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In this study, 6:2 and 8:2 polyfluoroalkyl phosphate diester (diPAP) were individually investigated in lysimeters under near-natural conditions. Leachate was sampled for 2 years, as was the soil after the experiment. In the leachate of the diPAP spiked soils, perfluorocarboxylic acids (PFCAs) of different chain lengths were detected [23.2% (6:2 diPAP variant) and 20.8% (8:2 diPAP variant) of the initially applied molar amount]. After 2 years, the soils still contained 36-37% 6:2 diPAP and 41-45% 8:2 diPAP, respectively, in addition to smaller amounts of PFCAs (1.5 and 10.6%, respectively). Amounts of PFCAs found in the grass were low (

Soil respiration is an important pathway of soil organic carbon losses in temperate grasslands; however, it is rarely studied across broad management intensity gradients in a landscape. Using the soda-lime method, we measured in-situ soil CO2 efflux with single measurements of long exposure time (i.e. 3 day long) in 150 grasslands in three German regions in early summer 2018 and 2019. The grasslands ranged from unfertilized and grazed grasslands to intensively fertilized and frequently harvested ones. To assess effects of grazing and fertilization intensities and plant diversity on soil CO2 efflux, we used Structural Equation Modeling to account for direct effects and indirect effects through soil and plant organic matter quantity and quality. Soil CO2 efflux was suppressed by limited water availability caused by naturally occurring droughts in both study years. Under the prevailing environmental conditions, grazing intensity, plant biomass and plant C:N ratio were not related to soil CO2 efflux. In contrast, fertilization intensity was positively associated with soil CO2 efflux (standardized coefficient of net effect: + 0.04 in 2018 and + 0.03 in 2019). This was because fertilization led to lower plant species richness and, thus, to lower C:N ratios in soils, which were associated with higher soil CO2 efflux (plant species richness net effect: −0.09 in 2018 and −0.18 in 2019; soil C:N ratio direct effect: −0.23 in 2018 and −0.33 in 2019). Intensively managed grasslands have higher soil respiration than extensively managed, plant species-rich grasslands even under the extreme conditions of natural droughts., Nutrient Cycling in Agroecosystems, 124, ISSN:1385-1314, ISSN:1573-0867

To improve our understanding of early microbial colonization of pristine minerals and their group-specific C utilization, we exposed minerals (illite/goethite/quartz) amended with artificial root exudates (ARE, glucose, and citric acid) in grassland soils for a period of 24 weeks. FTIR spectra indicated that mineral-associated ARE were used within the first 2 weeks of exposure and were replaced by other carbohydrates derived from living or dead cells as well as soil-borne C sources transported into the mineralosphere after heavy rain events. Fungi and Gram-positive bacteria incorporated ARE-derived C more rapidly than Gram-negative bacteria. Gram-negative bacteria presumably profited indirectly from the ARE by cross-feeding on mineral-associated necromass of fungi and Gram-positive bacteria. The Gram-negative bacterial phyla Verrucomicrobia, Planctomycetes, Gemmatimonadetes, Armatimonadetes, and Chloroflexi showed a positive correlation with Gram-negative PLFA abundances. After 24 weeks of exposure in the grassland soils, abundances of soil microorganisms in the mineralosphere reached only 3.1% of the population density in soil. In conclusion, both bacteria and fungi slowly colonize new surfaces such as pristine minerals, but quickly assimilate artificial root exudates, creating an active microbial community in the mineralosphere.

