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Drilosphere is the zone of interaction between earthworms and soil environments. Besides contributing to soil microporosity, the drilosphere provides habitat for a wide array of micro-organisms. There is growing evidence that the impact of earthworms on soil functions is mediated by microbial populations in the soil. It is important for agroecosystems that the soil quality is maintained for processes like decomposition, soil formation, and several other soil ecological properties. This review focuses on the structure and function of the Drilosphere, as well as its significance as a microbial 'hotspot'; which contributes to the agroecosystem earthworm-driven Drilosphere is a key to agroecosystem, boosting soil health and productivity through improved water movement, enhanced fertility, and nutrient cycling. Their mucus stabilizes soil and promote microbial activity, while interactions with plant roots supports nutrient exchange. Leveraging these dynamics can advance sustainable farming and yield higher agricultural productivity. [ABSTRACT FROM AUTHOR]

Contamination of soils with antimony (Sb) is becoming increasingly severe and widespread, and the associated ecological risks cannot be ignored. To evaluate how different Sb forms affected the earthworm Eisenia fetida in soil, the biomarker response index (BRI), effect addition index (EAI), and microbial diversity were characterized after single and joint application of Sb(III) and Sb(V). The results showed that Sb(III) was better enriched by earthworms than Sb(V). The metallothionein (MT) content in earthworms increased under Sb stress, and the superoxide dismutase (SOD), catalase (CAT), and glutathione S-transferase (GST) activities also showed an increasing trend, suggesting waken-up antioxidant capacity. Severe alterations for health status were observed under combined treatment. Additionally, the EAI indicated that Sb(III) and Sb(V) had synergistic and antagonistic effects at low and high concentrations, respectively. The bacterial populations in the drilosphere (gut and burrow lining) appeared to be more susceptible to Sb contamination than in the non-drilosphere, their specific microecology may be an important factor in soil Sb migration and transformation. The abundance of Actinobacteria exhibited a significant decrease with increasing concentrations of single Sb(III) and Sb(V), while the abundance of Bacteroidia increased. The correlation heatmap showed that Sphingobacterium faecium was highly tolerant to Sb. These results provide not only an important basis for the ecological risk assessment of Sb in the soil environment but also new insights into the altered drilosphere bacterial communities under Sb stress.

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Although the potential of vermiremediation for restoring metal-contaminated soils is promising, the effects of earthworms on the availability of soil metals are still debatable. Most previous studies considered the soil as a "whole black box." Mobilization or immobilization of metals are affected by earthworm activities within drilosphere hotspots under different soil conditions, which has not been specifically studied. Therefore, an improved 2D terrarium was designed to study the impact of earthworm activities on cadmium (Cd) fate in the drilosphere hotspots (burrow wall soils, burrow casts, and surface casts) of different artificially spiked Cd treatments (CK: 0 mg kg-1; LM: 1 mg kg-1; and HM: 5 mg kg-1) with different organic amendments (2% and 10%). The results revealed that Cd increased earthworm activities with the highest cast production in HM and the highest burrow length in LM. Earthworms exhibited a stronger tendency to reduce total Cd concentration by 4.48–13.58% in casts of LM soils, while 3.37–5.22% in burrow walls under HM treatments. Overall, earthworms could increase the availability of Cd in casts under all conditions (55.46–121.01%). The organic amendments decreased the total Cd concentration and increased the availability of Cd in the disturbed soil. A higher amount of organic amendment significantly decreased total Cd concentration of the drilosphere by 1.16–5.83% in LM and HM treatments, while increasing DTPA-Cd concentrations in all components by 23.13–55.20 %, 14.63–35.11%, and 3.30–11.41% in CK, LM, and HM treatments, respectively, except for earthworm non-disturbed soil and no-earthworm soil in HM treatments. Redundancy analysis (RDA) revealed that the moisture, pH, and total carbon contents in soil are the main factors affecting Cd bioavailability. In this study, we decoded the "black box" of soil by making it relatively simple to better understand the effects and mechanisms of earthworm activities on soil metal availability and consequently provided comprehensive insights for using earthworms in soil vermiremediation. [ABSTRACT FROM AUTHOR]

