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Abstract Organismal functional strategies form a continuum from slow- to fast-growing organisms, in response to common drivers such as resource availability and disturbance. However, whether there is synchronisation of these strategies at the entire community level is unclear. Here, we combine trait data for >2800 above- and belowground taxa from 14 trophic guilds spanning a disturbance and resource availability gradient in German grasslands. The results indicate that most guilds consistently respond to these drivers through both direct and trophically mediated effects, resulting in a ‘slow-fast’ axis at the level of the entire community. Using 15 indicators of carbon and nutrient fluxes, biomass production and decomposition, we also show that fast trait communities are associated with faster rates of ecosystem functioning. These findings demonstrate that ‘slow’ and ‘fast’ strategies can be manifested at the level of whole communities, opening new avenues of ecosystem-level functional classification.

Biostimulants (Bio-effectors, BEs) comprise plant growth-promoting microorganisms and active natural substances that promote plant nutrient-acquisition, stress resilience, growth, crop quality and yield. Unfortunately, the effectiveness of BEs, particularly under field conditions, appears highly variable and poorly quantified. Using random model meta-analyses tools, we summarize the effects of 107 BE treatments on the performance of major crops, mainly conducted within the EU-funded project BIOFECTOR with a focus on phosphorus (P) nutrition, over five years. Our analyses comprised 94 controlled pot and 47 field experiments under different geoclimatic conditions, with variable stress levels across European countries and Israel. The results show an average growth/yield increase by 9.3% (n=945), with substantial differences between crops (tomato > maize > wheat) and growth conditions (controlled nursery + field (Seed germination and nursery under controlled conditions and young plants transplanted to the field) > controlled > field). Average crop growth responses were independent of BE type, P fertilizer type, soil pH and plant-available soil P (water-P, Olsen-P or Calcium acetate lactate-P). BE effectiveness profited from manure and other organic fertilizers, increasing soil pH and presence of abiotic stresses (cold, drought/heat or salinity). Systematic meta-studies based on published literature commonly face the inherent problem of publication bias where the most suspected form is the selective publication of statistically significant results. In this meta-analysis, however, the results obtained from all experiments within the project are included. Therefore, it is free of publication bias. In contrast to reviews of published literature, our unique study design is based on a common standardized protocol which applies to all experiments conducted within the project to reduce sources of variability. Based on data of crop growth, yield and P acquisition, we conclude that application of BEs can save fertilizer resources in the future, but the efficiency of BE application depends on cropping systems and environments.

Relationships between biodiversity and phosphorus cycling and the underlying processes are complex. Here the authors analyse a biodiversity manipulation experiment and an agricultural management gradient to show how plant and mycorrhizal fungal diversity promote phosphorus exploitation.

Adoption of microbial consortia as plant growth-promoting microorganisms (PGPMs) instead of single-strain inoculants is discussed as an approach to increase the efficiency and flexibility of PGPM-assisted production strategies. This study provides the functional characterisation of a commercial microbial consortia product (MCP) in a series of greenhouse experiments with maize on a silty-loam field soil (pH 5.9). A 60%-increased abundance of bacteria that could be cultivated after rhizosphere extraction was measured after MCP inoculation at the end of the 42-days culture period. MCP inoculation did not stimulate shoot biomass production of maize fertilised with nitrate, but growth improvement was recorded in combination with stabilised ammonium, especially with reduced phosphorus (P) supply. The MCP inoculant improved the acquisition of ammonium-N but also increased shoot-P. MCP inoculation stimulated root length development under reduced P supply with stabilised ammonium by 52%. This was accompanied by the increased auxin production capacity of rhizosphere bacteria. C-, N-, and P-turnover in the rhizosphere were little affected by the MCP inoculation, as deduced from the analysis of activities of extracellular soil enzymes. The findings suggest that the form of N supply is crucial for the efficiency of plant-MCP interactions.