Per- and polyfluoroalkyl substances (PFAS) are environmentally ubiquitous, anthropogenic substances with adverse effects on organisms, which shows the need to study their environmental fate and leaching behavior. In the present soil columns study, the leaching behavior and fate of nontransformable and transformable (precursors) were investigated. Ten nontransformable PFAS in two different soils, two precursors and two field soils, which were already contaminated with a mixture of PFAS, and two uncontaminated controls, were set up for a time span of 2 years. At the end of the study, the molecular balance could not be closed for nontransformable PFAS. This effect was positively correlated to the fluorinated carbon chain length. The precursors, which were both polyfluoroalkyl phosphate diesters (diPAP), had different transformation products and transformation rates, with a higher rate for 6:2 diPAP than 8:2 diPAP. After 2 years, amounts of diPAP were still found in the soil with no significant vertical movement, showing high adsorption to soils. Transformation products were estimated to be simultaneously formed. They were predominantly found in the percolation water; the amounts left in soil were negligible. Up to half of the initial precursor amounts could not be balanced and were considered missing amounts. The results of contaminated field soil experiments showed the challenge to estimate PFAS leaching without knowing all occurring precursors and complex transformation dynamics. For this purpose, it was shown that a broad examination of contaminated soil with different analytical methods can help with qualitative estimations of leaching risks. For a better quantitative estimation, analytical determination of more PFAS and a quantification of the missing amounts are needed. Environ Toxicol Chem 2022;41:2065-2077. © 2022 The Authors. Environmental Toxicology and Chemistry published by Wiley Periodicals LLC on behalf of SETAC.

Soil scientists are accustomed to geostatistical methods and tools such as semivariograms and kriging for analysis of observational data. Such methods assume and exploit that observations are spatially correlated. Conversely, analysis of variance (ANOVA) of designed experiments assumes that observations from different experimental units are independent, an assumption that is justified based on randomization. It may be beneficial, however, to perform an ANOVA assuming a geostatistical covariance model. Also, it is increasingly common to have multiple observations per experimental unit. Simple ANOVA assuming independence of observations is not appropriate for such data. Instead, a linear mixed model accounting for correlation among observations made on the same plot is required for proper analysis. The purpose of this paper is to demonstrate the benefits of integrating geostatistical covariance structures and ANOVA procedures into a linear mixed modelling framework. Two examples from designed experiments are considered in detail, making a link between terminologies and jargon used in geostatistical analysis on the one hand and linear mixed modelling on the other hand. We provide code in R and SAS for both examples in two supporting companion documents. HIGHLIGHTS: Analysis of variance and geostatistical analysis can be joined in a mixed model. Randomization justifies the independence assumption in analysis of variance. Geostatistical models imply a correlation of errors and can improve efficiency. Lacking randomization, spatial correlation can be accounted for in a mixed model.

Grassland management intensity influences nutrient cycling both directly, by changing nutrient inputs and outputs from the ecosystem, and indirectly, by altering the nutrient content, and the diversity and functional composition of plant and microbial communities. However, the relative importance of these direct and indirect processes for the leaching of multiple nutrients is poorly studied. We measured the annual leaching of nitrate, ammonium, phosphate and sulphate at a depth of 10 cm in 150 temperate managed grasslands using a resin method. Using Structural Equation Modeling, we distinguished between various direct and indirect effects of management intensity (i.e. grazing and fertilization) on nutrient leaching. We found that management intensity was positively associated with nitrate, ammonium and phosphate leaching risk both directly (i.e. via increased nutrient inputs) and indirectly, by changing the stoichiometry of soils, plants and microbes. In contrast, sulphate leaching risk was negatively associated with management intensity, presumably due to increased outputs with mowing and grazing. In addition, management intensification shifted plant communities towards an exploitative functional composition (characterized by high tissue turnover rates) and, thus, further promoted the leaching risk of inorganic nitrogen. Plant species richness was associated with lower inorganic nitrogen leaching risk, but most of its effects were mediated by stoichiometry and plant community functional traits. Maintaining and restoring diverse plant communities may therefore mitigate the increased leaching risk that management intensity imposes upon grasslands., Nutrient Cycling in Agroecosystems, 123 (3), ISSN:1385-1314, ISSN:1573-0867

