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.     

Herbivory of an invasive slug in a model grassland community can be affected by earthworms and mycorrhizal fungi

Invasion of non-native species is among the top threats for the biodiversity and functioning of native and agricultural ecosystems worldwide. We investigated whether the herbivory of the slug Arion vulgaris (formerly Arion lusitanicus; Gastropoda), that is listed among the 100 worst alien species in Europe, is affected by soil organisms commonly present in terrestrial ecosystems (i.e. earthworms—Annelida: Lumbricidae and arbuscular mycorrhizal fungi—AMF, Glomerales). We hypothesized that slug herbivory would be affected by soil organisms via altered plant nutrient availability and plant quality. In a greenhouse experiment, we created a simple plant community consisting of a grass, a forb, and a legume species and inoculated these systems with either two earthworm species and/or four AMF taxa. Slugs were introduced after plants were established. Earthworms significantly reduced total slug herbivory in AMF-inoculated plant communities (P?=?0.013). Across plant species, earthworms increased leaf total N and secondary metabolites, AMF decreased leaf thickness. Mycorrhizae induced a shift in slug feeding preference from non-legumes to legumes; the grass was generally avoided by slugs. AMF effects on legume herbivory can partly be explained by the AMF-induced increase in total N and decrease in C/N ratio; earthworm effects are less clear as no worm-induced alterations of legume plant chemistry were observed. The presence of earthworms increased average AMF colonization of plant roots by 140 % (P?<?0.001). Total shoot mass was significantly increased by AMF (P?<?0.001). These data suggest that the feeding behavior of this invasive slug is altered by a belowground control of plant chemical quality and community structure.     

Plant community impacts on the structure of earthworm communities depend on season and change with time

Declining plant diversity potentially threatens essential ecosystem functions driven by the decomposer community, such as litter decomposition and nutrient cycling. Currently, there is no consensus on the interrelationships between plant diversity and decomposer performance and previous studies highlighted the urgent need for long-term experiments.In the Jena Experiment we investigated the long-term impacts of plant community characteristics on the structure of earthworm communities representing key decomposers in temperate grassland. We repeatedly sampled plots varying in plant species richness (1-16 species), plant functional group richness (1-4 groups), and presence of certain plant functional groups (grasses and legumes) three, four, and six years after establishment of the experiment in spring and autumn.The results show that earthworm performance is essentially driven by the presence of certain plant functional groups via a variety of mechanisms. Plant productivity (root biomass) explained most of the detrimental grass impacts (decrease in earthworm performance), while beneficial legume effects likely were linked to high quality inputs of plant residues (increase in earthworm performance). These impacts depended on the functional group of earthworms with the strongest effects on surface feeding anecic earthworms and minor effects on soil feeding endogeic species. Remarkably, effects of plant community characteristics on the composition and age structure of earthworm communities varied between seasons. Moreover, plant diversity effects reported by a former study decreased and detrimental effects of grasses increased with time.The results indicate that plant community characteristics, such as declining diversity, indeed affect the structure of earthworm communities; however, loss of key plant functional groups is likely to be more important than plant species number per se. However, in frequently disturbed ecosystems plant species richness might be important for the recovery and resilience of belowground functions. Moreover, the results accentuate the importance of long-term repeated measurements to fully appreciate the impacts of plant community composition and diversity on ecosystem properties. Single point observations may be misleading and potentially mask the complexity of above-belowground interrelationships.     