Using a mesocosm experiment, we investigated the individual and interaction effects of two earthworm species with contrasting behaviour on soil structure and water transfers. The anecic species Amynthas zenkevichi (Thai, 1982) and the polyhumic endogeic species Pontoscolex corethrurus (Müller, 1857) were incubated in repacked soil columns alone or together for three months under laboratory conditions. The volume of belowground casts, empty burrows and lateral soil compaction were assessed using X-ray computed tomography. The production of surface casts and the amount of food ingested were also recorded. The soil moisture at 7 cm depth and water evaporation of the whole column were monitored regularly. Soil water infiltration was assessed using the Beerkan method at the end of the experiment. A. zenkevichi burrows were less numerous (25 vs. 85), more continuous (41 vs. 0 cm3), more connected from the surface to the bottom of the columns (17 vs. 0 cm3) and more compacted laterally (243 vs. 92 cm3) than those of P. corethrurus. Conversely, P. corethrurus burrows were more abundant in the top 5 cm of the columns and more backfilled by casts than those of A. zenkevichi (36 vs. 5 %). Both species ingested buffalo dung provided at the soil surface and produced surface casts at similar rates. Interactions resulted in an increase in surface activity of more than 40 % and a decrease in the depth and continuity of burrow systems. The water infiltration rate was increased by 3.5 times (compared to the control soil without earthworms) by A. zenkevichi burrows and was not modified by interactions. P. corethrurus increased the cumulative water evaporation by 10 % and decreased soil moisture by 3 % (compared to the control soil without earthworms), whereas A. zenkevichi had marginal effects on these parameters. Globally, interactions led to a slight positive synergistic effect on soil resistance to water loss by evaporation, which was likely related to the increase in surface casting activity. To conclude, this study stresses the importance of considering interactions between earthworms in soil and the need to confirm our findings under natural conditions. • The two species had contrasting belowground activities but similar surface activities. • Anecics had a larger impact on water infiltration than endogeics. • Only the endogeics increased soil water evaporation and decreased soil moisture. • Interactions increased surface activity and decreased the depth of burrows. • The two species together had a positive effect on soil resistance to water loss by evaporation. [ABSTRACT FROM AUTHOR]

Contamination of soils with antimony (Sb) is becoming increasingly severe and widespread, and the associated ecological risks cannot be ignored. To evaluate how different Sb forms affected the earthworm Eisenia fetida in soil, the biomarker response index (BRI), effect addition index (EAI), and microbial diversity were characterized after single and joint application of Sb(III) and Sb(V). The results showed that Sb(III) was better enriched by earthworms than Sb(V). The metallothionein (MT) content in earthworms increased under Sb stress, and the superoxide dismutase (SOD), catalase (CAT), and glutathione S-transferase (GST) activities also showed an increasing trend, suggesting waken-up antioxidant capacity. Severe alterations for health status were observed under combined treatment. Additionally, the EAI indicated that Sb(III) and Sb(V) had synergistic and antagonistic effects at low and high concentrations, respectively. The bacterial populations in the drilosphere (gut and burrow lining) appeared to be more susceptible to Sb contamination than in the non-drilosphere, their specific microecology may be an important factor in soil Sb migration and transformation. The abundance of Actinobacteria exhibited a significant decrease with increasing concentrations of single Sb(III) and Sb(V), while the abundance of Bacteroidia increased. The correlation heatmap showed that Sphingobacterium faecium was highly tolerant to Sb. These results provide not only an important basis for the ecological risk assessment of Sb in the soil environment but also new insights into the altered drilosphere bacterial communities under Sb stress., Competing Interests: The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (© 2024 The Authors.)

The drilosphere is described as a microbial hotspot in soil, which is rich in labile nutrients. However, phosphorus (P) quality and availability change along the drilosphere from labile organically-bound P in upper layers to immobilized Ca-phosphate with increasing depth. In this study, we postulated (1) that microbial P turnover is generally accelerated in the drilosphere compared to bulk soil as a result of an increase in labile nutrients, and (2) that P mineralization is dominant in upper layers of the drilosphere and P solubilization in deeper layers of the drilosphere, respectively. Furthermore, we hypothesized (3) that the enhanced P mobilization in the drilosphere favors copiotrophic and earthworm gut derived bacteria like Pseudomonadaceae , which distinguishes the community composition from that of the bulk soil. We investigated the effect of earthworm activities on potential phosphatase activities, abundance and diversity of bacterial communities involved in P mobilization (gcd , phoD, appA, phnX, phoN) and uptake (pstS, pitA) along a soil depth gradient in a short rotation coppice in Northeast Germany. Potential phosphatase activities and gene abundances revealed increasing potentials for organic P mineralization and high-affinity P uptake via the Pst system in the drilosphere below 50 cm depth. Conversely, in the upper soil layers potential for bacterial P mobilization were similar between drilosphere and bulk soil. We also observed drilosphere-related changes in the composition of bacterial communities involved in P solubilization and low-affinity P uptake via the Pit system below 60 cm depth. In the drilosphere compared to bulk soil, Pseudomonadaceae were enriched and played a major role in solubilization and low-affinity uptake of P, while bacteria potentially involved in N turnover like Bradyrhizobiaceae and Phyllobacteriaceae were less abundant. Our study suggested that earthworms increased the potential enzyme activity and abundance of bacteria carrying the Pho-regulon genes in the drilosphere of the deeper soil layers. Furthermore, earthworm-derived bacteria were enriched in the drilosphere at the same soil depth, especially those involved in P solubilization and carrying the Pit system. • Increased potential for bacterial P mineralization and uptake via the Pst system in the drilosphere below 50 cm depth • Enhanced P turnover potential was highly associated with P scarcity • Earthworm-induced changes in bacterial community composition involved in P solubilization and uptake via the Pit system in subsoil • Earthworm-derived Pseudomonadaceae highly contributed to P solubilization and uptake via the Pit system in the drilosphere subsoil [ABSTRACT FROM AUTHOR]

