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Enhanced Silicate Weathering (ESW) has emerged as a promising Carbon Dioxide Removal (CDR) technology. However, it is still not clear which factors could maximize ESW rates. Previous studies have shown the potential of soil biota to amplify mineral weathering rates. Among the relevant soil biota are earthworms, which have been found to enhance nutrient release from soil minerals. This is indicative of their ability to increase weathering rates and further inorganic carbon (C) sequestration. Here we aim to accelerate ESW rates in a bio-reactor through earthworm activity. First, we identified the optimal conditions for earthworm survival and activity in an environment exclusively composed of ground silicate rocks and organic substrate. Second, we determined to what extent earthworms can enhance inorganic C sequestration in such a system. We carried out 5 rounds of 2-month experiments in a climate chamber at 25°C. The set-up of the experiments consisted of 200 columns, each topped by a sprinkler connected to an irrigation system, which allowed for different water irrigation rates and watering frequencies. The leachate of each column was collected in a jerrycan kept at 4˚C. Within this set-up, we used two endogeic earthworm species (Aporrectodea Caliginosa and Allobophora Chlorotica) at different densities (10, 20, and 30 earthworms kg-1 soil), three types of rock flours (Basalt, Lava and Dunite) of two grain sizes (0.063 and 1.5 mm), one organic source (straw) and two water irrigation rates (125 and 250 ml day-1 kg-1 soil) at three watering frequencies (1, 2 and 5 times/day). Rock flours were used not only as single type or single size, but also as mixtures of types and/or sizes. At the end of each experiment, we measured earthworm survival and activity, and inorganic C sequestration rates by summing cumulative dissolved inorganic C in the leachate and the newly formed solid inorganic C content. We found no differences in survival and activity between the two earthworm species, but we did find an optimum for both parameters at a density of 10 earthworms kg-1 soil. Earthworms showed a clear preference for a mixture of grain sizes compared to single size, and for single mineral type compared to a mixture of mineral types. The response of earthworm survival and activity to the two water irrigation rates was similar, but at a water irrigation rate of 250 ml day-1 kg-1 soil a frequency of 5 times/day resulted in higher activity. Preliminary results indicate that earthworms increase mineral weathering rates and thereby sequester inorganic C. We demonstrate that earthworms can thrive in a fully artificial environment designed to ESW rates, removing one hurdle for designing a bio-reactor aimed at optimizing carbon sequestration.

Flooding threatens the functioning of managed grasslands by decreasing primary productivity and increasing nitrogen losses, notably as the potent greenhouse gas nitrous oxide (N2O). Sowing species with traits that promote flood resilience and mitigate flood-induced N2O emissions within these grasslands could safeguard their productivity while mitigating nitrogen losses. We tested how plant traits and resource acquisition strategies could predict flood resilience and N2O emissions of 12 common grassland species (eight grasses and four legumes) grown in field soil in monocultures in a 14-week greenhouse experiment. We found that grasses were more resistant to flooding while legumes recovered better. Resource-conservative grass species had higher resistance while resource-acquisitive grasses species recovered better. Resilient grass and legume species lowered cumulative N2O emissions. Grasses with lower inherent leaf and root δ13C (and legumes with lower root δ13C) lowered cumulative N2O emissions during and after the flood. Our results highlight the differing responses of grasses with contrasting resource acquisition strategies, and of legumes to flooding. Combining grasses and legumes based on their traits and resource acquisition strategies could increase the flood resilience of managed grasslands, and their capability to mitigate flood-induced N2O emissions. A free Plain Language Summary can be found within the Supporting Information of this article.

Minimizing nitrogen (N) losses and increasing plant N uptake in agroecosystems is a major global challenge. Ecological concepts from (semi)natural grasslands suggest that manipulating plant community composition using plant species with different traits may represent a promising opportunity to face this challenge. Here, we translate these trait-based concepts to agricultural systems in a field experiment, aiming to reveal the main determinants of how plant community composition regulates N-cycling in intensively managed grasslands.We focused on key N pools (plant N from soil and from biological N-fixation, soil mineral N and N2O emissions) as well as on biological drivers of N-cycling in soil (abundance of N-cycling microbial communities, earthworm populations and arbuscular mycorrhizal fungi), using three common grass and one legume species in monoculture, two- and four-species mixtures. We hypothesized that: (a) plant species mixtures increase plant N uptake, reduce soil mineral N concentrations and N2O emissions and promote the abundance of biological N-cyclers; (b) legume presence stimulates N pools, fluxes and biological N-cycling activity, (c) but in combination with a grass with acquisitive traits, more N is retained in the plant community, while N2O emissions are reduced.We found that mixtures increased plant N and lowered the soil mineral N pool compared to monocultures. However, plant species identity played an overarching role: Legume presence increased N2O emissions, plant N pools, soil mineral N and the abundance of N-cycling microbes and earthworms. Combining the legume with a grass with low leaf dry matter content and high root length density (and with high root biomass) reduced the higher soil mineral N and N2O emissions induced by the legume, while harnessing positive effects on plant N pools and biological N-fixation.Synthesis and applications. Our results show the potential of plant community composition to steer N-cycling in fertilized agroecosystems, paving the way for a more biologically based agriculture. Legumes will play a crucial role, but selecting an optimum companion species is key for the sustainability of the agroecosystem.

Microbial necromass is regarded as a central pool of soil organic carbon, whose management is critical in efforts to reduce atmospheric CO2 concentrations and mitigate climate change. However, recent concepts on soil organic matter formation have ignored one of the most important factors for the formation and stabilization of microbial necromass in many soils: earthworms. Based on recent evidence, we conceptualize how the ingestion and mixing of mineral particles and organic matter by earthworms temporarily convert the egested soil to a hotspot of quick and efficient microbial growth and turnover, in which increased amounts of necromass tightly bind to mineral surfaces and stabilize within aggregates. We further stress the low dependence of this process on the quality of pre-existing soil organic matter (in contrast to the assumptions of recent concepts) and its high relevance to the resilience of soil carbon to external disturbances in extensive regions of the soil remote from classical hotspots of microbial necromass formation. We finally provide suggestions on how to close remaining research gaps.