蚯蚓与菌根提高玉米生长和氮磷吸收的互补效应

【目的】蚯蚓和丛枝菌根真菌处于不同的营养级,但在促进植物生长和提高土壤肥力等方面却都发挥着积极作用。研究蚯蚓菌根互作及其对玉米吸收土壤中的氮、磷养分的影响,可为提升土壤生物肥力及促进农业的可持续发展提供理论依据。【方法】本研究采用田间盆栽方式,以玉米为供试作物,研究蚯蚓(Eisenia fetida)与丛枝菌根真菌(Glomus intraradices)互作及其对玉米养分吸收的影响。试验设置P 25和175 mg/kg两个水平。每个磷水平进行接种与不接种菌根真菌以及添加与不添加蚯蚓,共8个处理。调查了玉米生长、养分吸收以及真菌浸染和土壤养分的有效性。【结果】两个磷水平下,蚯蚓和菌根在增加玉米地上部和根系生物量方面有显著正交互作用(P0.05)。接种菌根真菌的各处理显著增加了玉米的侵染率及泡囊丰度、根内菌丝丰度等菌根指标。同时添加蚯蚓和接种菌根真菌的处理(AM+E)显著提高了菌根的侵染率、菌丝密度、丛枝丰度和根内菌丝丰度但是泡囊丰度有所下降。两种磷水平下,AM+E处理玉米地上部和地下部含氮量和含磷量均显著高于其他三个处理。在低磷条件下,地上部氮磷总量的增加分别是添加蚯蚓和接菌的作用;而地下部磷总量的增加主要是菌根真菌的作用。在高磷条件下,单加蚯蚓显著增加玉米氮磷的总量,而接种菌根真菌对玉米氮磷吸收的影响未达显著性水平。在高磷条件下,单加蚯蚓的处理显著提高玉米地上地下部生物量(P0.05),而单接菌的处理效应不显著,蚯蚓菌根互作通过提高土壤微生物量碳、氮实现对玉米生长和养分吸收的调控。在低磷条件下,单接菌显著提高了玉米的生物量(P0.05),单加蚯蚓的处理具有增加玉米生物量的趋势。菌根真菌主要促进玉米对磷的吸收,蚯蚓主要矿化秸秆和土壤中的氮磷养分增加土壤养分的有效性,蚯蚓菌根互作促进了玉米根系对土壤养分的吸收并形成氮磷互补效应。【结论】无论在高磷还是低磷水平下,蚯蚓菌根相互作用都提高了玉米地上地下部生物量、氮磷吸收量同时提高了土壤微生物量碳、氮。蚯蚓菌根相互作用对植物生长的影响取决于土壤养分条件。在高磷条件下(氮相对不足),蚯蚓菌根互作通过调控土壤微生物量碳、氮调控玉米生长和养分吸收。低磷条件下,菌根主要发挥解磷作用,蚯蚓主要矿化秸秆和土壤中的氮素,蚯蚓和菌根互补调控土壤中氮、磷,从而促进植物的生长和养分吸收。     

Decomposer community complexity affects plant competition in a model early successional grassland community

The effects of a simple decomposer community, consisting of collembola, enchytraeids and earthworms on the performance of plants in a model Mediterranean early successional grassland community were investigated. Interactions of plant functional groups and species within the soil decomposer community resulted in significant shifts of plant competitive strength and therefore are likely to affect the rate of the successional development of plant communities.Depending on the functional role of the decomposer taxa involved, either legumes, or non-leguminous forbs were more negatively affected. Therefore, increasing species numbers with in the decomposer community gradually reduced the relative contribution of forbs (legumes and non-leguminous forbs), although fundamentally different mechanisms were responsible for these effects. Surprisingly, not the biomass-dominant annelid taxa, but collembola had the strongest effects on plant performance. The grass-to-forb ratio, an indicator of successional change, shifted from 3 to 4 in presence of collembola, suggesting that indirect effects on microbial symbionts of plants were more important than classic decomposer effects via increased and more constant nutrient availability to plants. Total plant productivity, however, was not affected, since grasses gained competitive advantage and compensated with increased growth for the negative animal effects on forb performance. Our results highlight the importance of specific functional groups among decomposers for structuring grassland plant communities. However, great plasticity in the plant community through negative covariance between forbs and grasses, partly compensated for decomposer effects at the plant canopy level.     

Earthworms can modify effects of hydrochar on growth of Plantago lanceolata and performance of arbuscular mycorrhizal fungi