Earthworms, as ecosystem engineers, are known to inoculate organic residues with microorganisms, consuming and processing mineral soil and its organic matter, which may lead to changes in soil quality. We conducted a field experiment, in randomized complete blocks, using soil mesocosms incubated with 4 different densities of earthworms: L1 = 100 earthworms m−2, L2 = 500 earthworms m−2, L3 = 1000 earthworms m−2, and C0 = control without earthworms. All earthworm populations consisted of epigeic, endogeic and anecic earthworms in a ratio of 2:3:3, respectively. The earthworms used were Lumbricus rubellus , Octolasium cyaneum , Aporrectodea calliginosa , and Lumbricus friendi. Physical (aggregate stability and structure heterogeneity), chemical (pH CaCl 2 , Mineral-N, Olsen-P, exchangeable K+, Ca+2, Mg+2, Na+ and Al) and biological indicators (microbial biomass) were assessed to evaluate the changes in soil quality. After applying an equivalent of 18 tons ha−1 of aged horse manure to each mesocosm, soils were sampled at two depths: 0–25 cm and 25–50 cm and on two occasions: prior to inoculating the soil with earthworms, T0, and 96 days after the soil was inoculated with earthworms, T1. Earthworms were found to increase soil mineral nitrogen (NO 3 + NH 4) and exchangeable calcium (Ca+2) and decrease exchangeable aluminum (Al+3) and soil microbial biomass (g C g−1). These results could imply that earthworms promote the mineralization of organic nitrogen, increase the cation content, and decrease the solubility of the principal toxic elements in soil (Al+3). Finally, the incorporation of earthworms at the 0–25 cm depth proved to increase the quality of the degraded soil, which was reflected in the chemical quality parameters. [Display omitted] • Use of soil indicators to assess quality. • Earthworms decrease the solubility of exchangeable Al in soil. • High earthworm density increases the quality of a degraded volcanic soil. [ABSTRACT FROM AUTHOR]

The positive effect of earthworms on soil processes and plant growth has been extensively documented. The capacity of earthworms to decompose organic matter has been attributed to the microbial communities that inhabit their digestive track or the structures they build, which in turn contribute to make up the drilosphere, a hotspot for microbial activity. However, how earthworms modify the structure of soil microbial communities and how these changes affect soil microbial processes is still unclear. Do earthworms reduce microbial abundance and activity because they feed on microorganisms or do they select and stimulate specific microbial groups? We hypothesise that “the effect of earthworms on nutrient cycling and plant growth is not only a direct effect but is mainly mediated indirectly, via modifications of the microbial community.” The objective of this review is to synthesize the existing literature concerning the influence of earthworms on the structure and function of soil microbial communities, as well as to understand how earthworm-induced changes in the soil microbiota would in turn impact soil processes, particularly those occurring in the rhizosphere and involved in plant growth and health. Recent reports have shown that specific bacterial groups consistently increase in soils where earthworms are present, regardless of the earthworm functional group. The extent of this increase seems to be dependent upon the type of substrate under study. Our synthesis also reveals that endogeic and anecic earthworms regularly induce an increase in soil nutrients, whilst this positive effect is not as evident in the presence of epigeic earthworms. The effect of earthworms on nutrient cycling has been further investigated with microbial functional genes, although existing reports largely focus on nitrogen cycling. Earthworms seem to enhance denitrification, most likely through the increase in organic compounds due to organic matter decomposition. By enhancing soil nutrient availability, earthworms indirectly promote plant growth, which has also been attributed to the induction of signal molecules. However, no experiment to date has been able to prove a direct causal relationship between specific signal molecules, earthworms and plant growth promotion. Finally, we propose a framework for earthworm-microbiota interactions and recommend further research.