In the search for a more sustainable form of agriculture, a better recycling of major nutrients is essential. For phosphorus (P), one of the most limiting factors to better recycling is chemical adsorption to reactive soil particles, which seriously restricts P supply to plants in many soils. It has been known for some time that earthworms can temporarily increase soil P availability in their casts. However, the exact pathways behind this effect are unclear, making it difficult to infer under which conditions earthworms may significantly contribute to P recycling. In two greenhouse experiments, we studied the occurrence of earthworm-induced increased P availability (i) across a range of common earthworm species; and (ii) across four soils with different physico-chemical characteristics. In the first experiment we analyzed casts of eight common Dutch earthworm species for P pools and related soil properties. For all species, pH in casts was higher than in the bulk soil (up to 1.6 pH unit). Dissolved Organic Carbon (DOC) concentrations were an order of magnitude higher in the casts, and directly available P (defined as water-soluble ortho-phosphate) up to two orders of magnitude. Although these effects were significant for all earthworm species, significant changes were found between the species that could not be explained by conventional earthworm feeding guilds. In the second experiment, we tested effects of three different earthworm species across four soils differing in texture, metal oxide composition, P availabilty and pH. We found a significant effect of earthworms on P availability in all soils, but the extent of this effect varied. Using surface complexation modeling we evaluated the relative importance of the various possible mechanisms. We concluded that the effect of pH on P desorption was relatively small. Increased mineralization of organic P did play an important role; as did competitive desorption of DOC to metal oxides. However, our study also showed a new important pathway: a reduction in reactive surface area of soil metal (hydr)oxides during earthworm gut passage. As this decrease was important in iron (hydr)oxide-dominated soils but not in aluminum (hydr)oxide-dominated soils, we suggest that earthworms have the largest potential to affect soil P availability in the former soils.

Negative Emission Technologies (NETs) are urgently needed if we want to keep global temperature increase below 1.5 °C. Enhanced Silicate Weathering (ESW) is a NET with as yet unknown potential to mitigate climate change. There are indications that ESW rates can be amplified by biotic activity, including that of earthworms. Earlier studies have suggested various pathways through which earthworms might enhance weathering rates, including the grinding of minerals in their gizzard, the stimulation of microbial communities in their gut, as well as the production of mucus rich in organic acids and digestive enzymes. Within this research, we aim to unravel the mechanisms through which earthworms increase mineral weathering rates, and ultimately to develop a bio-reactor in which these processes are optimized. As a first step, we carried out two experiments to determine the suitability of two earthworm species and to establish the optimal conditions for earthworms and mineral weathering in a small bio-reactor. The first study tested the potential of two endogeic earthworm species, Aporrectodea caliginosa and Allolobophora chlorotica, in a system with two types of rock flours (dunite and basalt) and three organic sources (hay, straw and co-digestate solid). The results showed that both earthworm species can thrive and remain active in the bio-reactor in the presence of basalt mixed with either co-digestate or straw. In the second study we tested the tolerance of the same two earthworm species to two temperatures exceeding earthworms ambient levels (20°C and 25°C) and two flow rates (50 ml/day and 80 ml/day) in a system with basalt and straw. The results showed that both earthworm species can survive and remain active at the highest temperature level and the highest flow rate. Our findings demonstrate that earthworms are suitable for use in a bio-reactor and can tolerate physical conditions which are known to stimulate weathering. Future studies will elucidate to what extent earthworms can enhance weathering.

Intensively managed grasslands are large sources of the potent greenhouse gas nitrous oxide (N2O) and important regulators of methane (CH4) consumption and production. The predicted increase in flooding frequency and severity due to climate change could increase N2O emissions and shift grasslands from a net CH4 sink to a source. Therefore, effective management strategies are critical for mitigating greenhouse gas emissions from flood-prone grasslands. We tested how repeated flooding affected the N2O and CH4 emissions from 11 different plant communities (Festuca arundinacea, Lolium perenne, Poa trivialis, and Trifolium repens in monoculture, 2- and 4-species mixtures), using intact soil cores from an 18-month old grassland field experiment in a 4-month greenhouse experiment. To elucidate potential underlying mechanisms, we related plant functional traits to cumulative N2O and CH4 emissions. We hypothesized that traits related with fast nitrogen uptake and growth would lower N2O and CH4 emissions in ambient (non-flooded) conditions, and that traits related to tissue toughness would lower N2O and CH4 emissions in flooded conditions. We found that flooding increased cumulative N2O emissions by 97 fold and cumulative CH4 emissions by 1.6 fold on average. Plant community composition mediated the flood-induced increase in N2O emissions. In flooded conditions, increasing abundance of the grass F. arundinacea was related with lower N2O emissions; whereas increases in abundance of the legume T. repens resulted in higher N2O emissions. In non-flooded conditions, N2O emissions were not clearly mediated by plant traits related with nitrogen uptake or biomass production. In flooded conditions, plant communities with high root carbon to nitrogen ratio were related with lower cumulative N2O emissions, and a lower global warming potential (CO2 equivalent of N2O and CH4). We conclude that plant functional traits related to slower decomposition and nitrogen mineralization could play a significant role in mitigating N2O emissions in flooded grasslands.

Biochar is proposed as an option to sequester carbon (C) in soils and promote other soil-based ecosystem services. However, its impact on soil biota from micro to macroscale remains poorly understood. We investigated biochar effects on the soil biota across the soil food web, on plant community composition and on biomass production. We conducted a field experiment in a nature restoration grassland testing four treatments: two biochar types (herbaceous feedstock pyrolyzed at 400 °C or 600 °C – hereafter B400 and B600), and a positive (i.e. unpyrolysed biochar feedstock, hereafter Hay) and negative (no addition) control. Responses of plants and soil biota were evaluated one and three years after establishing the treatments. Soil pH and K concentrations increased significantly in the B600 treatment. Mite abundances were significantly higher in B400 whereas nematode abundances were highest in Hay (1st year) and lowest in B400 (3rd year). Other soil fauna groups (enchytraeids and earthworms) varied more between years than between treatments. Legume cover increased significantly in the biochar treatments but this effect was transient. Legumes, grasses and primary productivity also showed a statistically significant Treatment x Year interaction due to transitory effects that were no longer present by the 3rd year. Our results suggest that biochar produced from meadow cuttings and applied at the 10 t/ha rate cause transitory impacts on soil biota abundance and plant communities over the 3-year timeframe used for this experiment. Therefore, this type of biochar could potentially be used for soil carbon sequestration, with minimal impacts on soil biota abundance or diversity, within the groups studied here, or plant biodiversity and productivity. Further research is required to investigate the longer-term impacts of this potential soil C storage sink.

