecological group response to interactions between soil tillage, crop, and organic matter management in conventional and organic farming has implications for management to maximise soil ecosystem functions.     

Influence of acetamiprid on soil enzymatic activities and respiration

The effect of a new pesticide, acetamiprid, applied at normal field concentration (0.5 mg kg⁻¹ dried soil) and at high concentration (5 and 50 mg kg⁻¹ dried soil), on soil enzyme activities and soil respiration in upland soil was studied. The results showed that acetamiprid had a strong negative influence on soil respiration and phosphatase activity, and the enzyme activities in soil treated with 5 and 50 mg kg⁻¹ dry soil were significantly (P < 0.05) lower than the CK over the course of incubation. The 7-, 14-, and 35-day EC10 for phosphatase were 11, 15, and 11 mg kg⁻¹ dry soil, respectively. The 21-day EC10 and EC50 for soil respiration was 0.005 and 83 mg kg⁻¹ dry soil. The activity of dehydrogenase was enhanced after acetamiprid application for 2 weeks and the enzyme activities in samples treated with 0.5, 5 and 50 mg kg⁻¹ dry soil was about 2.5-, 1.5- and 2-fold to that of the control on sample day 28. Variance of urease and catalase had no distinct relationship with the application concentration. The activity of proteinase was not significantly affected within the first 2 weeks but inhibited from the fourth week after acetamiprid application and was only 0.45-fold to that of the control on sample day 28. Overall, acetamiprid at normal field dose would not pose a toxicological threat to soil enzymes, but a certain potential threat to soil respiration.     

Methyl-mercury affects microbial activity and biomass,bacterial community structure but rarely the fungal community structure

Monomethyl-mercury is one of the most toxic compounds. Methylation of Hg usually appears under anoxic conditions. In Swiss forest soils, methyl-Hg concentrations of up to 3 μg kg⁻¹ soil dw have been observed, but the impact of methyl-Hg on soil microorganisms have rarely been examined so far. In this study, we investigated the effect of increasing concentrations of methyl-Hg (0, 5, 20, 90 μg kg⁻¹ soil dw) on the microbial communities in various forest soils differing in their physico-chemical properties. Experiments were conducted in microcosms under controlled conditions and the basal respiration (BR), the microbial biomass carbon (MBC) and the bacterial and fungal community structures using T-RFLP-profiling were investigated. BR was significantly affected by methyl-Hg. In general, the BR increased with increasing methyl-Hg concentrations, whereas the MBC was significantly reduced. Bacterial communities were more sensitive to methyl-Hg than fungal communities. In five out of seven soils, the bacterial community structures differed significantly between the treatments whereas the fungal communities did not. The impact of methyl-Hg on the soil bacterial communities was site specific. In one soil, a methyl-Hg concentration of already 5 μg kg⁻¹ soil dw significantly affected the relative abundance of 13% bacterial operational taxonomic units (OTU), whereas in other soils concentrations of even 90 μg kg⁻¹ soil dw rarely affected the abundance of OTUs. In this study, for the first time, the impact of methyl-Hg on soil bacterial and fungal communities in forest soils was assessed. We showed that its impact strongly depends on the physico-chemical conditions of the soil and that bacterial communities were more sensitive to methyl-Hg than fungi.     

Effects of azoxystrobin,chlorothalonil, and ethoprophos on the reproduction of three terrestrial invertebrates using a natural Mediterranean soil

The potential terrestrial toxicity of three pesticides, azoxystrobin, chlorothalonil, and ethoprophos was evaluated using reproduction ecotoxicological tests with different non-target species: the collembolan Folsomia candida, the earthworm Eisenia andrei, and the enchytraeid Enchytraeus crypticus. All reproduction tests were performed with natural soil from a Mediterranean agricultural area (with no pesticide residues) in order to improve the relevance of laboratory data to field conditions. Controls were performed with natural and standard artificial soil (OECD 10% OM). The fungicide azoxystrobin showed the highest toxicity to earthworms (EC₅₀ = 42.0 mg a.i. kg⁻¹ dw soil). Collembolans were the most sensitive taxa in terms of sublethal effects of chlorothalonil with an EC₅₀ of 31.1 mg a.i. kg⁻¹ dw soil followed by the earthworms with an EC₅₀ of 40.9 mg a.i. kg⁻¹ dw soil. The insecticide ethoprophos was the most toxic to collembolans affecting their reproduction with an EC₅₀ of 0.027 mg a.i. kg⁻¹ dw soil. Enchytraeids were generally the least sensitive of the three species tested for long-term effects. Earthworms were not always the most sensitive species, emphasizing the need to increase the number of mandatory assays with key non-target organisms in the environmental risk assessment of pesticides.     