Most subsoil carbon (C) turnover occurs in biopore hotspots such as root channels and earthworm burrows. Biopores allocate large C amounts into the subsoil, where a vast capacity for long-term C sequestration is predicted. We hypothesise that organic matter (OM) cycling in biopores depends on their origin. Earthworm and root biopores were induced under field conditions and were sampled from the subsoil (45–75 and 75–105 cm) after two years of biopore formation. The effects of biopore formation on OM decomposition were studied by biomarkers: neutral sugars, cutin and suberin-derived lipids, lignin-derived phenols and free lipids. The degradation stage of OM was biopore type-specific but was only governed by the soil depth in root biopores. Degradation of OM increased from earthworm biopores to root biopores and bulk soil. Hemicelluloses (GM/AX ratio) were more strongly degraded than lignin side-chains (relative change from initial values). Two years of microbial processing during biopore formation increased the GM/AX ratio in earthworm biopores from 0.65 to 1.05 and in root biopores from 0.15 to 1.35 (both relative to source biomasses). Root biopores and bulk soil had the highest GM/AX ratios (1.2–1.3), hinting to rapid processing of plant residues and accumulation of microbial residues. The regular, frequent OM inputs by earthworms stimulated microbial growth and processing of mostly bioavailable OM and, thus, relatively enriched more persistent OM (e.g. lignin). Syringyl subunits of lignin underwent low (ratio changed from 0.35 to 0.55 relative to initial input) and vanillyl subunits underwent almost no processing in earthworm biopores indicating the preferential microbial utilisation of the easily available compounds frequently replenished by earthworm activity. After two years of decomposition of the root detritus, mainly structural plant material was enriched in root biopores. Short periods (6 months) of earthworm activity effectively recharged the highly processed OM in root biopores with fresh OM. In total, deep-rooting catch crops and short-term earthworm activities promote C accumulation in the subsoil followed by biopore-specific microbial processing predominantly governed by the C input frequency. As root biopores are up to 40 times more common than earthworm biopores, they dominate the OM input into subsoils. Such C inputs create several years lasting hotspots for preferential root growth and nutrient mobilisation in the subsoil. We conclude that root- and earthworm-derived biopores are vertical pathways for plant C from the soil surface into the subsoil and for intensive processing of litter C and sequestration of microbial necromass. [ABSTRACT FROM AUTHOR]

The current study describes quantitatively the interactions between (a) activity of extracellular enzymes: β‐glucosidase (BG), protease (Prot), alkaline phosphatase (Alk P), and acid phosphatase (Acid P); (b) intracellular activities: respiration, dehydrogenase activity (DHA); and (c) C and N contents of soil: Corg—organic carbon, Cmic—microbial biomass, ratio Cmic:Corg, Ntot—total nitrogen, and ratio C:N in earthworm‐affected and surrounding loess soil under a pear orchard. The soil tested was collected from: burrow walls (0–3 mm) and a transition zone (3–7 mm) both forming drilosphere, bulk soil (20 mm from the burrow wall), and cast aggregates. The enzyme activities were presented as total 'tot' (per gram of dry soil mass) and specific 'spec' (per Cmic). It was shown that soil Cmic, basal respiration, BGtot, Prottot, and Alk Ptot were significantly positively correlated with Corg, Cmic:Corg, Ntot, and C:N (r = 0.627–0.976). However, the specific BGspec, Protspec, and Acid Pspec were negatively correlated, while specific respiration (qCO2) and Alk Pspec were not correlated with most soil characteristics. The relationships between DHAtot, DHAspec, and all soil characteristics were positive and had the highest correlation coefficients (r = 0.703–0.989). Response ratio of the enzyme activity was more strongly affected by enzyme activity location (extracellular vs. intracellular) and presentation method (total vs. specific), than by earthworm‐affected soil compartments. The results suggest a need to consider the method of enzyme activity presentation (specific vs. total) that supports understanding ecologically relevant microbial processes in earthworm‐inhabited orchard soils. [ABSTRACT FROM AUTHOR]

Purpose: Dynamical in situ observation of biological and climatic structuring factors involved in pedogenesis has not previously been possible in a way that would consider the early stages of pedogenesis. If studies have explored the effect of pedogenetic factors on soil structure, none have succeeded in ranking them in view of the intensity of their effects. We propose a novel approach for describing the aggregation process for a constructed Technosol obtained from a process of pedological engineering.Materials and methods: We focus on agents including plants, macrofauna, and water, and we use (i) a dynamic in situ observation and (ii) the quantification of the evolution of selected descriptors of pores and aggregates. They are quantified from high-resolution images obtained with the Soilinsight® device. Associating those images with each other, movies of interactions between soil and organisms over a 14-month non-destructive soil evolution experiment are made.Results and discussion: Agents influencing aggregation—plant roots, earthworms, and water—can be ranked according to their impact on soil structure. During the studied period of evolution, wetting-drying cycles are the first to operate. The intensity of their action on soil structure is dominant at the very first stages of pedogenesis. Despite this ranking of agents, over the long term, plants and earthworms have a more intense effect on soil structure than wetting-drying cycles.Conclusions: The method applied to observe and quantify soil structure dynamics is thus proposed as a helpful approach to modeling other processes involved in soil functioning and evolution in relation to their ability to fulfill ecosystem services. [ABSTRACT FROM AUTHOR]

Arbuscular mycorrhizal fungi are recognized as important drivers of plant health and productivity in agriculture but very often existing knowledge is limited to the topsoil. With growing interest in the role of subsoil in sustainable agriculture, we used high-throughput Illumina sequencing on a set of samples encompassing drilosphere, rhizosphere and bulk soil, in both top- and subsoil. Our results show subsoil AMF communities harbor unique Operational Taxonomic Units (OTUs) and that both soil depths differ in community structure both at the OTU and family level. Our results emphasize the distinctness of subsoil AMF communities and the potential role of subsoil as a biodiversity reservoir. [ABSTRACT FROM AUTHOR]