Nitrous oxide (N2O) emissions from agricultural soils are the main source of atmospheric N2O, a potent greenhouse gas and key ozone-depleting substance. Several agricultural practices with potential to mitigate N2O emissions have been tested worldwide. However, to guide policymaking for reducing N2O emissions from agricultural soils, it is necessary to better understand the overall performance and variability of mitigation practices and identify those requiring further investigation. We performed a systematic review and a second-order meta-analysis to assess the abatement efficiency of N2O mitigation practices from agricultural soils. We used 27 meta-analyses including 41 effect sizes based on 1119 primary studies. Technology-driven solutions (e.g. enhanced-efficiency fertilizers, drip irrigation, and biochar) and optimization of fertilizer rate have considerable mitigation potential. Agroecological mitigation practices (e.g. organic fertilizer and reduced tillage), while potentially contributing to soil quality and carbon storage, may enhance N2O emissions and only lead to reductions under certain pedoclimatic and farming conditions. Other mitigation practices (e.g. lime amendment or crop residue removal) led to marginal N2O decreases. Despite the variable mitigation potential, evidencing the context-dependency of N2O reductions and tradeoffs, several mitigation practices may maintain or increase crop production, representing relevant alternatives for policymaking to reduce greenhouse gas emissions and safeguard food security.

Microbial necromass is a central component of soil organic matter (SOM), whose management may be essential in mitigating atmospheric CO2 concentrations and climate change. Current consensus regards the magnitude of microbial necromass production to be heavily dependent on the carbon use efficiency of microorganisms, which is strongly influenced by the quality of the organic matter inputs these organisms feed on. However, recent concepts neglect agents relevant in many soils: earthworms. We argue that the activity of earthworms accelerates the formation of microbial necromass stabilized in aggregates and organo-mineral associations and reduces the relevance of the quality of pre-existing organic matter in this process. Earthworms achieve this through the creation of transient hotspots (casts) characterized by elevated contents of bioavailable substrate and the efficient build-up and quick turnover of microbial biomass, thus converting SOM not mineralized in this process into a state more resistant against external disturbances, such as climate change. Promoting the abundance of earthworms may, therefore, be considered a central component of management strategies that aim to accelerate the formation of stabilized microbial necromass in wide locations of the soil commonly not considered hotspots of microbial SOM formation.

The key nutrient phosphorus (P) binds strongly to reactive soil particles, which makes it poorly available for plant uptake. In the search for sustainable ways to overcome a resulting P shortage, it has been shown that earthworms can increase the pool of plant available P and enhance plant P uptake under controlled (greenhouse) conditions. To validate these findings under field conditions and to study the effect of earthworm community composition, we conducted a mesocosm-field experiment on grassland. Mesocosms containing a sandy soil with a low P-status and communities of five earthworm species common to the Netherlands (Lumbricus rubellus, Aporrectodea caliginosa, Allolobophora chlorotica, Lumbricus terrestris and Aporrectodea longa; monocultures, three- or five-species mixtures and controls without earthworms) were installed in a field. Aboveground biomass production and P uptake of Lolium perenne were monitored for over two years. Earthworm community composition varied between the start and the end of the experiment, but multiple linear regression on the final earthworm communities yielded strong indications that earthworms increased both biomass production (R2adj = 0.52, p < 0.001, n = 76) and P uptake (R2adj = 0.48, p < 0.001, n = 76). The species A. longa and L. terrestris were most influential for this earthworm effect. We did not observe any general relations between the number of earthworm species in a community and a P effect. Our results suggest that earthworms can indeed increase grass biomass production and P uptake on a low P soil in the field, and can thereby contribute to making the P nutrition of our agricultural systems more sustainable.

Reactive nitrogen (N) losses, and in particular nitrous oxide losses, from dairy grasslands on peat soils are generally high as a result of relative high soil organic matter contents, potential N mineralisation rates and shallow groundwater levels. Effects of the inclusion of the temperate forage species plantain (Plantago lanceolata L.) (PL), which produces secondary compounds with biological nitrification inhibition capacity, on the fate of soil mineral N were studied in a combined mesocosm and field experiment. The experiments comprised four treatments differing in intentional herbage share of plantain versus perennial ryegrass (Lolium perenne L.) (100%PL, 66%PL, 33%PL and 0%PL). Potential nitrification in the mesocosm experiment was significantly lower at 100%PL versus 0%PL (p = 0.018), but soil nitrate concentrations were not. Nitrous oxide fluxes reduced by 39% (p = 0.021) in the presence of plantain in the field experiment, without an obvious link to the quantity of plantain. N use efficiency of plantain tended to increase with the quantity of plantain in the sward in the mesocosm experiment (p = 0.098), but not in the field experiment. Our results suggest that the presence of plantain can affect the fate of soil mineral N of dairy grasslands on peat soils.

International audience; A number of negative emission technologies (NETs) have been proposed to actively remove CO2 from the atmosphere, with enhanced silicate weathering (ESW) as a relatively new NET with considerable climate change mitigation potential. Models calibrated to ESW rates in lab experiments estimate the global potential for inorganic carbon sequestration by ESW at about 0.5–5 Gt CO2 year−1, suggesting ESW could be an important component of the future NETs mix. In real soils, however, weathering rates may differ strongly from lab conditions. Research on natural weathering has shown that biota such as plants, microbes, and macro-invertebrates can strongly affect weathering rates, but biotic effects were excluded from most ESW lab assessments. Moreover, ESW may alter soil organic carbon sequestration and greenhouse gas emissions by influencing physicochemical and biological processes, which holds the potential to perpetuate even larger negative emissions. Here, we argue that it is likely that the climate change mitigation effect of ESW will be governed by biological processes, emphasizing the need to put these processes on the agenda of this emerging research field.

Sustainability of agricultural systems is at stake, as phosphorus (P) is a non-renewable resource while its global reserves are limited. Stimulating earthworm activity can be a technology to increase the level of readily plant-available phosphate (PO$_4$). However, conclusive evidence on the mechanisms underlying an earthworm-enhanced PO$_4$ solubility is yet missing. This study aimed to reveal possibly overlooked pathways by which earthworms affect PO$_4$ solubility, and quantify the relative importance of all contributing mechanisms. Therefore, we set up a greenhouse pot experiment in which we investigated the large increase in water-extractable PO$_4$ in casts of three earthworm species (Lumbricus rubellus, Aporrectodea caliginosa, Lumbricus terrestris) in soils with either predominantly Fe- or Al-(hydr)oxides. Oxalate-extractable PO$_4$ was increased in earthworm casts compared to bulk soil which can be attributed to the mineralisation of natural organic matter (NOM). Surface complexation modelling was used to elucidate the mechanisms that control earthworm-enhanced PO$_4$ solubility. The results of our modelling showed that the increase in pH in earthworm casts relative to bulk soil affects PO$_4$ solubility only to a minor extent. Besides NOM mineralisation, two major mechanisms contributing to earthworm-enhanced PO$_4$ solubility are (i) a decrease in the reactive surface area (RSA) of the metal-(hydr)oxide fraction; and (ii) a decrease in the competition between NOM and PO$_4$ for binding sites of the metal-(hydr)oxides. As the newly discovered decrease of the RSA was only found for Fe-(hydr)oxide-dominated soils, earthworms have the largest potential to enhance PO$_4$ solubility in those soils.