Use of urea coated with natural products to inhibit urea hydrolysis and nitrification in soil

Laboratory incubation experiments were conducted with uncoated urea or urea coated with dementholized oil (DMO), pitch (the mint oil discard), terpenes (the products of menthol mint oil), or dicyandiamide (DCD) to study the retardation of urea hydrolysis and nitrification in soil. Two levels (0.5 and 1 %) of coating were tested. Urea was applied at a rate of 200 mg kg⁻¹ of dry weight of soil. The urea hydrolysis and nitrification processes were inhibited by all three natural products. All the three natural products viz., DMO, terpenes, and pitch significantly retarded urease activity of soil.     

Importance of heterotrophic nitrification and dissimilatory nitrate reduction to ammonium in a cropland soil: Evidences from a 15N tracing study to literature synthesis

Future climate change is predicted to influence soil moisture regime, a key factor regulating soil nitrogen (N) cycling. To elucidate how soil moisture affects gross N transformation in a cultivated black soil, a ¹⁵N tracing study was conducted at 30%, 50% and 70% water-filled pore space (WFPS). While gross mineralization rate of recalcitrant organic N (N_rec) increased from 0.56 to 2.47 mg N kg⁻¹ d⁻¹, the rate of labile organic N mineralization declined from 4.23 to 2.41 mg N kg⁻¹ d⁻¹ with a WFPS increase from 30% to 70%. Similar to total mineralization, no distinct moisture effect was found on total immobilization of ammonium, which primarily entered the N_rec pool. Nitrate (NO₃⁻) was mainly produced via autotrophic nitrification, which was significantly stimulated by increasing WFPS. Unexpectedly, heterotrophic nitrification was observed, with the highest rate of 1.06 mg N kg⁻¹ d⁻¹ at 30% WFPS, contributing 31.8% to total NO₃⁻ production, and decreased with WFPS. Dissimilatory nitrate reduction to ammonium (DNRA) increased from near zero (30% WFPS) to 0.26 mg N kg⁻¹ d⁻¹ (70% WFPS), amounting to 16.7–92.9% of NO₃⁻ consumption. A literature synthetic analysis from global multiple ecosystems showed that the rates of heterotrophic nitrification and DNRA in test soil were comparative to the forest and grassland ecosystems, and that heterotrophic nitrification was positively correlated with precipitation, soil organic carbon (SOC) and C/N, but negatively with pH and bulk density, while DNRA showed positive relationships with precipitation, clay, SOC, C/NO₃⁻ and WFPS. We suggested that low pH and bulk density and high SOC and C/N in test soil might favor heterotrophic nitrification, and that C and NO₃⁻ availability together with anaerobic condition were crucial for DNRA. Overall, our study highlights the role of moisture in regulating gross N turnover and the importance of heterotrophic nitrification for NO₃⁻ production under low moisture and DNRA for NO₃⁻ retention under high moisture in cropland.     

Earthworm colonisation of abandoned arable soil polluted by copper

The distribution, density and biomass of earthworms were investigated at the copper polluted site, Hygum (Denmark). In 1994, shortly after farming of the area was abandoned, only four earthworm species were present and their distribution was restricted to areas where copper concentration did not exceed 200 mg kg^?1 dry soil. Sixteen years later (in 2010), without any agricultural activity, ten species of earthworms were found, in particular, epigeic species were present where soil copper concentrations reached >1000 mg kg^?1 dry soil.     

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.     

