Food preferences of earthworms for soil fungi

Soil fungi are considered to be an important food source for earthworms. Selection experiments were carried out in order to study the preferences of earthworm species for a variety of soil fungi. Nine fungal species (Cladosporium cladosporioides, Rhizoctonia solani, Mucor sp., Trichoderma viride, Fusarium nivale, Phlebia radiata, Glaeophyllum trabeum, Coniophora puteana, Coriolus versicolor) were grown separately in centrifuge tubes on sterilized sand with potato dextrose. Tubes containing different fungal species, 8–9 per experiment, were arranged in a food choice arena. The preference for the fungi of 5 different earthworm species (Lumbricus terrestris, Lumbricus castaneus, Aporrectodea caliginosa, Aporrectodea rosea, Octolasion cyaneum) was tested by adding one specimen per chamber. Removal of sand from the tubes within 6 days was used as the indicator of preference by earthworms. The food preference of earthworms irrespective of ecological group followed a general pattern. F. nivale and C. cladosporioides were the preferred fungal species, followed by fast-growing species such as Mucor sp. and R. solani. In contrast, basidiomycetes were generally refused. The epigeic species L. rubellus had the strongest preference for a single fungal species, in contrast the endogeic species A. rosea fed more evenly on different fungal species. We conclude that early successional fungal species are used as cues by earthworms to detect fresh organic resources in soil.     

Reproduction and growth of three deep-burrowing earthworms (Lumbricidae) in laboratory culture in order to assess production for soil restoration

Laboratory experiments were conducted to examine the growth and reproduction of three deep-burrowing lumbricids, Aporrectodea longa, Lumbricus terrestris, and Octolasion cyaneum. The reproductive output was recorded as 18.8, 38.0, and 32.3 cocoons per worm per year for A. longa, L. terrestris, and O. cyaneum, respectively. For the same species, maturity was reached at a mean mass of 3.9, 5.0 and 2.4 g, within 3 months from the hatchling stage by L. terrestris and within 4 months by the other two species. The hatching success of cocoons at 15 and 20°C was within the range of 70–80% for each species, except A. longa at the higher temperature, where a viability of 47% was recorded. Twenty percent of viable O. cyaneum cocoons produced twin hatchlings, compared with only one percent for A. longa and L. terrestris. A combination of these results suggests that a complete life-cycle for each species could be achieved within 6 months (L. terrestris and A. longa) or 7–8 months (O. cyaneum). Each species has particular life-cycle strategies that would aid survival and colonisation, under field conditions, if inoculated into restored soils.     

Culture of commercially obtained Lumbricus terrestris L.: Implications for sub-lethal ecotoxicological testing

The use of commercially purchased or field-collected earthworms of unknown age, exposure or pre-treatment in sub-lethal ecotoxicological studies is questionable. In this study, adult (clitellate) Lumbricus terrestris, obtained from 5 commercial suppliers in the UK and also field collected, were kept under controlled environmental conditions (15 °C and 24 h darkness) in a sterilised loam soil and fed horse manure. Survival, biomass and cocoon production was monitored every 4 weeks over 1 y. Marked differences were recorded in survival rates (ranging from 40-100% after 40 weeks) and cocoon production (ranging from 15.1 to 32.2 worm⁻¹ y⁻¹) between treatments. Biomass in all treatments (mean mass 4.32-5.61 g at the outset) increased with time to week 20 (maximum 6.7 g) and then declined steadily (3.23-4.7 g at week 52). This pattern was also observed in cocoon production and was considered to be a function of an initial period of acclimation (0-12 weeks) followed by a period of high production (12-36 weeks) under optimal conditions and then fatigue (36-52 weeks) caused by reproductive exhaustion. Results suggest that earthworm origin may influence the validity and reproducibility of sub-lethal ecotoxicological studies and where applicable laboratory-reared earthworms of known age and history are recommended as test subjects.     