The impact of local biodiversity loss on ecosystem functioning is well established, but the role of larger-scale biodiversity dynamics in the delivery of ecosystem services remains poorly understood. Here we address this gap using a comprehensive dataset describing the supply of 16 cultural, regulating and provisioning ecosystem services in 150 European agricultural grassland plots, and detailed multi-scale data on land use and plant diversity. After controlling for land-use and abiotic factors, we show that both plot-level and surrounding plant diversity play an important role in the supply of cultural and aboveground regulating ecosystem services. In contrast, provisioning and belowground regulating ecosystem services are more strongly driven by field-level management and abiotic factors. Structural equation models revealed that surrounding plant diversity promotes ecosystem services both directly, probably by fostering the spill-over of ecosystem service providers from surrounding areas, and indirectly, by maintaining plot-level diversity. By influencing the ecosystem services that local stakeholders prioritized, biodiversity at different scales was also shown to positively influence a wide range of stakeholder groups. These results provide a comprehensive picture of which ecosystem services rely most strongly on biodiversity, and the respective scales of biodiversity that drive these services. This key information is required for the upscaling of biodiversity–ecosystem service relationships, and the informed management of biodiversity within agricultural landscapes.

Plant–soil feedbacks (PSFs) underlying grassland plant richness and productivity are typically coupled with nutrient availability; however, we lack understanding of how restoration measures to increase plant diversity might affect PSFs. We examined the roles of sward disturbance, seed addition and land-use intensity (LUI) on PSFs. We conducted a disturbance and seed addition experiment in 10 grasslands along a LUI gradient and characterized plant biomass and richness, soil microbial biomass, community composition and enzyme activities. Greater plant biomass at high LUI was related to a decrease in the fungal to bacterial ratios, indicating highly productive grasslands to be dominated by bacteria. Lower enzyme activity per microbial biomass at high plant species richness indicated a slower carbon (C) cycling. The relative abundance of fungal saprotrophs decreased, while pathogens increased with LUI and disturbance. Both fungal guilds were negatively associated with plant richness, indicating the mechanisms underlying PSFs depended on LUI. We show that LUI and disturbance affect fungal functional composition, which may feedback on plant species richness by impeding the establishment of pathogen-sensitive species. Therefore, we highlight the need to integrate LUI including its effects on PSFs when planning for practices that aim to optimize plant diversity and productivity., New Phytologist, 236 (5), ISSN:0028-646X, ISSN:1469-8137

Bacteria colonize reactive minerals in soils where they contribute to mineral weathering and transformation. So far, the specificity, patterns and dynamics of mineral colonization have rarely been assessed under natural conditions. High throughput Illumina sequencing was employed to investigate the bacterial communities assembling on illite and goethite during exposure to natural grassland soils. Two different types of organic carbon sources, simple carbon compounds representing root exudates and detritus of two dominant grassland plant species were applied, and their effects on the temporal dynamics of bacterial communities were investigated. The observed temporal patterns suggest that the surfaces of de novo exposed minerals in soils drive the establishment of bacterial communities and override the effect of the type of carbon sources and of other environmental properties. Mineral colonization was selective and specific bacterial sequence variants exhibited distinct colonization patterns, among which early, intermittent, and late colonizers could be distinguished. Based on our results, soil minerals are not only colonized by specific bacterial communities but enable a succession of different bacterial communities. Our results thereby expand the concept of the mineralosphere and provide novel insights into mechanisms of community assembly in the soil ecosystem.

Summary Soils provide a heterogeneous environment varying in space and time; consequently, the biodiversity of soil microorganisms also differs spatially and temporally. For soil microbes tightly associated with plant roots, such as arbuscular mycorrhizal fungi (AMF), the diversity of plant partners and seasonal variability in trophic exchanges between the symbionts introduce additional heterogeneity. To clarify the impact of such heterogeneity, we investigated spatiotemporal variation in AMF diversity on a plot scale (10 × 10 m) in a grassland managed at low intensity in southwest Germany. AMF diversity was determined using 18S rDNA pyrosequencing analysis of 360 soil samples taken at six time points within a year. We observed high AMF alpha‐ and beta‐diversity across the plot and at all investigated time points. Relationships were detected between spatiotemporal variation in AMF OTU richness and plant species richness, root biomass, minimal changes in soil texture and pH. The plot was characterized by high AMF turnover rates with a positive spatiotemporal relationship for AMF beta‐diversity. However, environmental variables explained only ≈20% of the variation in AMF communities. This indicates that the observed spatiotemporal richness and community variability of AMF was largely independent of the abiotic environment, but related to plant properties and the cooccurring microbiome.