Biological and physical effects of non-native earthworms on nitrogen cycling in riparian soils

Global nitrogen cycling is being altered by anthropogenic disturbances including invasion by non-native species. European and Asian earthworms have invaded northern temperate forests in North America with dramatic consequences for litter thickness, forest floor plant diversity, and soil nitrogen cycling. Invasive earthworms present at the boundary of terrestrial and aquatic ecosystems (i.e., riparian zones) may alter the flux of nitrogen into adjacent aquatic ecosystems. We examined how nitrogen cycling in riparian soil responds to amendments of invasive earthworms or artificial earthworm burrows. In earthworm-free riparian plots (0.25 m²), we established treatments of invasive earthworms (60 g fresh mass·m⁻²), artificial burrows (120 m⁻²), or control plots and sampled the plots after 30 days. Before and after treatment application we measured major soil characteristics (water-filled pore space, organic matter, and pH), nitrogen pools (exchangeable NH₄⁺ and NO₃⁻), and nitrogen transformation rates (net N-mineralization, net nitrification, and denitrification). Exchangeable NH₄⁺ and NO₃⁻ changed through time but did not differ among treatments. Net N-mineralization and net nitrification rates did not change through time and were similar across all treatments. However, denitrification rates in plots with added earthworms were 4 times greater than rates in control and burrow-only plots, which represents a large rapid increase in gaseous nitrogen flux out of these riparian soils. For all response variables, artificial burrows responded similarly to control plots, suggesting that earthworm biological activity (i.e., feeding, excretion, and mucus production) rather than physical effects (i.e., burrowing and soil aeration) drove the changes in nitrogen cycling. Examination of soil nitrogen pool and flux measurements suggest that this increase in denitrification was coupled with NH₄⁺ consumption by nitrifying bacteria, but future studies are needed to confirm this hypothesis. We conclude that the activity of invasive earthworms in riparian zones can increase the flux of N out of riparian zones, but the hydrologic context of the riparian zone (e.g., pore-water residence time) ultimately controls whether denitrification or nitrate leaching is the dominant flux of N.     

Can commercial mulches be reservoirs of invasive earthworms? Promotion of ligninolytic enzyme activity and survival of Amynthas agrestis (Goto and Hatai, 1899)

Amynthas agrestis is an exotic, invasive earthworm in North America that has been associated with horticulture settings as well as damage to forest soil. An experiment was conducted to find out whether A. agrestis, an earthworm commonly found in mulches in Vermont, stimulates ligninolytic enzymes in the presence of commercial wood mulches. Mesocosms filled with a sandy loam soil were topped with either spruce, cedar or pine mulch. Half of the mesocosms received juvenile A. agrestis, the other half did not. After 7 weeks soils were analyzed for phenoloxidase and peroxidase activity. Most A. agrestis survived and developed into adults during the incubation period. Significantly greater phenoloxidase activity was detected in soils with A. agrestis than without earthworms. Mean (standard deviation) phenoloxidase activities in the presence of A. agrestis were 0.15 (±0.10), 1.14 (±0.46), 2.71 (±0.98) μmol g⁻¹ h⁻¹ for pine, spruce and cedar respectively, and 0.012 (±0.023), 0.25 (±0.25), 0.78 (±0.45) μmol g⁻¹ h⁻¹ in the absence of A. agrestis. There was significantly greater peroxidase activity for the pine and spruce treatment when earthworms were present. Mean peroxidase activities were 0.47 (±0.21), 0.94 (±0.29), 1.20 (±0.77) μmol h⁻¹ g⁻¹ soil for pine, spruce and cedar, respectively for soils with A. agrestis and 0.15 (±0.10), 0.37 (±0.10), 0.63 (±0.30) μmol h⁻¹ g⁻¹ soil in the absence of earthworms. The increased ligninolytic activity in combination with successful maturation of juveniles into adult A. agrestis suggests that mulch can be habitat for these invasive earthworms. This finding is supported by a survey of master gardeners in Vermont and New Hampshire 20% of whom reported to have seen these earthworms mainly in their gardens and mulched beds.