Hydrothermal carbonization (HTC) is a method to produce carbonized material at relatively low temperatures (180–250 °C) under pressure and aqueous conditions. The product is called hydrochar and can be used as a soil amendment. However, applied in high dosages it may have detrimental effects on plants or soil biota. The potential impact of hydrochar amendment on beneficial soil organisms such as arbuscular mycorrhizal fungi (AMF) and earthworms and their interactions are not well understood. The goal of the present study was to determine effects of hydrochar on plant growth and soil biota and to evaluate interactions of earthworms and hydrochar on plant and AMF performance and to identify underlying mechanisms. In a greenhouse experiment, we investigated the effect of hydrochar at different addition rates (control, 1% and 10%, v/v) with or without the earthworm Aporrectodea caliginosa on the growth of Plantago lanceolata L. and the performance of its AMF. We observed a positive interaction between earthworms and 10% hydrochar on shoot and root biomass: added as a single treatment hydrochar had a negative effect on plant growth at this dosage, but plant biomass increased significantly when hydrochar was added together with earthworms. Root colonization by AMF increased significantly with increasing concentration of hydrochar, but was not affected by earthworms. Contrastingly, extraradical hyphal length of AMF was reduced by earthworms, but not affected by hydrochar. Thus, hydrochar and earthworms affected the performance of AMF, albeit of different AMF structures and in different directions. Our results indicate that earthworms may play an important role in alleviating the negative impacts of high dosages of hydrochar on plant growth; such interactions should move into focus of future research on potential effects of HTC materials.     

Changes in nutrient pools,microbial biomass and microbial activity in soils after transit through the gut of three endogeic earthworm species of the genus Postandrilus Qui and Bouché, 1998

Purpose

Endogeic earthworms play a significant role in biogeochemical cycles due to the large amount of soil they ingest, and because after transit through their guts, casts usually show differences in nutrient contents and microbial populations with bulk soil. Here, we studied how three endogeic earthworm species, Postandrilus majorcanus, Postandrilus sapkarevi and Postandrilus palmensis, inhabiting soils in Majorca island (Balearic Islands, W Mediterranean), modify nutrient pools and microbial communities of soil.

Materials and methods

To do this, we analysed C, N and P pools, microbial biomass (phospholipid fatty acids, PLFA) and microbial activity (fluorescein diacetate hydrolysis, FDA) in paired samples of bulk soil and fresh casts.

Results and discussion

The mineral and organic N contents were generally enhanced in casts produced by all three earthworm species. However, inorganic P and organic C contents were only higher in P. sapkarevi (32 %, only P) and P. majorcanus casts (100 % for both soil nutrient pools) than in bulk soil. Bacterial and fungal biomass were only higher than in bulk soil in P. majorcanus casts (65 and 100 %, respectively), but without effects on microbial activity, that was lower in P. palmensis casts (26 %). Earthworm gut transit strongly influenced the soil microbial community structure, resulting in differences between casts and soils.

Conclusions

The increased nutrient mineralization (6-, 1.3- and 1.4-fold for N, C and P, respectively) in casts produced by these earthworm species is of particular importance because of the amount of casts released and the seasonal variations in earthworm activity, which may favour plant growth.     

Additive and interactive effects of functionally dissimilar soil organisms on a grassland plant community

The productivity and diversity of plant communities are affected by soil organisms such as arbuscular mycorrhizal fungi (AMF), root herbivores and decomposers. However, it is unknown how interactions between such functionally dissimilar soil organisms affect plant communities and whether the combined effects are additive or interactive. In a greenhouse experiment we investigated the individual and combined effects of AMF (five Glomus species), root herbivores (wireworms and nematodes) and decomposers (collembolans and enchytraeids) on the productivity and nutrient content of a model grassland plant community as well as on soil microbial biomass and community structure. The effects of the soil organisms on productivity (total plant biomass), total root biomass, grass and forb biomass, and nutrient uptake of the plant community were additive. AMF decreased, decomposers increased and root herbivores had no effect on productivity, but in combination the additive effects canceled each other out. AMF reduced total root biomass by 18%, but decomposers increased it by 25%, leading to no net effect on total root biomass in the combined treatments. Total shoot biomass was reduced by 14% by root herbivores and affected by an interaction between AMF and decomposers where decomposers had a positive impact on shoot growth only in presence of AMF. AMF increased the shoot biomass of forbs, but reduced the shoot biomass of grasses, while root herbivores only reduced the shoot biomass of grasses. Interactive effects of the soil organisms were detected on the shoot biomasses of Lotus corniculatus, Plantago lanceolata, and Agrostis capillaris. The C/N ratio of the plant community was affected by AMF.In soil, AMF promoted abundances of bacterial, actinomycete, saprophytic and AMF fatty acid markers. Decomposers alone decreased bacterial and actinomycete fatty acids abundances but when decomposers were interacting with herbivores those abundances were increased. Our results suggests that at higher resolutions, i.e. on the levels of individual plant species and the microbial community, interactive effects are common but do not affect the overall productivity and nutrient uptake of a grassland plant community, which is mainly affected by additive effects of functionally dissimilar soil organisms.     