It has long been established that earthworms beneficially affect plant growth. This is to a large extent due to the high fertility of their casts. However, it is not clear how fertile casts are compared to bulk soil, and how their fertility varies between earthworm feeding guilds and with physico-chemical soil properties. Using meta-analysis, we quantified the fertility of earthworm casts and identified its controlling factors. Our analysis included 405 observations from 81 articles, originating from all continents except Antarctica. We quantified cast fertility by determining the enrichment of earthworm casts relative to the bulk soil (“relative cast fertility”; RCF) for total organic carbon (TOC), total phosphorus (P) and total nitrogen (N) concentrations, as well as for plant available pools of N (total mineral N) and P (available P: P-Olsen, P-Bray or comparable metrics), C-to-N ratio and microbial biomass C. In addition to these response variables, we studied four additional ones closely related to soil fertility: pH-H2O, clay content, cation exchange capacity (CEC), and base saturation. With the exception of C-to-N ratio, microbial C and clay content, all studied response variables were significantly increased in casts compared to the bulk soil. Increases in total elemental concentrations (TOC, total P and total N), which are the result of preferential feeding or concentration processes, were comparable and ranged between 40 and 48%. Nutrient availability, which is to a large extent the result of (bio)chemical transformation processes in the earthworm gut, was increased more strongly than total elemental concentrations (241% and 84% for mineral N and available P, respectively). Increases in pH (0.5 pH units), cation exchange capacity (40%), and base saturation (27%) were also large and significant. None of the soil-related possible controlling factors could satisfactorily explain the variation in RCF; plant presence (or other sources of organic C input such as residue application) was the only controlling factor that consistently increased RCF across soil properties. With the exception of available P, none of the studied response variables could be linked to earthworm feeding guild. Our results show that earthworm casts are much more fertile than bulk soil for almost all analysed cast fertility properties. However, these positive RCFs are to a large extent dependent upon the presence of plants. In general, earthworm feeding guild or specific physico-chemical soil properties could not explain the large variability in RCF for the various response variables. Therefore, we hypothesize that RCF effects depend on intricate interactions between earthworm species traits and specific soil properties. Understanding these interactions requires trait-based approaches combined with mechanistic modelling of biochemical processes in the earthworm gut and casts.

How much C can be stored in agricultural soils worldwide to mitigate rising carbon dioxide (CO2) concentrations, and at what cost? This question, because of its critical relevance to climate policy, has been a focus of soil science for decades. The amount of additional soil organic C (SOC) that could be stored has been estimated in various ways, most of which have taken the soil as the starting point: projecting how much of the SOC previously lost can be restored, for example, or calculating the cumulative effect of multiple soil management strategies. Here, we take a different approach, recognizing that photosynthesis, the source of C input to soil, represents the most fundamental constraint to C sequestration. We follow a simple “Fermi approach” to derive a rough but robust estimate by reducing our problem to a series of approximate relations that can be parameterized using data from the literature. We distinguish two forms of soil C: ‘ephemeral C’, denoting recently-applied plant-derived C that is quickly decayed to CO2, and ‘lingering C,’ which remains in the soil long enough to serve as a lasting repository for C derived from atmospheric CO2. First, we estimate global net C inputs into lingering SOC in croplands from net primary production, biomass removal by humans and short-term decomposition. Next, we estimate net additional C storage in cropland soils globally from the estimated C inputs, accounting also for decomposition of lingering SOC already present. Our results suggest a maximum C input rate into the lingering SOC pool of 0.44 Pg C yr−1, and a maximum net sequestration rate of 0.14 Pg C yr−1 – significantly less than most previous estimates, even allowing for acknowledged uncertainties. More importantly, we argue for a re-orientation in emphasis from soil processes towards a wider ecosystem perspective, starting with photosynthesis.

Sustainable agricultural practices aim to ensure the rebuilt of soil organic carbon (SOC) stocks and to sustain soil fertility. One of the levers is the use of carbon and nutrient inputs in the form of organic amendments, such as farmyard manure, slurry and biogas digestate. These organic fertilizers represent a promising alternative to the mineral fertilizers, which are mainly made from non-renewable resources. The use mineral fertilizers is indeed associated with an excessive use of natural resources and a loss of biodiversity. The effect of organic amendments compared with traditional mineral fertilizers on SOC stocks and soil fertility are uncertain in the longer-term. We aimed at investigating the effects of mineral and organic fertilizers (i.e., manure, pig slurry and biogas digestate) on topsoil and subsoil biogeochemistry, after eight years of application. For this purpose, we sampled soil cores down to a depth of one meter in a randomized field experiment in Germany, running since 2011. A full-profile assessment of the carbon and nitrogen distribution, stability and bioavailability was achieved using a combination of classical bulk physico-chemical analyses (e.g., SOC and nitrogen contents, texture, pH, bulk density) and state-of-the-art imaging techniques. Selected samples were analysed for aggregate size distribution, as well as organic carbon and nitrogen contents and allocation within these aggregates. Further, undisturbed core-samples were scanned using a hyperspectral camera in the Vis-NIR range to reveal hotspots of carbon storage at the soil profile scale. Soil carbon distribution was predicted as a function of spectral response, using a variety of machine learning approaches. The application of organic fertilizers (whatever their nature) resulted in higher SOC contents in the first 10 cm, as compared to the control and the mineral fertilizer treatments. The SOC stocks were + 21-33 % higher in the soil treated with organic fertilizers as compared to the control treatment. The application of mineral fertilizer or digestate, as compared to the control, resulted in higher relative amount of microaggregates (versus macroaggregates) (+ 19-40 %) in the soil down to 80 cm. These results will provide essential information to develop management strategies that increase nutrient recycling as well as SOC stocks.