Influence of soil ingestion by earthworms on the availability of potassium in soil: An incubation experiment

Summary An incubation experiment was conducted to study the changes that occur in the K status of soil due to earthworm activity. Samples of Tokomaru silt loam soil were inoculated with the common pasture earthworm species Aporrectodea caliginosa and incubated for 21 days. Aliquots of moist soil were analyzed for exchangeable K by leaching with neutral molar ammonium acetate at 1:50 soil solution ratio. Extraction with boiling 1 M nitric acid at 1:100 soil solution ratio for 20 min was used to determine available non-exchangeable K. The results indicated that the exchangeable K content increased significantly due to earthworm activity but nitric acid-extractable K did not change significantly. It is inferred that earthworms increase the availability of K by shifting the equilibrium among the forms of K from relatively un-available forms to more available forms in the soil chosen for the study.     

Critical body residues (CBRs) for ecotoxicological soil quality assessment: copper in earthworms

The term ‘critical body residue’ (CBR) was defined as the lowest observed total body concentration of a contaminant in an organism, which is associated with the occurrence of adverse toxic effects in either individuals or populations of a defined age or stage of development. In this study, internal toxicity thresholds were determined for copper in the clitellated adult stage of earthworms (Lumbricus rubellus and Aporrectodea caliginosa). The objective was to assess the applicability of CBRs as a practical tool in soil quality assessment of contaminated sites and as a means of a sustainable protection of earthworm fauna. Laboratory studies showed that body concentrations of Cu were generally in agreement with the chemically available CaCl₂-extractable fraction in soil, but that there was also some evidence of internal pH-related homeostatic regulation. Toxicological correlates of body Cu concentrations with adverse effects on cocoon production (fecundity) suggested an approximate sublethal internal threshold of about 40 mg kg⁻¹, with mortality occurring at about 60 mg kg⁻¹. Adult L. rubellus sampled from areas with a wide range of metal pollution showed body Cu concentrations with a minimum of 8 mg kg⁻¹ and a maximum of 60 mg kg⁻¹. Beyond this apparent physiological tolerance range, environmental management directed at optimal earthworm population survival may not be sustainable in contaminated fields. Studies of L. rubellus colonizing a metal-contaminated experimental sludge-treated field showed that a reduced rate of colonization can already be associated with an average body Cu concentration of 25 mg kg⁻¹. However, in this particular field situation mixture effects of other metals that were also present in the soil and the occurrence of avoidance behaviour during colonization may have contributed to this low internal toxicity threshold. It is concluded that the CBR approach seems to be a feasible option for use as a tool in a bioavailability-based soil quality assessment, even for essential trace metals like copper, but that further insight may be needed to establish the uncertainty and reliability of the application in environmental quality assessment and decision making.     

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

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

A review of visual soil evaluation techniques for soil structure

Soil structure forms a key component of soil quality, and its assessment by semi‐quantitative visual soil evaluation (VSE) techniques can help scientists, advisors and farmers make decisions regarding sampling and soil management. VSE techniques require inexpensive equipment and generate immediate results that correlate well with quantitative measurements of physical and biochemical properties, highlighting their potential utility. We reviewed published VSE techniques and found that soils of certain textures present problems and a lack of research into the influence of soil moisture content on VSE criteria. Generally, profile methods evaluate process interactions at specific locations within a field, exploring both intrinsic aspects and anthropogenic impacts. Spade methods focus on anthropogenic characteristics, providing rapid synopses of soil structure over wider areas. Despite a focus on structural form, some methods include criteria related to stability and resiliency. Further work is needed to improve existing methods regarding texture influences, on‐farm sampling procedures and more holistic assessments of soil structure.     