Effects of epigeic earthworm (Eisenia fetida) and arbuscular mycorrhizal fungus (Glomus intraradices) on enzyme activities of a sterilized soil–sand mixture and nutrient uptake by maize

A pot experiment was conducted to investigate the effect of epigeic earthworm (Eisenia fetida) and arbuscular mycorrhizal (AM) fungi (Glomus intraradices) on soil enzyme activities and nutrient uptake by maize, which was grown on a mixture of sterilized soil and sand. Maize plants were grown in pots inoculated or not inoculated with AMF, treated or not treated with earthworms. Wheat straw was added as a feed source for earthworms. Mycorrhizal colonization of maize was markedly increased in AM fungi inoculated pots and further increased by addition of epigeic earthworms. AM fungi and epigeic earthworms increased maize shoot and root biomass, respectively. Soil acid phosphatase activity was increased by both earthworms and mycorrhiza, while urease and cellulase activities were only affected by earthworms. Inoculation with AM fungi significantly (p?<?0.001) increased the activity of soil acid phosphatase but decreased soil available phosphorus (P) and potassium (K) concentrations at harvest. Addition of earthworms alone significantly (p?<?0.05) increased soil ammonium-N content, but decreased soil available P and K contents. AM fungi increased maize shoot weight and root P content, while earthworms improved N, P, and K contents in shoots. AM fungi and earthworm interactively increased maize shoot and root biomass through their regulation of soil enzyme activities and on the content of available soil N, P, and K.     

Quantifying surface and subsurface cast production by earthworms under controlled laboratory conditions

Earthworm casts, formed when organic substrates and soil minerals pass through the digestive tract, may protect soil organic matter from biological degradation if they persist in the soil. Yet, the stability of casts is affected by their location in the soil profile because surface casts are exposed to more disruptive forces (wetting-drying, freezing-thawing) than subsurface casts. It is not known whether environmental conditions affect the proportions of surface and subsurface casts produced by earthworms. This study investigated how surface and subsurface cast production by juveniles of Aporrectodea spp. and Lumbricus spp. was affected by temperature. Two juveniles of Aporrectodea spp. or Lumbricus spp. were added to plexiglass chambers filled with soil, and five replicate chambers were incubated in the dark at 5°C, 10°C, 15°C or 20°C for 1 week. Most of the casts produced by Aporrectodea spp. and Lumbricus spp. were surface casts, with <10% of casts deposited below the soil surface. The earthworms studied produced more casts, and a greater proportion of surface casts, as the soil temperature increased. These results can be used to estimate the quantity of surface and subsurface casts produced by earthworm populations under field conditions and determine the susceptibility of cast-associated organic matter to decomposition in the medium- to long-term.     

Distribution of radioactive cesium in soil and its uptake by herbaceous plants in temperate pastures with different management after the Fukushima Dai-Ichi Nuclear Power Station accident