Land‐use intensification drives changes in microbial communities and the soil functions they regulate, but the mechanisms underlying these changes are poorly understood as land use can affect soil communities both directly (e.g. via changes in soil fertility) and indirectly (e.g. via changes in plant inputs). The speed of microbial responses is also poorly understood. For instance, whether it is long‐term legacies or short‐term changes in land‐use intensity that drive changes in microbial communities. To address these topics, we measured multiple microbial functions, bacterial and fungal biomass and abiotic soil properties at two time intervals 3 years apart. This was performed in 150 grassland sites differing greatly in management intensity across three German regions. Observed changes in microbial soil properties were related to both long‐term means and short‐term changes in: abiotic soil properties, land‐use intensity, community abundance‐weighted means of plant functional traits and plant biomass properties in regression and structural equation models. Plant traits, particularly leaf phosphorus, and soil pH were the best predictors of change in soil microbial function, as well as fungal and bacterial biomass, while land‐use intensity showed weaker effects. Indirect legacy effects, in which microbial change was explained by the effects of long‐term land‐use intensity on plant traits, were important, thus indicating a time lag between plant community and microbial change. Whenever the effects of short‐term changes in land‐use intensity were present, they acted directly on soil microorganisms. Synthesis. The results provide new evidence that soil communities and their functioning respond to short‐term changes in land‐use intensity, but that both rapid and longer time‐scale responses to changes in plant functional traits are at least of equal importance. This suggests that management which shapes plant communities may be an effective means of managing soil communities and the functions and services they provide.

Evidence from experimental and established grasslands indicates that plant biodiversity can modify the water cycle. One suspected mechanism behind this is a higher infiltration capacity (νB) and hydraulic conductivity (K) of the soil on species‐rich grasslands. However, in established and agriculturally managed grasslands, biodiversity effects cannot be studied independent of land‐use effects. Therefore, we investigated in established grassland systems how land‐use intensity and associated biodiversity of plants and soil animals affect νB and K at and close to saturation. On 50 grassland plots along a land‐use intensity gradient in the Biodiversity Exploratory Schwäbische Alb, Germany, we measured νB with a hood infiltrometer at several matrix potentials and calculated the saturated and unsaturated K. We statistically analysed the relationship between νB or K and land‐use information (e.g., fertilising intensity), abiotic (e.g., soil texture) and biotic data (e.g., plant species richness, earthworm abundance). Land‐use intensity decreased and plant species richness increased νB and K, while the direction of the effects of soil animals was inconsistent. The effect of land‐use intensity on νB and K was mainly attributable to its negative effect on plant species richness. Our results demonstrate that plant species richness was a better predictor of νB and K at and close to saturation than land‐use intensity or soil physical properties in the established grassland systems of the Schwäbische Alb., Deutsche Forschungsgemeinschaft http://dx.doi.org/10.13039/501100001659, Stiftung Landesbank Baden‐Württemberg, Swiss National Science Foundation http://dx.doi.org/10.13039/501100001711, http://doi.org/10.17616/R32P9Q

Acidobacteria occur in a large variety of ecosystems worldwide and are particularly abundant and highly diverse in soils. In spite of their diversity, only few species have been characterized to date which makes

Land-use intensification can increase provisioning ecosystem services, such as food and timber production, but it also drives changes in ecosystem functioning and biodiversity loss, which may ultimately compromise human wellbeing. To understand how changes in land-use intensity affect the relationships between biodiversity, ecosystem functions, and services, we built networks from correlations between the species richness of 16 trophic groups, 10 ecosystem functions, and 15 ecosystem services. We evaluated how the properties of these networks varied across land-use intensity gradients for 150 forests and 150 grasslands. Land-use intensity significantly affected network structure in both habitats. Changes in connectance were larger in forests, while changes in modularity and evenness were more evident in grasslands. Our results show that increasing land-use intensity leads to more homogeneous networks with less integration within modules in both habitats, driven by the belowground compartment in grasslands, while forest responses to land management were more complex. Land-use intensity strongly altered hub identity and module composition in both habitats, showing that the positive correlations of provisioning services with biodiversity and ecosystem functions found at low land-use intensity levels, decline at higher intensity levels. Our approach provides a comprehensive view of the relationships between multiple components of biodiversity, ecosystem functions, and ecosystem services and how they respond to land use. This can be used to identify overall changes in the ecosystem, to derive mechanistic hypotheses, and it can be readily applied to further global change drivers. The work has been supported by the DFG Priority Program 1374 “Infrastructure-Biodiversity-Exploratories”. S.S. was supported by the Spanish Government under Ramón y Cajal Contract RYC-2016-20604.