Biopores are hotspots of nutrient mobilisation and shortcuts for carbon (C) into subsoils. C processing relies on microbial community composition, which remains unexplored in subsoil biopores. Phospholipid fatty acids (PLFAs; markers for living microbial groups) and amino sugars (microbial necromass markers) were extracted from two subsoil depths (45-75 cm ; 75-105 cm) and three biopore types: (I) drilosphere of Lumbricus terrestris L., (II) 2-year-old root biopores and (III) 1.5-year-old root biopores plus six 6 months of L. terrestris activities. Biopore C contents were at least 2.5 times higher than in bulk soil, causing 26-35 times higher Σ PLFAs g soil. The highest Σ PLFAs were found in both earthworm biopore types; thus, the highest soil organic matter and nutrient turnover were assumed. Σ PLFAs was 33% lower in root pores than in earthworm pores. The treatment affected the microbial community composition more strongly than soil depth, hinting to similar C quality in biopores: Gram-positives including actinobacteria were more abundant in root pores than in earthworm pores, probably due to lower C bioavailability in the former. Both earthworm pore types featured fresh litter input, promoting growth of Gram-negatives and fungi. Earthworms in root pores shifted the composition of the microbial community heavily and turned root pores into earthworm pores within 6 months. Only recent communities were affected and they reflect a strong heterogeneity of microbial activity and functions in subsoil hotspots, whereas biopore-specific necromass accumulation from different microbial groups was absent. [ABSTRACT FROM AUTHOR]

Numerous researches have been conducted on the effects of biotic and abiotic-induced aging on the physicochemical characteristics and functions of biochar; however, the impacts of earthworm-induced aging on biochar have not been reported. Hence, we conducted a microscopic experiment simulating a 'drilosphere' to explore the influence of earthworm activity on the natural aging of rice husk biochar (RHBC) through the difference in biochar characteristics after aging in drilosphere and non-drilosphere. The earthworm activity increases the available nitrogen (AN) and dissolved organic matter (DOM) contents of aged RHBC and changes its composition. The increase of DOM and AN content may recruit more microorganisms to colonize biochar and accelerate the biological oxidation of biochar. Furthermore, earthworm activity significantly increased the contents of oxygen (O) and O-containing functional groups in the aged RHBC and decreased the stability (aromaticity) of the aged RHBC, suggesting that the earthworm activity accelerates the natural aging of biochar. Earthworm feeding promotes physical damage to biochar. Besides, the earthworm activity decreased the pH, hydrophilicity and specific surface area (SSA) of aged RHBC but enhanced the adsorption capacity of aged RHBC for heavy metals. The higher content of O-containing functional groups on the surface of drilosphere-aged RHBC was the main reason for its higher adsorption performance. Earthworm feeding promotes physical damage to biochar. These results indicate that earthworm activity can accelerate the natural aging of biochar and alter its physicochemical characteristics and functions. This study illustrates how biochar characteristics change in earthworm-soil systems, which will help scientifically evaluate the long-term effectiveness of biochar. [Display omitted] • Earthworm activity changes the composition of soluble components of aged biochar. • Earthworm activity accelerates the aging of biochar and reduces its stability. • Earthworm activity changes the physical and chemical properties of aged biochar. • Earthworm activity improves the capacity of aged biochar to adsorb heavy metals. [ABSTRACT FROM AUTHOR]

Soil microbial communities provide a wide range of soil functions including nutrient cycling, soil formation, and plant growth promotion. On the small scale, nutrient rich soil hotspots developed from soil animal or plant activity are important drivers for microbial communities and their activity pattern. Nevertheless, in subsoil, the spatial heterogeneity of microbes with diverging lifestyles has been barely considered so far. In this study, the phylogenetic composition of the bacterial and archaeal microbiome based on 16S rRNA gene pyrosequencing was investigated in the soil compartments bulk soil, drilosphere, and rhizosphere in topsoil and in the subsoil of an agricultural field. With co-occurrence network analysis, the spatial separation of typically oligotrophs and heterotrophs in subsoil and hotspots was assessed. Four co-occurring bacterial communities were identified and attributed to bulk topsoil, bulk subsoil, drilosphere, and rhizosphere. The bacterial phyla Proteobacteria and Bacteroidetes, which represent many copiotrophic bacteria, are affiliated to the hotspot communities – the rhizosphere and drilosphere – both in topsoil and subsoil. Acidobacteria, Actinobacteria, Gemmatimonadetes, Planctomycetes, and Verrucomicrobia with many oligotrophic bacteria, are the abundant groups of the bulk subsoil community. The bacterial core microbiome in this soil was estimated and only covers 7.6% of the bacterial sequencing reads but includes both oligotrophic and copiotrophic bacteria. Instead, the archaeal core microbiome includes 56% of the overall archaeal diversity and comprises only the ammonium oxidizing Nitrososphaera. Thus, the spatial variability of nutrient quality and quantity strongly shapes the bacterial community composition and their interaction in subsoil, whereas archaea are a stable backbone of the soil prokaryotes.