Increased human‐derived nitrogen (N) deposition to terrestrial ecosystems has resulted in widespread phosphorus (P) limitation of net primary productivity. However, it remains unclear if and how N‐induced P limitation varies over time. Soil extracellular phosphatases catalyze the hydrolysis of P from soil organic matter, an important adaptive mechanism for ecosystems to cope with N‐induced P limitation. Here we show, using a meta‐analysis of 140 studies and 668 observations worldwide, that N stimulation of soil phosphatase activity diminishes over time. Whereas short‐term N loading (≤5 years) significantly increased soil phosphatase activity by 28%, long‐term N loading had no significant effect. Nitrogen loading did not affect soil available P and total P content in either short‐ or long‐term studies. Together, these results suggest that N‐induced P limitation in ecosystems is alleviated in the long‐term through the initial stimulation of soil phosphatase activity, thereby securing P supply to support plant growth. Our results suggest that increases in terrestrial carbon uptake due to ongoing anthropogenic N loading may be greater than previously thought. This study was funded by Aarhus University Centre for Circular Bioeconomy, Aarhus University Research Foundation AUFF Starting Grants (AUFF-E-2019-7-1), and Marie Skłodowska-Curie Individual Fellowship H2020-MSCA-IF-2018 (no. 839806). Ji Chen acknowledges funding support from the National Natural Science Foundation of China (41701292) and China Postdoctoral Science Foundation (2017M610647, 2018T111091) when constructing the databases. César Terrer was supported by a Lawrence Fellow award through Lawrence Livermore National Laboratory (LLNL). This work was performed under the auspices of the U.S. Department of Energy by LLNL under contract DEAC52-07NA27344 and was supported by the LLNL-LDRD Program under Project No. 20-ERD-055. Fernando T. Maestre was supported by the European Research Council (ERC Grant agreement 647038 [BIODESERT]) and Generalitat Valenciana (CIDEGENT/2018/041).

The increasing frequency of extreme weather events, such as floods, requires management strategies that promote resilience of grassland productivity. Mixtures of plant species may better resist and recover from flooding than monocultures, as they could combine species with stress-coping and resource acquisition traits. This has not yet been tested in intensively managed grasslands despite its relevance for enhancing agroecosystem resilience. Using intact soil cores from an 18-month-old field experiment, we tested how 11 plant communities (Festuca arundinacea, Lolium perenne, Poa trivialis and Trifolium repens in monoculture, two- and four-species mixtures) resist and recover from repeated flooding in a 4-month greenhouse experiment. We found that plant community composition, not whether the community was a mixture or monoculture, influenced the community's resistance to flooding, although most communities were able to resist and recover from both floods. The plant community's position on the leaf economic spectrum in flooded conditions was related to its resistance to and recovery from flooding. Resistance to and recovery from a severe flood were related to flood-induced intraspecific trait variation, causing a shift in the community's position on the leaf resource economic spectrum. In flooded conditions, resource-conservative communities (characterized by low specific leaf area, low leaf nitrogen content and high leaf dry matter content) better resisted and recovered from flooding. The community's position on the root resource economic spectrum was less connected to the community's resistance and recovery. Synthesis and applications. Our study shows that in flooded conditions, resource-conservative plant communities are more resilient to flooding than resource-acquisitive communities in an intensively managed grassland. This suggests that plant community position on the leaf economic spectrum, as well as species’ flood-induced intraspecific variation, should be considered when designing grasslands to withstand increasing flood frequency and severity.

Studies on soil organic carbon (SOC) stocks mostly focus on topsoils (< 30 cm). However, 30 to 63% of the SOC are stored in the subsoils (30 to 100 cm), and the factors controlling SOC storage in subsoils may be substantially different than in topsoils. The low mean SOC content in subsoils makes its quantification and characterization challenging. Thus, new approaches are required to depict the SOC stocks distribution in full soil profile. Hyperspectral imaging of soil core samples can provide high spatial resolution of the vertical distribution of SOC in a soil profile. The main objective of the ongoing study, within the Horizon 2020 European Project Circular Agronomics, is to apply laboratory hyperspectral imaging with a variety of machine learning approaches for the mapping of OC distribution in undisturbed soil cores. Soil cores were collected down to a depth of one meter in grasslands of 15 organic farms located in the Lungau Valley, in Austria. Some samples were divided into five depths in the field for classical bulk soil measurements (total carbon and nitrogen, texture, pH, EC and bulk density) on disturbed samples. Undisturbed soil cores were sliced vertically for laboratory hyperspectral imaging in the range of Vis-NIR (400-1000 nm). We were able to reveal the hotspots of OC and map the OC distribution in soil profile by applying a variety of machine learning approaches (i.e. partial least square and random forest regression) as a function of spectral responses. A digital elevation model was further exploited to investigate the effects of topographical factors such as elevation, aspect and slope on SOC profile distribution. Landsat 8 data were also used to depict the spatial variability of land insensitive cover/vegetation in study area.

Soil faunal activity can be a major control of greenhouse gas (GHG) emissions from soil. Effects of single faunal species, genera or families have been investigated, but it is unknown how soil fauna diversity may influence emissions of both carbon dioxide (CO2, end product of decomposition of organic matter) and nitrous oxide (N2O, an intermediate product of N transformation processes, in particular denitrification). Here, we studied how CO2 and N2O emissions are affected by species and species mixtures of up to eight species of detritivorous/fungivorous soil fauna from four different taxonomic groups (earthworms, potworms, mites, springtails) using a microcosm set‐up. We found that higher species richness and increased functional dissimilarity of species mixtures led to increased faunal‐induced CO2 emission (up to 10%), but decreased N2O emission (up to 62%). Large ecosystem engineers such as earthworms were key drivers of both CO2 and N2O emissions. Interestingly, increased biodiversity of other soil fauna in the presence of earthworms decreased faunal‐induced N2O emission despite enhanced C cycling. We conclude that higher soil fauna functional diversity enhanced the intensity of belowground processes, leading to more complete litter decomposition and increased CO2 emission, but concurrently also resulting in more complete denitrification and reduced N2O emission. Our results suggest that increased soil fauna species diversity has the potential to mitigate emissions of N2O from soil ecosystems. Given the loss of soil biodiversity in managed soils, our findings call for adoption of management practices that enhance soil biodiversity and stimulate a functionally diverse faunal community to reduce N2O emissions from managed soils., We tested the relation between diversity of soil fauna and greenhouse gas emissions. We found that higher faunal diversity decreased emissions of the potent greenhouse gas nitrous oxide. Our results suggest that maintaining soil biodiversity is crucial to reduce soil greenhouse gas emissions