Landscape-scale species richness of earthworms in the Porongurup Range,Western Australia: influence of aspect,soil fertility,and vegetation type

Indigenous Australian earthworms comprise a patchily collected taxon that is likely to have high diversity and pose difficulties for systems of biodiversity prediction or monitoring that use surrogates. This is because surrogates for earthworm diversity have scarcely been identified at landscape scales in Australian contexts. In the present paper we examine the diversity of earthworms in a 26 km² remnant of native vegetation. Comparison of diversity within the Porongurup Range to diversity of other earthworm faunas demonstrates the sampling intensities required to adequately sample regions of 10⁰–10¹ km². In addition diversity patterns were examined in relation to habitat as described by landscape context, vegetation cover and soil characteristics. Species accumulation in samples from the northern aspect of the Porongurup Range was less than expected from a random distribution of species. Combined Karri-, and Marri- and Karri-vegetated sites supported significantly more earthworm species on the southern aspect of the Porongurup Range than expected from accumulation in randomly ordered samples. Sites carrying Jarrah as the only dominant canopy had fewer earthworm species than would be expected from accumulation in randomly ordered samples. Jarrah overstorey sites and southern aspect Karri sites correspond to extremes in a continuum of soil and landscape characteristics. Earthworm responses to landscape and vegetation units in the region warrant further investigation.     

Effects of earthworms on decomposition and metal availability in contaminated soil: Microcosm studies of populations with different exposure histories

Population-specific differences in the responses of earthworms to simultaneous exposure to Cu and Zn were studied in microcosm experiments. Two populations of Aporrectodea caliginosa tuberculata (Eisen) with different metal exposure histories were chosen for the studies. Microcosms were prepared containing either uncontaminated soil or soils with low or high combined Cu/Zn -concentrations (79/139 or 178/311 mg kg⁻¹ dry mass of soil, respectively). Earthworms from each population were introduced to the microcosm treatments with some microcosms serving as controls without earthworms. One series of microcosms was destructively sampled after 16 weeks incubation in a climate chamber. Survival, growth, reproduction and decomposition by earthworms in each treatment were measured. An additional microcosm series was sampled for soil and earthworm measurements at four weeks intervals to determine temporal changes in the availability of metals in the soils and their accumulation into earthworms. Cu and Zn were sequentially extracted from the soil samples of both microcosm series to estimate mobility and availability of the metals in the soil. Earthworms with long-term exposure history to metal-contaminated soil seemed to tolerate higher soil metal concentrations than earthworms without earlier exposure. Both earthworms and metals affected soil respiration (CO₂ production) and nitrogen mineralization. In addition, earthworms seemed to decrease the mobility and bioavailability of metals in the soil through their burrowing activity.     

Potential of earthworms to accelerate removal of organic contaminants from soil: A review

Earthworms can accelerate the removal of contaminants from soil. Earthworms change the physical and chemical properties of soil by mixing it with organic material and through their burrowing they improve aeration and render contaminants available for microorganisms. The presence of earthworms in contaminated soil indicate that they can survive a wide range of different organic contaminants, such as pesticides, herbicides, polycyclic aromatic hydrocarbons (PAHs), polychlorinated biphenyls (PCBs), and crude oil, at least when concentrations of the contaminant are not too high. The improvement of the soil due to their activity and the microorganisms in their digestive track can contribute to the accelerated removal of contaminants from soil, but sometimes their casts adsorb the pollutant so that its dissipation is delayed. There are limits, however, on how earthworms can be used to remediate soil, which will be discussed in this review.     

Gut load and food-retention time in the earthworms Lumbricus festivus and L. castaneus: A field study

Summary Lumbricus festivus and L. castaneus consume dung. In the field, below cow pats, their gut loads were about 0.15 and 0.14 g dry weight g^-1 ash-free dry weight of worm, respectively, but in free soil the loads were higher, about 0.21 and 0.19 g g^-1 ash-free dry weight of worm. The gut contents of dung were lighter than the total ingested material, at about 0.10 and 0.07 g dry weight g^-1 ash-free dry weight of worm, respectively. Field experiments showed that the retention time of dung ranged from >9 to 15 h for L. festivus, and from >3 to 6 h for L. castaneus. The experiments also indicated that L. festivus exploited 20- and 36-day-old dung in different ways, since the gut load was lower in those worms consuming 20-day-old dung than in those consuming 36- to 40-day-old dung. On the basis of these results the calculated consumption rate for L. festivus is 0.08 g dung day^-1 g live weight of worm^-1, and for L. castaneus 0.15 g dung day^-1, with retention times assumed to be at maximum, 15 h, for L. festivus and 6 h for L. castaneus. These calculations indicate that our field population of worms (75 g m^-2) consumes 10–15 t dung ha^-1 180 days^-1, corresponding to the amount of dung produced by 2–3 dairy cows.     