Abstract

The accident at Fukushima Dai-Ichi Nuclear Power Station (NPS) extensively contaminated the agricultural land in the Tohoku region of Japan with radioactive cesium [sum of cesium-134 (¹³⁴Cs) and cesium-137 (¹³⁷Cs)]. We evaluated the status of radioactive cesium (Cs) contamination in soil and plants at the Field Science Center of Tohoku University, northern Miyagi prefecture, 150 km north of the NPS. In seven pastures with different management, we examined: (1) the distribution of radioactive Cs in soil, (2) the concentration of radioactive Cs in various herbaceous plant species and (3) the change in radioactive Cs content of plants as they matured. We collected samples of litter, root mat layer (root mat soil and plant roots), and subsurface soil (0–5 cm beneath the root mat) at two to three locations in each pasture in December 2011 and May 2012. The aboveground parts of herbaceous plants (four grasses, two legumes, and one forb species) were collected from May 9 to June 20, 2012, at 14-d intervals, from one to five fixed sampling locations in each pasture. The distribution of radioactive Cs in soil differed among pastures to some degree: a large proportion of radioactive Cs was distributed in the root mat layer. Pasture management greatly influenced the radioactive Cs content of herbaceous plants (p < 0.001); plant species had less influence. Radioactive Cs content was highest (> 3 kBq kg^?1 dry weight) on May 9 and significantly decreased with maturity (p < 0.001) for most of the pastures, whereas it remained low (0.04–0.18 kBq kg^?1 dry weight) throughout the measurement period in the pasture where composted cattle manure was applied. The soil-to-plant transfer factor was negatively correlated to pH(H₂O) (R² = 0.783, p < 0.001) and exchangeable K content (R² = 0.971, p < 0.001) of root mat soils, which suggests that surface application of composted cattle manure reduces plant uptake of radioactive Cs by increasing the exchangeable K content of the soil. The radioactive Cs content of plants decreased with plant maturity; its degree of decrease (May 9 to June 6) was smaller in legumes (80.6%) than grasses (55.5%) and the forb (58.6%). Radioactive Cs content decreased with plant maturity; also, the proportion remaining in the aboveground plant was higher in legumes (80.6%) than grasses (55.5%) and the forb (58.6%).     

The earthworm (Aporrectodea caliginosa) primes the release of mobile and available micronutrients in soil

The objectives of our study were to quantify the impact of endogeic earthworms Aporrectodea caliginosa (Savigny) on iron (Fe), manganese (Mn) and zinc (Zn) mobility and availability in soil. Dried rye straw (Cecale cereale L.), clover aboveground parts (Trifolium pratense L.) or calcium carbonate were added to determine the effects on soil micronutrient mobility. To test the importance of soil–water saturation mediated by earthworms, soil samples were modified to 60% (control) and 100% (as in casts) water holding capacity (WHC). To assess availability of micronutrients, a cucumber plant (Cucumis sativus L.) bioassay were used. Earthworm casts had generally higher amounts of water-soluble micronutrients compared with bulk soils regardless of their moisture contents. The increased micronutrient mobility was more pronounced in casts from soil samples amended with plant residues (especially with straw) and was significantly higher than mobility in control soil for at least 1 week after the casts were deposited. Pre-incubation of soils amended with clover or straw with living earthworms for 4 weeks produced an increase in both shoot biomass and translocation rate of micronutrients (Mn, Zn) into xylem sap of cucumber compared to soils not worked by earthworms. The earthworm-mediated plant performances were determined 4 weeks after the earthworms were removed. The results demonstrated that earthworms can significantly impact the formation of mobile and available micronutrients in a soil. The relationship between micronutrient availability to cucumber plants and earthworm contribution to nitrogen (N) mineralization and micronutrient mobility are discussed.     

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.     

Inoculating maize fields with earthworms (Aporrectodea trapezoides) and an arbuscular mycorrhizal fungus (Rhizophagus intraradices) improves mycorrhizal community structure and increases plant nutrient uptake

Earthworms and arbuscular mycorrhizal fungi (AMF) are important macrofauna and microorganisms of the rhizosphere. The effect of the inoculation of soil with earthworms (Aporrectodea trapezoides) and mycorrhiza (Rhizophagus intraradices) on the community structure of mycorrhizal fungi and plant nutrient uptake was determined with split plots in a maize field. Maize plants were inoculated or not inoculated with AMF, each treated with or without earthworms. Wheat straw was added as a feed source for earthworms. Inoculating AMF significantly increased maize yield (p?<?0.05), and the yield was further enhanced by the addition of earthworms. Alkaline phosphomonoesterase activities, soil microbial biomass carbon (SMBC) and nitrogen (SMBN) increased with the addition of both earthworms and AMF. Soil inorganic N and available K were positively affected by earthworms, while available P showed a negative relationship with AMF. Treatment with both AMF and earthworms increased shoot and root biomass as well as their N and P uptake by affecting soil phosphomonoesterase and urease activities, SMBC, SMBN, and the content of available nutrients in soil. The applied fungal inoculants were successfully traced by polymerase chain reaction with novel primers (AML1 and AML2) which target the small subunit rRNA gene. The amplicons were classified by restriction fragment length polymorphism and sequencing. Moreover, field inoculation with inocula of non-native isolates of R. intraradices appeared to have stimulated root colonization and yield of maize. Adding earthworms might influence native AMF community, and the corresponding abundance increased after earthworms were inoculated, which has positive effects on maize growth.     