Aerated topsoils are important sinks for atmospheric methane (CH4) via oxidation by CH4‐oxidizing bacteria (MOB). However, intensified management of grasslands and forests may reduce the CH4 sink capacity of soils. We investigated the influence of grassland land‐use intensity (150 sites) and forest management type (149 sites) on potential atmospheric CH4 oxidation rates (PMORs) and the abundance and diversity of MOB (with qPCR) in topsoils of three temperate regions in Germany. PMORs measurements in microcosms under defined conditions yielded approximately twice as much CH4 oxidation in forest than in grassland soils. High land‐use intensity of grasslands had a negative effect on PMORs (−40%) in almost all regions and fertilization was the predominant factor of grassland land‐use intensity leading to PMOR reduction by 20%. In contrast, forest management did not affect PMORs in forest soils. Upland soil cluster (USC)‐α was the dominant group of MOBs in the forests. In contrast, USC‐γ was absent in more than half of the forest soils but present in almost all grassland soils. USC‐α abundance had a direct positive effect on PMOR in forest, while in grasslands USC‐α and USC‐γ abundance affected PMOR positively with a more pronounced contribution of USC‐γ than USC‐α. Soil bulk density negatively influenced PMOR in both forests and grasslands. We further found that the response of the PMORs to pH, soil texture, soil water holding capacity and organic carbon and nitrogen content differ between temperate forest and grassland soils. pH had no direct effects on PMOR, but indirect ones via the MOB abundances, showing a negative effect on USC‐α, and a positive on USC‐γ abundance. We conclude that reduction in grassland land‐use intensity and afforestation has the potential to increase the CH4 sink function of soils and that different parameters determine the microbial methane sink in forest and grassland soils. Deutsche Forschungsgemeinschaft http://dx.doi.org/10.13039/501100001659 ESF Ministry of Education, Science and Culture of Mecklenburg‐Western Pomerania

Soil protists are increasingly appreciated as essential components of soil foodwebs; however, there is a dearth of information on the factors structuring their communities. Here we investigate the importance of different biotic and abiotic factors as key drivers of spatial and seasonal distribution of protistan communities. We conducted an intensive survey of a 10m2 grassland plot in Germany, focusing on a major group of protists, the Cercozoa. From 177 soil samples, collected from April to November, we obtained 694 Operational Taxonomy Units representing >6 million Illumina reads. All major cercozoan taxonomic and functional groups were present, dominated by the small flagellates of the Glissomonadida. We found evidence of environmental selection structuring the cercozoan communities both spatially and seasonally. Spatial analyses indicated that communities were correlated within a range of 3.5 meters. Seasonal variations in the abundance of bacterivores and bacteria, followed by that of omnivores suggested a dynamic prey-predator succession. The most influential edaphic properties were moisture and clay content, which differentially affected each functional group. Our study is based on an intense sampling of protists at a small scale, thus providing a detailed description of the biodiversity of different taxa/functional groups and the ecological processes involved in shaping their distribution.