Earthworms boost microbial activities and consequently create hotspots in soil. Although the presence of earthworms is thought to change the soil enzyme system, the distribution of enzyme activities inside worm burrows is still unknown. For the first time, we analyzed enzyme kinetics and visualized enzyme distribution inside and outside worm burrows (biopores) by in situ soil zymography. Kinetic parameters ( V max and K m ) of 6 enzymes – β-glucosidase (GLU), cellobiohydrolase (CBH), xylanase (XYL), chitinase (NAG), leucine aminopeptidase (LAP) and acid phosphatase (APT) – were determined in pores formed by Lumbricus terrestris L. In earthworm burrows, the spatial distributions of GLU, NAG and APT become observable in zymogram images. Zymography showed a heterogeneous distribution of hotspots in the rhizosphere and worm burrows. The hotspot areas were 2.4–14 times larger in the burrows versus reference soil (soil without earthworms). The significantly higher V max values for GLU, CBH, XYL, NAG and APT in burrows confirmed that earthworms stimulated enzyme activities. For CBH, XYL and NAG, the 2- to 3-fold higher K m values in burrows indicated different enzyme systems with lower substrate affinity compared to reference soil. The positive effects of earthworms on V max were cancelled by the K m increase for CBH, XYL and NAG at a substrate concentration below 20 μmol g −1 soil. The change of enzyme systems reflected a shift in dominant microbial populations toward species with lower affinity to holo-celluloses and to N-acetylglucosamine, and with higher affinity to proteins as compared to the reference soil. We conclude that earthworm burrows are microbial hotspots with much higher and denser distribution of enzyme activities than reference soil. [ABSTRACT FROM AUTHOR]

Copyright of Basic & Applied Ecology is the property of Urban & Fischer Verlag and its content may not be copied or emailed to multiple sites or posted to a listserv without the copyright holder's express written permission. However, users may print, download, or email articles for individual use. This abstract may be abridged. No warranty is given about the accuracy of the copy. Users should refer to the original published version of the material for the full abstract. (Copyright applies to all Abstracts.)

Microbial communities in soil provide a wide range of ecosystem services. On the small scale, nutrient rich hotspots in soil developed from the activities of animals or plants are important drivers for the composition of microbial communities and their functional patterns. However, in subsoil, the spatial heterogeneity of microbes with differing lifestyles has been rarely considered so far. In this study, the phylogenetic composition of the bacterial and archaeal microbiome based on 16S rRNA gene pyrosequencing was investigated in the soil compartments bulk soil, drilosphere, and rhizosphere in top- and in the subsoil of an agricultural field. With co-occurrence network analysis, the spatial separation of typically oligotrophic and copiotrophic microbes was assessed. Four bacterial clusters were identified and attributed to bulk topsoil, bulk subsoil, drilosphere, and rhizosphere. The bacterial phyla Proteobacteria and Bacteroidetes, representing mostly copiotrophic bacteria, were affiliated mainly to the rhizosphere and drilosphere--both in topsoil and subsoil. Acidobacteria, Actinobacteria, Gemmatimonadetes, Planctomycetes, and Verrucomicrobia, bacterial phyla which harbor many oligotrophic bacteria, were the most abundant groups in bulk subsoil. The bacterial core microbiome in this soil was estimated to cover 7.6% of the bacterial sequencing reads including both oligotrophic and copiotrophic bacteria. In contrast the archaeal core microbiome includes 56% of the overall archaeal diversity. Thus, the spatial variability of nutrient quality and quantity strongly shapes the bacterial community composition and their interaction in subsoil, whereas archaea build a stable backbone of the soil prokaryotes due to their low variability in the different soil compartments. [ABSTRACT FROM AUTHOR]

The spatial heterogeneity of nutrient turnover in subsoils has been rarely studied in the past, although drilosphere and rhizosphere are found to be important microbial hotspots in this oligotrophic environment. In this study, we measured different potential enzyme activities in different soil compartments of subsoil and topsoil. It could be shown that the activities of hydrolases, which cleave readily available organic substrates, are significantly higher in samples from the drilosphere and rhizosphere both in topsoil and subsoil. In bulk soil, hydrolase activities decrease with depth. In contrast, oxidative enzymes, which are involved in the decay of recalcitrant organic material, are released from the microbial community especially in the bulk fraction of subsoil. This emphasizes the importance of subsoil for nutrient acquisition and gives evidence for a distinct spatial separation of microbes with diverging lifestyles. [ABSTRACT FROM AUTHOR]