Flooded rice production is crucial to global food security, but there are associated environmental concerns. In particular, it is a significant source of methane (CH4) and nitrous oxide (N2O) emissions and a large consumer of water resources, while arsenic levels in the grain are a serious health concern. There is also a tendency to use more organic fertilisers to close nutrient cycles, posing a threat of even higher GHG emissions and grain arsenic levels. It has been shown that alternate wetting and drying (AWD) water management reduces both water use and GHG emissions, but success at maintaining yields varies. This study tested the effect of early AWD (e-AWD) versus continuous flooding (CF) water management practices on grain yields, GHG emissions and grain arsenic levels in a split-plot field experiment with organic fertilisers under organic management. The treatments included: i) farmyard manure, ii) compost, and iii) biogas digestate, alone or in combination with mineral fertiliser. The e-AWD water regime showed no difference in yield for the organic treatments. Yields significantly increased by 5–16 % in the combination treatments. Root biomass and length increased in the e-AWD treatments up to 72 and 41 %, respectively. The e-AWD water regime reduced seasonal CH4 emissions by 71–85 % for organic treatments and by 51–76 % for combination treatments; this was linked to a 15–47 % reduction in dissolved organic carbon (DOC), thereby reducing methanogenesis. N2O emissions increased by 23–305 % but accounted for

Because phosphorus (P) is one of the most limiting nutrients in agricultural systems, P fertilisation is essential to feed the world. However, declining P reserves demand far more effective use of this crucial resource. Here, we use meta-analysis to synthesize yield responses to P fertilisation in grasslands, the most common type of agricultural land, to identify under which conditions P fertilisation is most effective. Yield responses to P fertilisation were 40–100% higher in (a) tropical vs temperate regions; (b) grass/legume mixtures vs grass monocultures; and (c) soil pH of 5–6 vs other pHs. The agronomic efficiency of P fertilisation decreased for greater P application rates. Moreover, soils with low P availability reacted disproportionately strong to fertilisation. Hence, low fertiliser application rates to P-deficient soils result in stronger absolute yield benefits than high rates applied to soils with a higher P status. Overall, our results suggest that optimising P fertiliser use is key to sustainable intensification of agricultural systems.

Earthworm activity can strongly influence soil structure and organic matter (OM) dynamics of agricultural soils. Several short-term studies (≤90 days) have shown that earthworms can increase incorporation of residue carbon (C) into soil aggregates, suggesting reduced decomposition in the longer term. In contrast, another body of short-term studies reported that earthworms can increase carbon dioxide (CO2) emission from soils, thus suggesting increased decomposition in the longer term instead. To solve this controversy, we measured the effect of the epigeic Lumbricus rubellus (Hoffmeister) and the endogeic Aporrectodea caliginosa (Savigny) on the soil C balance in a unique 750-day mesocosm experiment, where loess soil was surface-applied with maize (Zea mays L.) residues every six months. Carbon inputs and outputs were strictly controlled: no soil C input through growing plants and no leaching of soil organic C. Flux measurements of CO2 were taken regularly and aggregate size distribution and total C and residue-derived C in the aggregate fractions (using the natural δ13C signature of maize) were measured after 185, 565 and 750 days. Both earthworm species increased cumulative CO2 emissions by at least 25%, indicating a higher C loss compared to the no-earthworm control. Yet, both earthworm species also increased the amount of soil C associated with the macroaggregate fraction after 750 days. L. rubellus increased the amount of residue-derived C in the macroaggregate fraction after 565 and after 750 days, whereas A. caliginosa increased residue-derived C in all the measured soil fractions after 750 days. Our results show that earthworms can simultaneously enhance CO2 emissions and C incorporation in aggregate fractions. However, over 750 days the presence of earthworms resulted in a lower total C content due to a higher overall OM decomposition rate. We therefore propose that under the most realistic incubation conditions so far (longer term and multiple residue applications), earthworms stimulate the mineralization of freshly added and older OM to a greater extent than that they stabilize residue-derived C inside biogenic aggregates. Future studies should focus on the balance between these processes in the presence of growing plants.

It is estimated that only 20% of fertilizers applied annually in the European agricultural systems are converted to finished products for human consumption. These low efficiencies result in large loss of nutrients into the environment with severe negative influences on soils, water and air, and constitute unacceptable health and environmental costs. In addition, around 45% of soils in the European countries have less than 2% organic carbon (OC). Low soil OC storage is linked with negative environmental impacts including soil and water quality, climate change and biodiversity. A relevant strategy to enhance soil OC is the transformation of waste products into organic amendments for application on soils. The aim of the H2020 European project Circular Agronomics is to provide a comprehensive synthesis of practical solutions to improve the current C, N and P cycling in European agro-ecosystems. This project explores the medium and long-term effects of new and classical organic fertilizers in six countries including Germany, Spain, Italy, Netherlands, Czech Republic and Austria. The study sites will be sampled before and after applying the new organic amendments using a hydraulic corer. A full profile assessment of the C, N and P distribution, stability and bioavailability will be released up to one meter depth using a combination of classical bulk chemical analyses and state-of-the-art imaging techniques. Undisturbed soil cores will be scanned using a hyperspectral camera to reveal hotspots of C, N and P storage in the soil profile, at the micro-scale. Soil C, N and P will be modelled as a function of spectral response using a variety of machine learning approaches. These results will provide essential information to develop management strategies that reduce nutrient surplus and increase C stocks.

Tropical peatlands in the Peruvian Amazon exhibit high densities of Mauritia flexuosa palms, which are often cut instead of being climbed for collecting their fruits. This is an important type of forest degradation in the region that could lead to changes in the structure and composition of the forest, quality and quantity of inputs to the peat, soil properties, and greenhouse gas (GHG) fluxes. We studied peat and litterfall characteristics along a forest degradation gradient that included an intact site, a moderately degraded site, and a heavily degraded site. To understand underlying factors driving GHG emissions, we examined the response of in vitro soil microbial GHG emissions to soil moisture variation, and we tested the potential of pneumatophores to conduct GHGs in situ. The soil phosphorus and carbon content and carbon-to-nitrogen ratio as well as the litterfall nitrogen content and carbon-to-nitrogen ratio were significantly affected by forest degradation. Soils from the degraded sites consistently produced more carbon dioxide (CO2) than soils from the intact site during in vitro incubations. The response of CO2 production to changes in water-filled pore space (WFPS) followed a cubic polynomial relationship with maxima at 60–70% at the three sites. Methane (CH4) was produced in limited amounts and exclusively under water-saturated conditions. There was no significant response of nitrous oxide (N2O) emissions to WFPS variation. Lastly, the density of pneumatophore decreased drastically as the result of forest degradation and was positively correlated to in situ CH4 emissions. We conclude that recurrent M. flexuosa harvesting could result in a significant increase of in situ CO2 fluxes and a simultaneous decrease in CH4 emissions via pneumatophores. These changes might alter long-term carbon and GHG balances of the peat, and the role of these ecosystems for climate change mitigation, which stresses the need for their protection.