Plant biomass, soil water content and soil N:P ratio regulating soil microbial functional diversity in a temperate steppe: A regional scale study

Soil microorganisms are influenced by various abiotic and biotic factors at the field plot scale. Little is known, however, about the factors that determine soil microbial community functional diversity at a larger spatial scale. Here we conducted a regional scale study to assess the driving forces governing soil microbial community functional diversity in a temperate steppe of Hulunbeir, Inner Mongolia, northern China. Redundancy analysis and regression analysis were used to examine the relationships between soil microbial community properties and environmental variables. The results showed that the functional diversity of soil microbial communities was correlated with aboveground plant biomass, root biomass, soil water content and soil N: P ratio, suggesting that plant biomass, soil water availability and soil N availability were major determinants of soil microbial community functional diversity. Since plant biomass can indicate resource availability, which is mainly constrained by soil water availability and N availability in temperate steppes, we consider that soil microbial community functional diversity was mainly controlled by resource availability in temperate steppes at a regional scale.     

利用土壤传递函数估算土壤水力学特性研究进展

Characterization of soil hydraulic properties is important to environment management; however, it is well recognized that it is laborious, time-consuming and expensive to directly measure soil hydraulic properties. This paper reviews the development of pedotransfer functions (PTFs) used as an alternative tool to estimate soil hydraulic properties during the last two decades. Modern soil survey techniques like satellite imagery/remote sensing has been used in developing PTFs. Compared to mechanistic approaches, empirical relationships between physical properties and hydraulic properties have received wide preference for predicting soil hydraulic properties. Many PTFs based on different parametric functions can be found in the literature. A number of researchers have pursued a universal function that can describe water retention characteristics of all types of soils, but no single function can be termed generic though van Genuchten (VG) function has been the most widely adopted. Most of the reported parametric PTFs focus on estimation of VG parameters to obtain water retention curve (WRC). A number of physical, morphological and chemical properties have been used as predictor variables in PTFs. Conventionally, regression algorithms/techniques (statistical/neural regression) have been used for calibrating PTFs. However, there are reports of utilizing data mining techniques, e.g., pattern recognition and genetic algorithm. It is inferred that it is critical to refine the data used for calibration to improve the accuracy and reliability of the PTFs. Many statistical indices, including root mean square error (RMSE), index of agreement (d), maximum absolute error (ME), mean absolute error (MAE), coefficient of determination (r²) and correlation coefficient (r), have been used by different researchers to evaluate and validate PTFs. It is argued that being location specific, research interest in PTFs will continue till generic PTFs are developed and validated. In future studies, improved methods will be required to extract information from the existing database.     

Proteolytic activity in soil: A review

The aim of this work is to review current knowledge on inputs, sources and regulation of protease activities in soils from different ecosystems, while exploring limitations to proteolysis and N mineralisation. Extracellular proteases enter the soil via microbial production and other sources, including plant root exudates, animal excrements, decomposition processes and leaching from agro-industrial fertilisers. The synthesis and activities of proteases in soil are regulated by many factors, including climate, soil properties and the presence of organic compounds of plant and microbial origin. Two particularly important areas for future research are the regulation of proteolysis by low-molecular-weight organic compounds, including amino acids, sugars, flavonoids, plant hormones and siderophores, as well as the identification and characterisation of proteinaceous protease inhibitors of plant and microbial origin in the soil. Despite all the work that has been performed on soil proteases, our understanding of the roles of extracellular plant root proteases in N nutrition is weak. Furthermore, the regulation of soil proteolytic activities of different ecosystems, especially in terms of pollutant inputs and the impact of climate change, requires investigation. Other areas that pose important questions for the future include assessments of protease inhibitor inputs to the soil, regulation of these inhibitors via naturally occurring soil organic compounds and the interactions between soil organisms.     