Bioporosity in subsoils is strongly influenced by either deep rooting taproots or earthworms, which can affect important properties for plant growth provided by soils. The open question of this study was, in how far the combination of X-ray computed tomography (XRCT) and endoscopy may support the analysis of soil bioporosity and the effects induced by earthworm activity. The hypothesis were that earthworms can re-open, close and re-connect pores as well as change the biopore wall properties (by leaving coatings at the wall) and that with a combination of XRCT and endoscopy much more detailed information about those changes can be observed. Soil monoliths were collected from a Haplic Luvisol (developed from loess) at the experimental station Klein Altendorf (University of Bonn, Germany). The microcosms were then prepared under laboratory conditions in terms of temperature, soil temperature gradient, relative humidity, illumination, watering and fertilizer. The microcosms were examined before cultivation and incubation, and re-examined after one-month earthworm incubation ( Lumbricus terrestris L.), three months wheat growth period and shoot harvesting. XRCT and endoscopy were used to qualitatively and quantitatively analyze the soil microcosms. After the incubation with earthworms, physical properties like porosity and accessible surface area of biopores have changed with respect to the original state. Coatings at the biopore walls resulted in smaller pore diameters of large biopores with a diameter >0.5 cm, while some of them have previously been pores with diameters <0.5 cm. Contrary to this, biopores with a diameter <0.5 cm were increased after earthworms, i.e. widened due earthworm passage. It was found that earthworms may have disconnected lateral pores that reach into the bulk soil from the vertical biopores by pore wall coatings. After incubation, biopores lined with fresh earthworm coatings in the monolith increased from 30% to 80%, which may indicate that earthworms potentially have a considerable influence on biopore properties and therefore the physical, chemical and microbiological environment which roots are exposed to in biopores. [ABSTRACT FROM AUTHOR]

Spatial and temporal dynamics of microbial community structure and function in subsoils have been rarely studied in the past. In this paper we present data on how bacterial communities as well as selected functional groups of microbes change in the rhizosphere, the drilosphere, and in bulk soil over time in topsoil as well as in subsoil. We show that the overall richness of bacteria and abundance of nitrifiers and denitrifiers decreases in bulk soil with soil depth. However, these effects were not or to a much lower degree observed in the rhizosphere and the drilosphere. Temporal fluctuations contributed by far less than spatial factors to the dynamics of bacterial communities and abundance of nitrifiers and denitrifiers in all compartments independent from the soil depth. [ABSTRACT FROM AUTHOR]

Abstract: The burrowing activity of earthworms leads to soil structural changes by compaction of surrounded soil matrix, which represents the so-called drilosphere. This is a distinct volume around the resulting macropore, which controls various soil processes. In the aim to detect the bulk density variation around earthworm burrows, two repacked soil columns from a silt loam with defined water content and bulk density were created, separately inoculated with the anecic earthworm species Lumbricus terrestris, stored under constant conditions for 7weeks and finally analyzed by means of X-ray computed tomography. A basic voxel approach was developed to conduct a gapless analysis of the drilosphere and surrounding soil matrix. The measured Hounsfield Units of basic voxels (pixel size of 0.25mm and slice thickness of 0.625mm) were converted into the corresponding dry bulk density (BD). The initial mean BD of an entire horizontal soil slice was 1.34Mgm−3 (core 1) and 1.38Mgm−3 (core 2). Due to burrowing L. terrestris compacted the inner boundary of the drilosphere (burrow wall) up to a BD of at least 1.75Mgm−3 which equates to more than 30% compared to the soil matrix. The BD decreased from the inner boundary of the drilosphere to its outer boundary, which is a transition zone to the surrounding soil matrix. However, the BD decrease was not concentrically uniform but revealed heterogeneous patterns of zones with different BD-classes. This locally heterogeneous BD distribution is an evidence of radial earthworm forces acting anisotropically. We conclude that BD heterogeneity in the drilosphere of L. terrestris might have notable implications for the understanding of lateral transfer of water and solutes in the soil profile. [Copyright &y& Elsevier]

Background and aims: The significance of biopores for nutrient acquisition from the subsoil depends on root-soil contact, which in turn is influenced by root architecture. The aim of this study was to detect differences regarding the architecture and root-soil contact of homorhizous barley roots ( Hordeum vulgare L.) and allorhizous oilseed rape roots ( Brassica napus L.) growing in biopores. Methods: In situ endoscopy was used as a technique that allows non-destructive display of pore wall characteristics and root morphology inside large biopores under field conditions. Results: For both crops, about 85 % of all roots did establish contact to the pore wall. However, according to their different root architecture, the two crops varied in their strategy of resource acquisition: While barley was characterized by thin vertical or ingrowing roots, most of them in direct contact to the pore wall, oilseed rape established contact to the pore wall predominantly via lateral roots. Conclusions: Root morphological and pore wall assessment with in situ endoscopy in combination with detailed studies of soil biochemical and soil physical parameters of the pore wall is considered an essential prerequisite for more precise future modelling of nutrient acquisition and uptake. [ABSTRACT FROM AUTHOR]