Soil protists are increasingly appreciated as essential components of soil foodwebs; however, there is a dearth of information on the factors structuring their communities. Here we investigate the importance of different biotic and abiotic factors as key drivers of spatial and seasonal distribution of protistan communities. We conducted an intensive survey of a 10m2 grassland plot in Germany, focusing on a major group of protists, the Cercozoa. From 177 soil samples, collected from April to November, we obtained 694 Operational Taxonomy Units representing >6 million Illumina reads. All major cercozoan groups were present, dominated by the small flagellates of the Glissomonadida. We found evidence of environmental filtering structuring the cercozoan communities both spatially and seasonally. Spatial analyses indicated that communities were correlated within a range of four meters. Seasonal variations of bactevirores and bacteria, and that of omnivores after a time-lapse, suggested a dynamic prey-predator succession. The most influential edaphic properties were moisture and clay content, which differentially affected each functional group. Our study is based on an intense sampling of protists at a small scale, thus providing a detailed description of the niches occupied by different taxa/functional groups and the ecological processes involved.

In heterogeneous environments such as soil it is imperative to understand the spatial relationships between microbial communities, microbial functioning and microbial habitats in order to predict microbial services in managed grasslands. Grassland land-use intensity has been shown to affect the spatial distribution of soil microorganisms, but so far it is unknown whether this is transferable from one geographic region to another. This study evaluated the spatial distribution of soil microbial biomass and enzyme activities involved in C-, N- and P-cycling, together with physico-chemical soil properties in 18 grassland sites differing in their land-use intensity in two geographic regions: the Hainich National Park in the middle of Germany and the Swabian Alb in south-west Germany. Enzyme activities associated with the C- and N-cycles, namely β-glucosidase, xylosidase and chitinase, organic carbon (Corg), total nitrogen (Nt), extractable organic carbon, and mineral nitrogen (Nmin) were higher in the Swabian Alb (Leptosols) than in the Hainich National Park (primarily Stagnosols). There was a negative relationship between bulk density and soil properties such as microbial biomass (Cmic, Nmic), urease, Corg, and Nt. The drivers (local abiotic soil properties, spatial separation) of the enzyme profiles (β-glucosidase, chitinase, xylosidase, phosphatase, and urease) were determined through a spatial analysis of the within site variation of enzyme profiles and abiotic properties, using the Procrustes rotation test. The test revealed that physical and chemical properties showed more spatial pattern than the enzyme profiles. β-glucosidase, chitinase, xylosidase, phosphatase, and urease activities were related to local abiotic soil properties, but showed little spatial correlation. Semivariogram modeling revealed that the ranges of spatial autocorrelation of all measured variables were site specific and not related to region or to land-use intensity. Nevertheless, land-use intensity changed the occurrence of spatial patterns measurable at the plot scale: increasing land-use intensity led to an increase in detectable spatial patterns for abiotic soil properties on Leptosols. The conclusion of this study is that microbial biomass and functions in grassland soils do not follow general spatial distribution patterns, but that the spatial distribution is site-specific and mainly related to the abiotic properties of the soils.

Management intensity modifies soil properties, e.g., organic carbon (Corg) concentrations and soil pH with potential feedbacks on plant diversity. These changes might influence microbial P concentrations (Pmic) in soil representing an important component of the P cycle. Our objectives were to elucidate whether abiotic and biotic variables controlling Pmic concentrations in soil are the same for forests and grasslands, and to assess the effect of region and management on Pmic concentrations in forest and grassland soils as mediated by the controlling variables. In three regions of Germany, Schwäbische Alb, Hanich-Dün, and Schorfheide-Chorin, we studied forest and grassland plots (each n = 150) differing in plant diversity and land-use intensity. In contrast to controls of microbial biomass carbon (Cmic), Pmic was strongly influenced by soil pH, which in turn affected phosphorus (P) availability and thus microbial P uptake in forest and grassland soils. Furthermore, Pmic concentrations in forest and grassland soils increased with increasing plant diversity. Using structural equation models, we could show that soil Corg is the profound driver of plant diversity effects on Pmic in grasslands. For both forest and grassland, we found regional differences in Pmic attributable to differing environmental conditions (pH, soil moisture). Forest management and tree species showed no effect on Pmic due to a lack of effects on controlling variables (e.g., Corg). We also did not find management effects in grassland soils which might be caused by either compensation of differently directed effects across sites or by legacy effects of former fertilization constraining the relevance of actual practices. We conclude that variables controlling Pmic or Cmic in soil differ in part and that regional differences in controlling variables are more important for Pmic in soil than those induced by management., Journal of Plant Nutrition and Soil Science, 181 (2), ISSN:1436-8730, ISSN:0044-3263, ISSN:1522-2624