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 Acidobacteria one of the most poorly understood phyla among the domain Bacteria. We used a culture-independent niche modeling approach to elucidate ecological adaptations and their evolution for 4,154 operational taxonomic units (OTUs) of Acidobacteria across 150 different, comprehensively characterized grassland soils in Germany. Using the relative abundances of their 16S rRNA gene transcripts, the responses of active OTUs along gradients of 41 environmental variables were modeled using hierarchical logistic regression (HOF), which allowed to determine values for optimum activity for each variable (niche optima). By linking 16S rRNA transcripts to the phylogeny of full 16S rRNA gene sequences, we could trace the evolution of the different ecological adaptations during the diversification of Acidobacteria. This approach revealed a pronounced ecological diversification even among acidobacterial sister clades. Although the evolution of habitat adaptation was mainly cladogenic, it was disrupted by recurrent events of convergent evolution that resulted in frequent habitat switching within individual clades. Our findings indicate that the high diversity of soil acidobacterial communities is largely sustained by differential habitat adaptation even at the level of closely related species. A comparison of niche optima of individual OTUs with the phenotypic properties of their cultivated representatives showed that our niche modeling approach (1) correctly predicts those physiological properties that have been determined for cultivated species of Acidobacteria but (2) also provides ample information on ecological adaptations that cannot be inferred from standard taxonomic descriptions of bacterial isolates. These novel information on specific adaptations of not-yet-cultivated Acidobacteria can therefore guide future cultivation trials and likely will increase their cultivation success.

Current farm systems rely on the use of Plant Protection Products (PPP) to secure high productivity and control threats to the quality of the crops. However, PPP use may have considerable impacts on human health and the environment. A study protocol is presented aiming to determine the occurrence and levels of PPP residues in plants (crops), animals (livestock), humans and other non-target species (ecosystem representatives) for exposure modelling and impact assessment. To achieve this, we designed a cross-sectional study to compare conventional and organic farm systems across Europe. Environmental and biological samples were/are being/will be collected during the 2021 growing season, at 10 case study sites in Europe covering a range of climate zones and crops. An additional study site in Argentina will inform the impact of PPP use on growing soybean which is an important European protein-source in animal feed. We will study the impact of PPP mixtures using an integrated risk assessment methodology. The fate of PPP in environmental media (soil, water and air) and in the homes of farmers will be monitored. This will be complemented by biomonitoring to estimate PPP uptake by humans and farm animals (cow, goat, sheep and chicken), and by collection of samples from non-target species (earthworms, fish, aquatic and terrestrial macroinvertebrates, bats, and farm cats). We will use data on PPP residues in environmental and biological matrices to estimate exposures by modelling. These exposure estimates together with health and toxicity data will be used to predict the impact of PPP use on environment, plant, animal and human health. The outcome of this study will then be integrated with socio-economic information leading to an overall assessment used to identify transition pathways towards more sustainable plant protection and inform decision makers, practitioners and other stakeholders regarding farming practices and land use policy.

Assessing agricultural sustainability is one of the most challenging tasks related to expertise and support methodologies because it entails multidisciplinary aspects and builds on cultural and value-based elements. Thus, agricultural sustainability should be considered a social concept, reliable enough to support decision makers and policy development in a broad context. The aim of this manuscript was to develop a methodology for the assessment of the sustainability of soil improving cropping systems (SICS) in Europe. For this purpose, a decision tree based on weights (%) was chosen because it allows more flexibility. The methodology was tested with data from the SoilCare Horizon 2020 study site in Germany for the assessment of the impact of the integration of cover crops into the crop rotation. The effect on the environmental indicators was slightly positive, but most assessed properties did not change over the short course of the experiment. Farmers reported that the increase in workload was outweighed by a reputation gain for using cover crops. The incorporation of cover crops reduced slightly the profitability, due to the costs for seeds and establishment of cover crops. The proposed assessment methodology provides a comprehensive summary to assess the agricultural sustainability of SICS.

Soils form the basis for agricultural production and other ecosystem services, and soil management should aim at improving their quality and resilience. Within the SoilCare project, the concept of soil-improving cropping systems (SICS) was developed as a holistic approach to facilitate the adoption of soil management that is sustainable and profitable. SICS selected with stakeholders were monitored and evaluated for environmental, sociocultural, and economic effects to determine profitability and sustainability. Monitoring results were upscaled to European level using modelling and Europe-wide data, and a mapping tool was developed to assist in selection of appropriate SICS across Europe. Furthermore, biophysical, sociocultural, economic, and policy reasons for (non)adoption were studied. Results at the plot/farm scale showed a small positive impact of SICS on environment and soil, no effect on sustainability, and small negative impacts on economic and sociocultural dimensions. Modelling showed that different SICS had different impacts across Europe—indicating the importance of understanding local dynamics in Europe-wide assessments. Work on adoption of SICS confirmed the role economic considerations play in the uptake of SICS, but also highlighted social factors such as trust. The project’s results underlined the need for policies that support and enable a transition to more sustainable agricultural practices in a coherent way.