Earthworm casts form stable aggregates in physically degraded soils

Topsoils affected by surface mining suffer severe physical degradation and lose most of their earthworm populations. After mining, replaced soils are planted to grassland and managed to improve soil structure. Earthworm inoculation into selected restored areas produced populations similar to those of undisturbed soils within 3 years. Soil properties in inoculated areas were compared with those of controls to evaluate the contribution of casts to bulk soil aggregation, and soil organic matter and root content responses to earthworm activity. Crumb porosity and coarse particle content were measured in water-stable macro-aggregates and earthworm casts to establish whether aggregates were formed by earthworms. Over a 5- to 6-year period, inoculation increased stable aggregation (>2 μm, >60 μm and >3 mm), even at 0- to 5-cm depth where it reduced soil organic matter content. Productivity and root content were also increased by inoculation; roots and organic matter were re-distributed to greater depth. Crumb porosity decreased with casts > aggregates (inoculated plots) > aggregates (control plots). Coarse particle content increased with casts < aggregates (inoculated plots) < aggregates (control plots). Coarse particle and porosity data were consistent with much of the newly aggregated soil being processed and formed by earthworms as casts. Whilst levels of soil organic matter were often closely associated with percentage stable aggregation, root content showed weaker associations. Aggregation percentage was most closely associated with abundance of Aporrectodea longa, although at particular depths significant correlations were also obtained for Aporrectodea caliginosa and Lumbricus terrestris. Results suggest that earthworms, rather than plant roots, initiate aggregation in severely degraded grassland soils.     

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.     

Adaptive attributes of tropical forage species to acid soils I. Differences in plant growth,nutrient acquisition and nutrient utilization among C4 grasses and C3 legumes

Low supply of nutrients is a major limitation of forage adaptation and production in acid soils of the tropics. A glasshouse study was conducted to find differences in plant growth, nutrient acquisition and use, among species of tropical forage grasses (with C₄ pathway of photosynthesis) and legumes (with C₃), when grown in two acid soils of contrasting texture and fertility. Twelve tropical forage legumes and seven tropical forage grasses were grown in sandy loam and clay loam Oxisols at low and high levels of soil fertility. After 83 days of growth, dry matter distribution among plant leaves, stems, and roots, leaf area production, shoot and root nutrient composition, shoot nutrient uptake, and nutrient use efficiency were measured. Soil type and fertility affected biomass production and dry matter partitioning between roots and shoots. The allocation of dry matter to root production was greater with low soil fertility, particularly in sandy loam. The grasses responded more than the legumes to increased soil fertility in both shoot and root biomass production. Leaf area production and the use of leaf biomass for leaf expansion (specific leaf area) were greater in legumes than in grasses, irrespective of soil type and fertility. But soil type affected shoot biomass production and nutrient uptake of the grasses more than those of the legumes. There were significant interspecific differences in terms of shoot nutrient uptake. The grasses were more efficient than legumes in nutrient use (grams of shoot biomass produced per gram of total nutrient uptake) particularly for nitrogen (N) and calcium (Ca).     

What are the limits of the drilosphere? An incubation experiment using Metaphire posthuma

The near infrared reflectance spectroscopy (NIRS) method was used in the present study to compare earthworm-made soil aggregates to aggregates found in the surrounding bulk soil. After initially assessing the daily cast production of Metaphire posthuma, boxes with soil incubated with M. posthuma and control soils were subjected to wetting in order to reorganize the soil structure. After two months of incubation, soil aggregates produced by earthworms (casts and burrows), soil aggregates that were appeared to be unaffected by earthworms (bulk soil without visible trace of earthworm bioturbation from the earthworm treatment) and soil aggregates that were entirely unaffected by earthworms (control – no earthworm – treatment) were sampled and their chemical signatures analyzed by NIRS. The production of below-ground and surface casts reached 14.9 g soil g worm^?1 d^?1 and 1.4 g soil g worm^?1 d^?1, respectively. Soil aggregates from the control soils had a significantly different NIRS signature from those sampled from boxes with earthworms. However, within the earthworm incubation boxes the NIRS signature was similar between cast and burrow aggregates and soil aggregates from the surrounding bulk soil. We conclude that the high cast production by M. posthuma and the regular reorganization of the soil structure by water flow in and through the soil lead to a relatively homogenous soil structure. Given these results, we question the relevance of considering the bulk soil that has no visible activity of earthworm activity as a control to determine the effect of earthworms on soil functioning.     