Temporal dynamics create unique and often ephemeral conditions that can influence soil microbial biogeography at different spatial scales. This study investigated the relation between decimeter to meter spatial variability of soil microbial community structure, plant diversity, and soil properties at six dates from April through November. We also explored the robustness of these interactions over time. An historically unfertilized, unplowed grassland in southwest Germany was selected to characterize how seasonal variability in the composition of plant communities and substrate quality changed the biogeography of soil microorganisms at the plot scale (10 m × 10 m). Microbial community spatial structure was positively correlated with the local environment, i.e. physical and chemical soil properties, in spring and autumn, while the density and diversity of plants had an additional effect in the summer period. Spatial relationships among plant and microbial communities were detected only in the early summer and autumn periods when aboveground biomass increase was most rapid and its influence on soil microbial communities was greatest due to increased demand by plants for nutrients. Individual properties exhibited varying degrees of spatial structure over the season. Differential responses of Gram positive and Gram negative bacterial communities to seasonal shifts in soil nutrients were detected. We concluded that spatial distribution patterns of soil microorganisms change over a season and that chemical soil properties are more important controlling factors than plant density and diversity. Finer spatial resolution, such as the mm to cm scale, as well as taxonomic resolution of microbial groups, could help determine the importance of plant species density, composition, and growth stage in shaping microbial community composition and spatial patterns.

The mineralo-sphere is an important micro habitat in terrestrial ecosystems. How different groups of microorganisms colonise mineral surfaces and whether the level of grassland land-use intensity (LUI) modifies this micro-habitat is not well known, however. We exposed mesh containers filled with pristine soil minerals (illite/goethite) mixed with 13 C labelled root litter of Dactylis glomerata and Lolium perenne in grassland soils of the “Schwabische Alb” (Germany) to characterise the succession of different microbial properties in the mineralo-sphere. The use of sites within the Biodiversity Exploratories made it possible to select five sites of low LUI and five sites of high LUI. After 1, 2, 7, 12 and 31 months of exposure in the grassland soils, we used physiological, microbiological and isotopic methods to elucidate in situ colonisation patterns, carbon use and levels of extracellular enzyme expression by soil microorganisms associated with mineral surfaces. Microorganisms slowly colonised pristine mineral surfaces and established functionally distinct communities over time. Fungi colonised mineral surfaces to a greater extent than bacteria, reaching 13.2% of control soils compared to 3.2% by bacteria after 31 months. Fungi also reached pristine mineral surfaces earlier than bacteria, probably due to their hyphal growth strategies, and made immediate use of the added complex root litter substrate. This result is evident by the incorporation of up to 74% root litter-derived C into the fungus-specific PLFA (18:2ω6,9) compared to 51% root litter-derived C in the bacteria-specific PLFAs. Both bacteria and fungi associated with minerals remained in an active state (high biomass-specific respiration, high bacterial and fungal growth rates) throughout the experimental period. Grassland LUI and physico-chemical properties of the adjacent soil modified both quantity and quality of substrates available to soil microorganisms in the mineralo-sphere. Since 13 C incorporation into microbial biomass was greater under low LUI than under high LUI, we conclude that microorganisms in low LUI sites had to rely on the added root material, while the carbon signal in microorganisms in the high LUI sites was diluted by alternative sources resulting from transport of dissolved organic carbon into the mineralo-sphere.