Food quality affects production of Lumbricus terrestris (L.) under controlled environmental conditions

Birch leaves and horse manure were used to determine the effects of food quality on growth and reproduction of laboratory-reared Lumbricus terrestris. Animals grew to maturity within 6 months but attained a significantly (p < 0.001) larger adult size with manure (6.17 g) versus leaves (4.20 g). Cocoon production by recently-mated adults maintained in isolation, fed with birch leaves or horse manure, resulted in 4.53 and 3.84 cocoons ind.⁻¹ month⁻¹ respectively, with an initial hatchability of 86%, falling to zero after 18 months. Re-mating of these known individuals permitted long term monitoring of reproductive output (to 30 months). For the whole experimental period, overall hatchability of the 2010 cocoons produced was 44.4%. Median incubation time of those cocoons that hatched within accepted norms (less than 5 months at 15 °C) was 103 days and was not influenced by adult food type. A proportion (35.5%) of cocoons took in excess of 12 months to hatch. Adult mortality was minimal (25%) during the long term experiment but abnormal cocoon production was recorded after 2 years. Overall results demonstrate that food quality can have a significant influence on somatic and reproductive production of L. terrestris and these data may aid construction of production models for this earthworm in ecosystems with contrasting food quality.     

Beyond microbial diversity for predicting soil functions: A mini review

Since the advent of sequencing technologies,the determination of microbial diversity to predict microbial functions,which are the major determinants of soil functions,has become a major topic of interest,as evidenced by the 900 publications dealing with soil metagenome published up to 2017.However,the detection of a gene in soil does not mean that the relative function is expressed,and the presence of a particular taxon does not mean that the relative functions determined in pure culture also occur in the studied soil.Another critical step is to link microbial community composition or function to the product analyzed to determine flux rates.Indeed,flux rates might not only be highly dynamic,but several metabolites can depend on different reactions,which makes the link to one process of interest difficult or even impossible.This review also discusses biases caused by sampling,storage of samples,DNA extraction and purification,sequencing(amplicon-vs.metagenome sequencing),and bioinformatic data analysis.Insights and the limits of predicting microbial interactions by network inference methods are critically discussed,and finally,future directions for a better understanding of soil functions by using measurements of microbial diversity are presented.     

Methods for the introduction of bacteria into soil: A review

Summary Literature on the use of microbial inoculants to increase crop yields, to control soil-borne plant diseases, or to degrade pollutants has been reviewed. Established inoculant technology based on Rhizobium/peat inoculants has been summarized. Special emphasis has been placed on the use of carrier materials for the delivery of microbial inoculants. Some new developments, e.g., the use of synthetic carriers, have been highlighted. The fact that not only inoculant survival in carrier materials should be studied, but also the ecological consequences of the introduction of bacteria, has been stressed.     

Potential of arbuscular mycorrhizal fungi for soil health: A review

Soil health is an important component of “One Health”. Soils provide habitat to diverse and abundant organisms. Understanding microbial diversity and functions is essential for building healthy soils towards sustainable agriculture. Arbuscular mycorrhizal fungi (AMF) form potentially symbiotic associations with approximately 80% of land plant species that are well recognized for carbon flux and nutrient cycling. In addition to disentangling the signaling pathways and regulatory mechanisms between the two partners, recent advances in hyphosphere research highlight some emerging roles of AMF and associated microbes in the delivery of soil functions. This paper reviews the contribution of AMF to soil health in agroecosystems, with a major focus on recent progress in the contribution of hyphosphere microbiome to nutrient cycling, carbon sequestration, and soil aggregation. The hyphosphere microbiome and fungal stimulants open avenues for developing new fertilizer formulas to promote AMF benefits. In practice, developing AMF-friendly management strategies will have long-term positive effects on sustainable agriculture aiming at simultaneously providing food security, increasing resource use efficiency, and maintaining environment integrity.