蚯蚓对土壤温室气体排放的影响及机制研究进展

土壤是温室气体的重要源和汇。蚯蚓是土壤物质循环的重要参与者,能够直接或间接影响土壤CO2、N2O和CH4等温室气体的产生和释放。蚯蚓呼吸产生的CO2,是土壤呼吸的重要组成部分;蚯蚓自身肠道、分泌液、消化物和排泄物等微环境促进反硝化过程释放N2O。蚯蚓还通过取食、掘穴、排泄等活动,改变土壤理化性质、微生物组成和活性及其他土壤动物的组成,影响地上植物生长,调节土壤分解、矿化、硝化、反硝化和甲烷生成及氧化等生态过程,间接影响土壤温室气体的排放。蚯蚓对土壤温室气体排放的影响逐渐受到重视,但目前研究仍以室内培养和单因子环境条件的模拟为主,缺少野外原位实验和多环境因子的交互实验研究。长期监测和同位素示踪技术,是深入探讨蚯蚓影响温室气体排放机制的重要手段。温室气体类型上,CO2和N2O是研究热点,CH4研究比较罕见。未来研究,应重视不同生态类群蚯蚓与土壤理化特征、微生物组成、其他类群土壤动物和地上植物间的交互作用,加强机制研究,并关注土壤污染环境下蚯蚓功能性状的变化;综合评价蚯蚓对土壤温室气体排放和土壤碳氮固定的影响,科学评估蚯蚓活动对土壤碳氮释放的促进或减缓作用。     

Stable isotope labelling of earthworms can help deciphering belowground–aboveground interactions involving earthworms,mycorrhizal fungi,plants and aphids

Functional relationships between belowground detritivores and/or symbionts and aboveground primary producers and their herbivores are not well studied. In a factorial greenhouse experiment we studied interactions between earthworms (addition/no addition of Lumbricus terrestris; Clitellata: Lumbricidae) and arbuscular-mycorrhizal fungi (AMF; with/without inoculation of Glomus mosseae; Glomerales: Glomeraceae) on the leguminous herb Trifolium repens (Fabales: Fabaceae) and associated plant aphids (Aphis gossypii, A. craccivora; Hemiptera: Aphidoidea). In order to be able to trace organismic interactions, earthworms were dual-labelled with stable isotopes (¹⁵N-ammonium nitrate and ¹³C-glucose). We specifically wanted to investigate whether (i) isotopic signals can be traced from the labelled earthworms via surface castings, plant roots and leaves to plant aphids and (ii) these compartments differ in their incorporation of stable isotopes. Our results show that the tested organismic compartments differed significantly in their ¹⁵N isotope enrichments measured seven days after the introduction of earthworms. ¹⁵N isotope incorporation was highest in casts followed by earthworm tissue, roots and leaves, with lowest ¹⁵N signature in aphids. The ¹³C signal in roots, leaves and aphids was similar across all treatments and is for this reason not recommendable for tracing short-term interactions over multitrophic levels. AMF symbiosis affected stable isotope incorporation differently in different subsystems: the ¹⁵N isotope signature was higher below ground (in roots) but lower above ground (leaves and aphids) in AMF-inoculated mesocosms compared to AMF-free mesocosms (significant subsystem × AMF interaction). Aphid infestation was unaffected by AMF and/or earthworms. Generally, these results demonstrate that plants utilize nutrients excreted by earthworms and incorporate these nutrients into their roots, leaf tissue and phloem sap from where aphids suck. Hence, these results show that earthworms and plant aphids are functionally interlinked. Further, ¹⁵N-labelling earthworms may represent a promising tool to investigate nutrient uptake by plants and consequences for belowground-aboveground multitrophic interactions.