Phosphorus (P) is an important nutrient for plant growth. However, P is often poorly available for uptake by roots because it strongly adsorbs to the soil mineral phase. Recent research shows that earthworms can temporally and locally increase P availability to plants. However, the pathways through which they do so are not fully understood, and it remains unclear to what extent this capacity varies among earthworm species. Here we study the variation among earthworm species with respect to readily-available P in casts as well as other physico-chemical cast properties, in a greenhouse pot experiment using a soil with a low P status. The earthworms belong to eight commonly occurring earthworm species in the Netherlands: two epigeic species (a mixture of the compost earthworms Dendrobaena veneta/Eisenia fetida; Lumbricus rubellus); four endogeic species (Allolobophora chlorotica, Aporrectodea caliginosa, Aporrectodea rosea, Octolasion lacteum); and two anecic species (Aporrectodea longa; Lumbricus terrestris). For all species, the pH in water extracts of earthworm cast (pH = 7.4–8.2) was significantly higher (p < 0.001) than for the control bulk soil (pH = 6.6) and differed significantly (p = 0.003) among earthworm species. Similarly, the dissolved organic carbon (DOC) concentration in the same water extracts was an order of magnitude higher in earthworm cast compared to the control bulk soil and varied among species (p < 0.001). The size of the total pool of reversibly adsorbed P in earthworm cast was greater than in the control bulk soil, but no significant differences were found among earthworm species. Differences among species were present for the readily-available P pools extracted from casts, including P-Olsen and water-extractable ortho-P. Water-extractable ortho-P concentrations were much higher in the casts of all species as compared to the control bulk soil (0.9–6.8 vs 0.06 mg l−1 or to 9.0–68 vs 0.6 mg kg−1). Highest ortho-P levels were measured in L. rubellus casts and the lowest in casts of A. chlorotica. A positive correlation between the concentrations of DOC and ortho-P was observed (R2 = 0.72, p < 0.001). The observed variation in all measured physico-chemical cast properties could not be explained by conventional ecological earthworm classifications. Our results show that the nature and magnitude of earthworm-induced increased P availability differs dramatically among earthworm species. This strongly suggests that, apart from its size, species composition of the earthworm community is key to optimizing P availability to plants.

Rice production systems are the largest anthropogenic wetlands on earth and feed more than half of the world's population. However, they are also a major source of global anthropogenic greenhouse gas (GHG) emissions. Several agronomic strategies have been proposed to improve water-use efficiency and reduce GHG emissions. The aim of this study was to evaluate the impact of water-saving irrigation (alternate wetting and drying (AWD) vs. soil water potential (SWP)), contrasting land establishment (puddling vs. reduced tillage) and fertiliser application methods (broadcast vs. liquid fertilisation) on water-use efficiency, GHG emissions and rice yield. The experiment was laid out in a randomised complete block design with eight treatments (all combinations of the three factors) and four replicates. AWD combined with broadcasting fertilisation was superior to SWP in terms of maintaining yield. However, seasonal nitrous oxide (N2O) emissions were significantly reduced by 64% and 66% in the Broadcast-SWP and Liquid fertiliser-SWP treatments, respectively, compared to corresponding treatments in AWD. The SWP also significantly reduced seasonal methane (CH4) emissions by 34 and 30% in the broadcast and liquid fertilisation treatments, respectively. Area-scaled GWPs were reduced by 48% and 54% in Broadcast-SWP and Liquid fertiliser-SWP treatments respectively compared to the corresponding treatments in AWD. Compared to AWD, the broadcast and liquid fertilisation in SWP irrigation treatments reduced yield-scaled GWPs by 46% and 37%, respectively. In terms of suitability, based on yield-scaled GWPs, the treatments can be ordered as follows: Broadcast-SWP < Broadcast-AWD = Liquid fertiliser-SWP < Liquid fertiliser-AWD. Growing-season water use was 15% lower in the SWP treatments compared with the water-saving AWD. Reduced tillage reduced additional water use during land preparation. The conclusions of this study are that improved water management and timely coordination of N fertiliser with crop demand can reduce water use, N loss via N2O emissions, and CH4 emissions.

Global rice production systems face two opposing challenges: the need to increase production to accommodate the world's growing population while simultaneously reducing greenhouse gas (GHG) emissions. Adaptations to drainage regimes are one of the most promising options for methane mitigation in rice production. Whereas several studies have focused on mid-season drainage (MD) to mitigate GHG emissions, early-season drainage (ED) varying in timing and duration has not been extensively studied. However, such ED periods could potentially be very effective since initial available C levels (and thereby the potential for methanogenesis) can be very high in paddy systems with rice straw incorporation. This study tested the effectiveness of seven drainage regimes varying in their timing and duration (combinations of ED and MD) to mitigate CH4 and N2O emissions in a 101-day growth chamber experiment. Emissions were considerably reduced by early-season drainage compared to both conventional continuous flooding (CF) and the MD drainage regime. The results suggest that ED + MD drainage may have the potential to reduce CH4 emissions and yield-scaled GWP by 85–90% compared to CF and by 75–77% compared to MD only. A combination of (short or long) ED drainage and one MD drainage episode was found to be the most effective in mitigating CH4 emissions without negatively affecting yield. In particular, compared with CF, the long early-season drainage treatments LE + SM and LE + LM significantly (p < 0.01) decreased yield-scaled GWP by 85% and 87% respectively. This was associated with carbon being stabilised early in the season, thereby reducing available C for methanogenesis. Overall N2O emissions were small and not significantly affected by ED. It is concluded that ED + MD drainage might be an effective low-tech option for small-scale farmers to reduce GHG emissions and save water while maintaining yield.

Grasslands are a major form of agricultural land use worldwide. Current and future declines of phosphorus (P) inputs into production grasslands necessitate a shift towards selecting grass species based on high efficiency under suboptimal, rather than optimal P conditions. It is therefore imperative to identify key root traits that determine P acquisition of grasses in soils with a low P status. In a 9-month greenhouse experiment, we grew eight common grass species and cultivars on a soil with a low P status and related root morphological traits to their performance under P-limiting conditions. We applied (P1) or withheld (P0) P fertilization while providing adequate amounts of all other nutrients. Omitting P fertilization greatly reduced yield and nutrient acquisition for the various grass species. Biomass production differed significantly (P < 0.1%) among species and P fertilization treatments, varying from 17.1 to 72.1 g pot−1 in the P0 treatment and from 33.4 to 85.8 g pot−1 in the P1 treatment. Root traits were species-specific and unresponsive to P fertilization, but overall we observed a trade-off between root biomass and specific root length. Structural equation modeling identified total root length as key factor with respect to resistance to P deficiency, especially when roots explored the subsoil. Optimizing root length and subsoil exploration could be the key to maintaining high productivity of production grasslands with decreasing P availability. This is relevant for both plant breeding programs and for composing seed mixtures.