Microbial degradation of 2-methyl-4-chlorophenoxyacetic acid ( MCPA) in soil is enhanced by earthworms and initiated by tfdA-like, cadA and r/sdpA gene encoding oxygenases. Copy numbers of such genes increased during MCPA degradation in soil, and MCPA stimulated transcription of tfdA-like and r/sdpA genes up to 4×. Transcription of cadA was detected in the presence of MCPA only. DNA stable isotope probing after consumption of 0.6-0.8 mg [ABSTRACT FROM AUTHOR]

Highlights: [•] The properties of casts collected at 20–30cm soil depth suggest they were mainly produced by endogeic species. [•] Carbon and nitrogen content and NIRS signature were similar between casts and surrounding soils. [•] The usefulness of using NIRS to differentiate cast aggregation may be context-dependent. [Copyright &y& Elsevier]

Abstract: In arable farming systems, the term ‘subsoil’ refers to the soil beneath the tilled or formerly tilled soil horizon whereas the latter one is denoted as ‘topsoil’. To date, most agronomic and plant nutrition studies have widely neglected subsoil processes involved in nutrient acquisition by crop roots. Based on our current knowledge it can be assumed that subsoil properties such as comparatively high bulk density, low air permeability, and poverty of organic matter, nutrients and microbial biomass are obviously adverse for nutrient acquisition, and sometimes subsoils provide as little as less than 10% of annual nutrient uptake in fertilised arable fields. Nevertheless, there is also strong evidence indicating that subsoil can contribute to more than two-thirds of the plant nutrition of N, P and K, especially when the topsoil is dry or nutrient-depleted. Based on the existing literature, nutrient acquisition from arable subsoils may be conceptualised into three major process components: (I) mobilisation from the subsoil, (II) translocation to the shoot and long-term accumulation in the Ap horizon and (III) re-allocation to the subsoil. The quantitative estimation of nutrient acquisition from the subsoil requires the linking of field experiments with mathematical modelling approaches on different spatial scales including Process Based Models for the field scale and Functional–Structural Plant Models for the plant scale. Possibilities to modify subsoil properties by means of agronomic management are limited, but ‘subsoiling’ – i.e. deep mechanical loosening – as well as the promotion of biopore formation are two potential strategies for increasing access to subsoil resources for crop roots in arable soils. The quantitative role of biopores in the nutrient acquisition from the subsoil is still unclear, and more research is needed to determine the bioaccessibility of nutrients in subsoil horizons. [Copyright &y& Elsevier]

Earthworms, arbuscular mycorrhiza fungi (AMF) and roots are important components of the belowground part of terrestrial ecosystem. However, their interacting effects on soil properties and plant growth are still poorly understood. A compartmental experimental design was used in a climate chamber in order to investigate, without phosphorus (P) addition, the single and combined effects of earthworms ( Allolobophora chlorotica), AMF ( Glomus intraradices) and roots ( Allium porrum) on soil structure, nutrient concentration and plant growth. In our experimental conditions, plant roots improved soil structure stability (at the level of macroaggregates) whereas earthworms decreased it. AMF had no effect on soil structure stability but increased P transfer from the soil to the plant and significantly increased plant biomass. Earthworms had no direct influence on P uptake or plant biomass, and the N/P ratio measured in the shoots indicated that P was limiting. Interactions between AMF and earthworms were also observed on total C and N content in the soil and on total root biomass. Their effects varied temporally and between the different soil compartments (bulk soil, rhizosphere and drilosphere). After comparison with other similar studies, we suggest that effects of earthworms and AMF on plant production may depend on the limiting factors in the soil, mainly N or P. Our experiment highlights the importance of measuring physical and chemical soil parameters when studying soil organism interactions and their influence on plant performance. [ABSTRACT FROM AUTHOR]

Abstract: The burrowing activity of earthworms creates a distinct area around the resulting macropores called the drilosphere, which controls various soil processes. Density and microstructure of the drilosphere were studied and compared with those of the surrounding soil. For this purpose soil cores were separately inoculated with the vertically burrowing earthworm species Lumbricus terrestris. After 70 days some cores were compacted by a hydraulic press (250 kPa) and all cores were analysed by means of X-ray computed tomography. Mean Hounsfield Units were measured for concentric ROI cylinders (ROI=region of interest) of increasing diameters located around vertical macropore sections within selected horizontal slices. Based on these data we estimated stepwise the distribution of bulk density from the inner boundary of the drilosphere to the surrounding soil. In uncompacted soil the bulk density of the drilosphere was increased by 11% over that of the surrounding soil. In cross section, drilosphere and burrow form a concentric area with a total radius up to 2.2 cm. Soil compaction increased the dry bulk density of soil and decreased the diameter of earthworm burrows. Moreover, we found a less dense part of soil between the dense drilosphere and the remaining soil of the compacted core. Scanning electron microscopy revealed that the coarse silt particles of the bulk soil were rearranged to a parallel orientation due to compaction whereas the microstructure of the drilosphere remained unchanged. In any case, the drilosphere revealed a very homogeneous and dense arrangement of silt particles. [Copyright &y& Elsevier]