EABIOmESUPDATCT3 INRA; The microbial groups of nitrogen fixers, ammonia oxidizers, and denitrifiers largely drive the inorganic nitrogen cycle in temperate terrestrial ecosystems. Their spatial and temporal dynamics, however, vary depending on the studied scale. The present study aimed to fill a knowledge gap by providing an explicit picture of spatial and temporal dynamics of a subset of these soil microorganisms at the plot scale. We selected an unfertilized perennial grassland, where nitrogen cycling is considered to be efficient and tightly coupled to plant growth. At six times over one growing season 60 soil samples were taken from a 10 m x 10 m area and abundances of marker genes for total archaea and bacteria (16S rRNA), nitrogen fixing bacteria (nifH), ammonia oxidizing archaea (amoA AOA) and bacteria (amoA AOB), and denitrifying bacteria (nirS, nirK and nosZ) were determined by qPCR. Potential nitrification activity (PNA) and denitrifying enzyme activity (DEA) were determined. Seasonal changes in abundance patterns of marker genes were detected, and were associated with changes in substrate availability associated with plant growth stages. Potential nitrification and denitrification enzyme activities were strongly spatially structured at the studied scale, corresponding to periods of rapid plant growth, June and October, and their spatial distributions were similar, providing visual evidence of highly localized spatial and temporal conditions at this scale. Temporal variability in the N -cycling communities versus the stability of their respective potential activities provided evidence of both short-lived temporal niche partitioning and a degree of microbial functional redundancy. Our results indicate that in an unfertilized grassland, at the meter scale, abundances of microbial N-cycling organisms can exhibit transient changes, while nitrogen cycling processes remain stable.

Characterization of spatial and temporal variation in grassland productivity and nutrition is crucial for a comprehensive understanding of ecosystem function. Although within-site heterogeneity in soil and plant properties has been shown to be relevant for plant community stability, spatiotemporal variability in these factors is still understudied in temperate grasslands. Our study aimed to detect if soil characteristics and plant diversity could explain observed small-scale spatial and temporal variability in grassland productivity, biomass nutrient concentrations, and nutrient limitation. Therefore, we sampled 360 plots of 20 cm × 20 cm each at six consecutive dates in an unfertilized grassland in Southern Germany. Nutrient limitation was estimated using nutrient ratios in plant biomass. Absolute values of, and spatial variability in, productivity, biomass nutrient concentrations, and nutrient limitation were strongly associated with sampling date. In April, spatial heterogeneity was high and most plots showed phosphorous deficiency, while later in the season nitrogen was the major limiting nutrient. Additionally, a small significant positive association between plant diversity and biomass phosphorus concentrations was observed, but should be tested in more detail. We discuss how low biological activity e.g., of soil microbial organisms might have influenced observed heterogeneity of plant nutrition in early spring in combination with reduced active acquisition of soil resources by plants. These early-season conditions are particularly relevant for future studies as they differ substantially from more thoroughly studied later season conditions. Our study underlines the importance of considering small spatial scales and temporal variability to better elucidate mechanisms of ecosystem functioning and plant community assembly., Plant Ecology, 217 (7), ISSN:1385-0237, ISSN:1573-5052

Increased warming in spring and prolonged summer drought may alter soil microbial denitrification. We measured potential denitrification activity and denitrifier marker gene abundances ( nirK , nirS , nosZ ) in grasslands soils in three geographic regions characterized by site-specific land-use indices (LUI) after warming in spring, at an intermediate sampling and after summer drought. Potential denitrification was significantly increased by warming, but did not persist over the intermediate sampling. At the intermediate sampling, the relevance of grassland land-use intensity was reflected by increased potential N2O production at sites with higher LUI. Abundances of total bacteria did not respond to experimental warming or drought treatments, displaying resilience to minor and short-term effects of climate change. In contrast, nirS- and nirK- type denitrifiers were more influenced by drought in combination with LUI and pH, while the nosZ abundance responded to the summer drought manipulation. Land-use was a strong driver for potential denitrification as grasslands with higher LUI also had greater potentials for N2O emissions. We conclude that both warming and drought affected the denitrifying communities and the potential denitrification in grassland soils. However, these effects are overruled by regional and site-specific differences in soil chemical and physical properties which are also related to grassland land-use intensity.

Interactions occur between two or more organisms affecting each other. Interactions are decisive for the ecology of the organisms. Without direct experimental evidence the analysis of interactions is difficult. Correlation analyses that are based on co-occurrences are often used to approximate interaction. Here, we present a new mathematical model to estimate the interaction strengths between taxa, based on changes in their relative abundances across environmental gradients.