Amendment of soils with plant residues is common practice for improving soil quality. In addition to stimulated microbial activity, the supply of fresh soluble organic (C) from litter may accelerate the microbial degradation of chemicals in soils. Therefore, the aim of this study was to test whether the maize litter enhances degradation of 4-chloro-2-methylphenoxyacetic acid (MCPA) and increases formation of non-toxic biogenic non-extractable residues (bioNERs). Soil was amended with 13C6-MCPA and incubated with or without litter addition on the top. Three soil layers were sampled with increasing distance from the top: 0–2 mm, 2–5 mm and 5–20 mm; and the mass balance of 13C6-MCPA transformation determined.Maize litter promoted microbial activity, mineralization of 13C6-MCPA and bioNER formation in the upper two layers (0–2 and 2–5 mm). The mineralization of 13C6-MCPA in soil with litter increased to 27% compared to only 6% in the control. Accordingly, maize addition reduced the amount of extractable residual MCPA in soil from 77% (control) to 35% of initially applied 13C6-MCPA. While non-extractable residues (NERs) were

The objective of our study was to test whether limited microbial degradation at low pesticide concentrations could explain the discrepancy between overall degradability demonstrated in laboratory tests and their actual persistence in the environment. Studies on pesticide degradation are often performed using unrealistically high application rates seldom found in natural environments. Nevertheless, biodegradation rates determined for higher pesticide doses cannot necessarily be extrapolated to lower concentrations. In this context, we wanted to (i) compare the kinetics of pesticide degradation at different concentrations in arable land and (ii) clarify whether there is a concentration threshold below which the expression of the functional genes involved in the degradation pathway is inhibited without further pesticide degradation taking place. We set up an incubation experiment for four weeks using 14C-ring labeled 2-methyl-4-chlorophenoxyacetic acid (MCPA) as a model compound in concentrations from 30 to 20,000 μg kg–1 soil. To quantify the abundance of putative microorganisms involved in MCPA degradation and their degradation activity, tfdA gene copy numbers (DNA) and transcripts (mRNA) were determined by quantitative real-time PCR. Mineralization dynamics of MCPA derived-C were analyzed by monitoring 14CO2 production and 14C assimilation by soil microorganisms. We identified two different concentration thresholds for growth and activity with respect to MCPA degradation using tfdA gene and mRNA transcript abundance as growth and activity indices, respectively. The tfdA gene expression started to increase between 1,000 and 5,000 μg MCPA kg–1 dry soil, but an actual increase in tfdA sequences could only be determined at a concentration of 20,000 μg. Accordingly, we observed a clear shift from catabolic to anabolic utilization of MCPA-derived C in the concentration range of 1,000 to 5,000 μg kg–1. Concentrations ≥1,000 μg kg–1 were mainly associated with delayed mineralization, while concentrations ≤1,000 μg kg–1 showed rapid absolute dissipation. The persistence of pesticides at low concentrations cannot, therefore, be explained by the absence of functional gene expression. Nevertheless, significant differences in the degradation kinetics of MCPA between low and high pesticide concentrations illustrate the need for studies investigating pesticide degradation at environmentally relevant concentrations.

Spatial and temporal processes shaping microbial communities are inseparably linked but rarely studied together. By Illumina 16S rRNA sequencing, we monitored soil bacteria in 360 stations on a 100 square meter plot distributed across six intra-annual samplings in a rarely managed, temperate grassland. Using a multi-tiered approach, we tested the extent to which stochastic or deterministic processes influenced the composition of local communities. A combination of phylogenetic turnover analysis and null modeling demonstrated that either homogenization by unlimited stochastic dispersal or scenarios, in which neither stochastic processes nor deterministic forces dominated, explained local assembly processes. Thus, the majority of all sampled communities (82%) was rather homogeneous with no significant changes in abundance-weighted composition. However, we detected strong and uniform taxonomic shifts within just nine samples in early summer. Thus, community snapshots sampled from single points in time or space do not necessarily reflect a representative community state. The potential for change despite the overall homogeneity was further demonstrated when the focus shifted to the rare biosphere. Rare OTU turnover, rather than nestedness, characterized abundance-independent β-diversity. Accordingly, boosted generalized additive models encompassing spatial, temporal and environmental variables revealed strong and highly diverse effects of space on OTU abundance, even within the same genus. This pure spatial effect increased with decreasing OTU abundance and frequency, whereas soil moisture – the most important environmental variable – had an opposite effect by impacting abundant OTUs more than the rare ones. These results indicate that – despite considerable oscillation in space and time – the abundant and resident OTUs provide a community backbone that supports much higher β-diversity of a dynamic rare biosphere. Our findings reveal complex interactions among space, time, and environmental filters within bacterial communities in a long-established temperate grassland.