Sampling and analysis or visual examination of soil to assess its status and use potential is widely practiced from plot to national scales. However, the choice of relevant soil attributes and interpretation of measurements are not straightforward, because of the complexity and site-specificity of soils, legacy effects of previous land use, and trade-offs between ecosystem services. Here we review soil quality and related concepts, in terms of definition, assessment approaches, and indicator selection and interpretation. We identify the most frequently used soil quality indicators under agricultural land use. We find that explicit evaluation of soil quality with respect to specific soil threats, soil functions and ecosystem services has rarely been implemented, and few approaches provide clear interpretation schemes of measured indicator values. This limits their adoption by land managers as well as policy. We also consider novel indicators that address currently neglected though important soil properties and processes, and we list the crucial steps in the development of a soil quality assessment procedure that is scientifically sound and supports management and policy decisions that account for the multi-functionality of soil. This requires the involvement of the pertinent actors, stakeholders and end-users to a much larger degree than practiced to date.

This is the author accepted manuscript. The final version is available from American Chemical Society via the DOI in this record.

Background and aimBiochar application to soil is widely claimed to increase plant productivity. However, the underlying mechanisms are still not conclusively described. Here, we aim to elucidate these mechanisms using stable isotope probing.Methods We conducted two experiments with uniquely double-labelled (15N and 13C) biochar and its feedstock (residue), applied separately at 15 Mg ha−1. Both experiments contained three treatments: biochar amendment (Biochar), unpyrolysed residue amendment (Residue) and a no addition control (Control). Experiment I was a 119 day pot experiment seeded with Lolium perenne. Experiment II was a 71 day incubation experiment without plants in which CO2 and N2O fluxes were measured.Results Both Biochar and Residue significantly increased aboveground productivity compared to Control (140% and 160%, respectively). Initial N immobilisation was stimulated in Residue, whereas not in Biochar. 13C–CO2 analysis confirmed that biochar was significantly more recalcitrant than residue. 15N analysis showed that 2% and 0.3% of grass N was derived from the amended material in Residue and Biochar, respectively.ConclusionsOur results suggest that biochar-induced yield increases derive from a combination of reduced N immobilization and a moderate N fertilization effect. Although in the short term biochar might offer benefits compared to residue incorporation, it is unlikely that biochar yield gains will be sustainable for the decades to centuries that biochar C can be expected to reside in soil.

This is the author accepted manuscript. The final version is available from American Chemical Society via the DOI in this record.

Applying biochar to soil is thought to have multiple benefits, from helping mitigate climate change [1, 2], to managing waste [3] to conserving soil [4]. Biochar is also widely assumed to boost crop yield [5, 6], but there is controversy regarding the extent and cause of any yield benefit [7]. Here we use a global-scale meta-analysis to show that biochar has, on average, no effect on crop yield in temperate latitudes, yet elicits a 25% average increase in yield in the tropics. In the tropics, biochar increased yield through liming and fertilization, consistent with the low soil pH, low fertility, and low fertilizer inputs typical of arable tropical soils. We also found that, in tropical soils, high-nutrient biochar inputs stimulated yield substantially more than low-nutrient biochar, further supporting the role of nutrient fertilization in the observed yield stimulation. In contrast, arable soils in temperate regions are moderate in pH, higher in fertility, and generally receive higher fertilizer inputs, leaving little room for additional benefits from biochar. Our findings demonstrate that the yield-stimulating effects of biochar are not universal, but may especially benefit agriculture in low-nutrient, acidic soils in the tropics. Biochar management in temperate zones should focus on potential non-yield benefits such as lime and fertilizer cost savings, greenhouse gas emissions control, and other ecosystem services.

As many soils are unable to supply sufficient amounts of phosphorus (P) to plants, P availability is often a growth-limiting factor. Literature shows that levels of readily available P can be considerably higher in earthworm casts than in the surrounding bulk soil, possibly resulting in increased plant P uptake. However, the underlying mechanisms through which this increase of readily available P occurs are unclear. In a greenhouse pot experiment with annual ryegrass (Lolium multiflorum) on a soil with low P availability, we tested whether the presence of the anecic earthworm Lumbricus terrestris resulted in increased grass growth and P uptake and explored the possible mechanisms behind such an effect. During the pot experiment, earthworm casts were collected and analysed for water-extractable P as well as for most other relevant ions. The earthworm's presence significantly increased grass yield from 9.80 to 10.80 g dry matter per pot (p = 0.044) and P uptake from 6.32 to 8.04 mg per pot (p = 0.002). Due to increased microbial activity, the solution chemistry in the water extracts of the casts was dramatically changed compared to the bulk soil samples. The concentrations of dissolved inorganic P in cast were enhanced by a factor ~ 30–1000 compared to the levels found in bulk soil. The pH in earthworm casts increased from 5.9 to values between 7.4 and 9.0 and the Ca concentrations decreased by about a factor ~ 2-3 compared to bulk soil samples. In addition, there was an increase in dissolved organic carbon (35 mg L− 1 in casts versus 3 mg L− 1 in soil). Surface complexation modelling, using the Charge Distribution model, suggests that, besides an increase in pH, particularly an enhanced interaction of dissolved organic matter with reactive surface of metal (hydr)oxides is likely to be a major driving force for the augmented release of orthophosphate (PO4) via competitive adsorption and desorption. This competition for adsorption sites is an alternative pathway, next to mineralisation of organic matter, through which earthworms can increase soil fertility.

Recent research has shown that microbial-feeding invertebrate soil fauna species can significantly contribute to N2O emissions. However, in soil food webs microbial-feeding soil fauna interact with each other and with their predators, which affects microbial activity. To date we lack empirical tests of whether or not these interactions play a significant role in N2O emissions from soil. Therefore we studied how interactions between soil microbes, two groups of microbial-feeding soil fauna (enchytraeids and fungivorous mites) and their predators (predatory mites) affect soil N2O emissions. We hypothesized that: 1) the presence of two microbial-feeding fauna groups (enchytraeids and fungivorous mites) together increase N2O emissions more than when only a single group is present; and 2) the addition of predatory mites further enhances N2O emissions. We assembled soil food webs consisting of soil microbes, enchytraeids, fungivorous and predatory mites in microcosms with sandy loamy soil and sterilised hay as a substrate for the soil microbes. N2O emissions were measured during 56 days. We found no support for our first yet support for our second hypothesis. Addition of predatory mites to microcosms with enchytraeids and fungivorous mites increased N2O emissions significantly from 135.3 to 482.1 mg N m-2, which was also significantly higher than the control without fauna (83 mg N m-2) (P