Phosphorus (P) is an essential nutrient, but European forest ecosystems are experiencing widespread declines in soil P concentrations. To clarify the roles of ectomycorrhizal (EM) and saprotrophic (SAP) fungi in P cycling in forest soils that differ in inorganic and organic P availability, we conducted an ingrowth tube (IGT) experiment over 18 months of exposure in five beech (Fagus sylvatica L.) forests in Germany. To separate the contributions of both fungal guilds in situ, two different types of IGTs were used: (i) open IGTs with micromesh windows to allow EM fungi to regrow into the tubes, and (ii) closed IGTs (without mesh windows) in which recolonization by EM hyphae was restricted. We then measured phosphomonoesterase, phosphodiesterase, peroxidase, and phenoloxidase activities in EM (open IGTs) and SAP (closed IGTs) dominated fungal assemblages. Shifts in fungal assemblages occurred over time, but the speed and extent of those shifts differed between the five forest sites. In three forest sites fungal community composition shifted toward greater dominance of EM fungi in open IGTs than in closed IGTs after a period of 18 months, but these changes were not apparent in two other sites. Although shifts in fungal community composition were noticeable between treatments, we found no effect on phosphomonoesterase activity. Since phosphomonoesterase is present in both fungal guilds, this indicates functional redundancy of fungi. Phosphodiesterase activity was slightly reduced in SAP dominated IGTs, indicating that SAP fungi contribute to phosphodiesterase activity to a lesser extent than do EM fungi. P cycling enzymes did not appear to have been influenced by the total P stocks of the forest sites but may have been affected by additional abiotic factors. Contrary to our expectation, phenoloxidase activity was unaffected by fungal community composition, and only peroxidase activity was higher in SAP dominated IGTs, indicating that the contribution of EM fungi to degradation of complex organic material is driven by the need to obtain nutrients in addition to C. Overall, the results of this study contribute to a better understanding of the roles of EM and SAP fungi in P cycling in forest soils.

Microplastics (MP) are pervasive in the environment. There is ample evidence of negative MP effects on biota in aquatic ecosystems, though little is known about MP effects in terrestrial ecosystems. Given numerous entry routes of MP into soils, soil organisms are likely to be exposed to MP. We compared potential toxicological effects of MP from (i) low-density polyethylene (LDPE) (mean diameter ± standard deviation: 57 ± 40 μm) and (ii) a blend of biodegradable polymers polylactide (PLA) and poly(butylene adipate-co-terephthalate) (PBAT) (40 ± 31 μm) on the reproduction and body length of the soil-dwelling bacterivorous nematode Caenorhabditis elegans. Feed suspensions without (control) or with MP (treatments) at concentrations of 1, 10, and 100 mg MP L–1 were prepared and nematodes were exposed to those suspensions on agar plates until completion of their reproductive phase (∼6 days). Using Nile red-stained PLA/PBAT MP particles and fluorescence microscopy, we demonstrated the ingestion of MP by C. elegans into pharynges and intestines. Under MP exposure, nematodes had fewer offspring (up to 22.9%) compared to nematodes in the control group. This decline was independent on the plastic type. We detected a tendency toward greater decreases in offspring at higher concentrations. Despite hints of negative effects on nematode body length under MP exposure, we could not derive a consistent pattern. We conclude that in MP-contaminated soils, the reproduction of nematodes, central actors in the soil food web, can be affected, with potentially negative implications for key soil functions, e.g., the regulation of soil biogeochemical cycles.

Crop productivity and plant physiology are affected by rising temperatures and altered precipitation patterns due to climate change. We studied the impacts of an increase in soil temperature of 2.5 °C, a decrease in summer precipitation amount of 25%, a reduction in summer precipitation frequency of 50%, and their interactions on photosynthesis, biomass production, and yield of spring barley (Hordeum vulgare L. cv. RGT Planet) in a temperate agricultural ecosystem near Stuttgart (Germany). Leaf gas exchange of barley appeared to be affected mainly by drought in the form of reduced precipitation frequency or by a combination of changes in soil temperature and precipitation patterns. In contrast, biomass production and yield parameters were more affected under soil warming alone. In addition, biomass of roots increased under soil warming at stem elongation. Stable grain yield was observed under reduced precipitation amount and also under increased evaporation through soil warming. These findings provide additional evidence that barley is relatively drought tolerant, which should be taken into consideration in the context of appropriate crop selection under climate change.

Microbial assimilation and stabilization of soil organic carbon (SOC) is an important process in global carbon cycling. For an improved understanding of climate-induced changes in ecosystem C dynamics, it is important to know the group-specific turnover of microbial C. Consequently, we wanted to answer the questions if fungi store newly assimilated C longer than bacteria and if climatic and edaphic properties of different regions affect microbial C assimilation and its subsequent release. This study presents results from a 112-day long field experiment where endogenous microorganisms from two agricultural soils were labeled with 13C labeled glucose to follow the dynamics of newly assimilated C. Both soils were representative for the respective study regions (Kraichgau and Swabian Alb). Whereas, microbial assimilation of newly added C was higher in Kraichgau than in Swabian Alb, the opposite result was obtained for the mean residence time (MRT) of microbial biomass C (76 days in Kraichgau and 93 days in Swabian Alb). The accelerated turnover rates of microbial C in the warmer soil of Kraichgau with lower clay content might be an important mechanism explaining the differences in SOC content between both regions. Gram-positive bacteria assimilated more 13C-glucose into their biomass than fungi and the MRT of C was higher in bacteria as compared to fungi in both regions. Beside these dynamic substrate utilization strategies, we observed possible cross feeding by gram-negative bacteria. Carbon MRT in fungi was region specific and was best represented by a two-pool model; the initial MRT ranged between 5 and 39 days and was, in the end, higher than 4 years. This hints toward a functional separation of the fungal community; fast growing fungi dominant in the early phase and internal redistribution of C in the second phase of decomposition. Our study identified microbial group and region specific MRT of freshly assimilated C as an important parameter, which might help to explain the commonly found variation in soil respiration and SOC stabilization.

Trait-based models have improved the understanding and prediction of soil organic matter dynamics in terrestrial ecosystems. Microscopic observations and pore scale models are now increasingly used to quantify and elucidate the effects of soil heterogeneity on microbial processes. Combining both approaches provides a promising way to accurately capture spatial microbial-physicochemical interactions and to predict overall system behavior. The present study aims to quantify controls on carbon (C) turnover in soil due to the mm-scale spatial distribution of microbial decomposer communities in soil. A new spatially explicit trait-based model (SpatC) has been developed that captures the combined dynamics of microbes and soil organic matter (SOM) by taking into account microbial life-history traits and SOM accessibility. Samples of spatial distributions of microbes at μm-scale resolution were generated using a spatial statistical model based on Log Gaussian Cox Processes which was originally used to analyze distributions of bacterial cells in soil thin sections. These μm-scale distribution patterns were then aggregated to derive distributions of microorganisms at mm-scale. We performed Monte-Carlo simulations with microbial distributions that differ in mm-scale spatial heterogeneity and functional community composition (oligotrophs, copiotrophs, and copiotrophic cheaters). Our modeling approach revealed that the spatial distribution of soil microorganisms triggers spatiotemporal patterns of C utilization and microbial succession. Only strong spatial clustering of decomposer communities induces a diffusion limitation of the substrate supply on the microhabitat scale, which significantly reduces the total decomposition of C compounds and the overall microbial growth. However, decomposer communities act as functionally redundant microbial guilds with only slight changes in C utilization. The combined statistical and process-based modeling approach derives distribution patterns of microorganisms at the mm-scale from microbial biogeography at microhabitat scale (μm) and quantifies the emergent macroscopic (cm) microbial and C dynamics. Thus, it effectively links observable process dynamics to the spatial control by microbial communities. Our study highlights a powerful approach that can provide further insights into the biological control of soil organic matter turnover.

The search for approaches to a holistic sustainable agriculture requires the development of new cropping systems that provide additional ecosystem services beyond biomass supply for food, feed, material, and energy use. The reduction of chemical synthetic plant protection products is a key instrument to protect vulnerable natural resources such as groundwater and biodiversity. Together with an optimal use of mineral fertilizer, agroecological practices, and precision agriculture technologies, a complete elimination of chemical synthetic plant protection in mineral-ecological cropping systems (MECSs) may not only improve the environmental performance of agroecosystems, but also ensure their yield performance. Therefore, the development of MECSs aims to improve the overall ecosystem services of agricultural landscapes by (i) improving the provision of regulating ecosystem services compared to conventional cropping systems and (ii) improving the supply of provisioning ecosystem services compared to organic cropping systems. In the present review, all relevant research levels and aspects of this new farming concept are outlined and discussed based on a comprehensive literature review and the ongoing research project “Agriculture 4.0 without Chemical-Synthetic Plant Protection”.

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 10 m2 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 m. 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.

Increasing air and soil temperatures and changes in precipitation patterns as consequences of climate change will affect crop production in agricultural ecosystems. The combined effects of soil warming and altered precipitation on the productivity and product quality of oil crops are not yet well studied. Winter oilseed rape (OSR) (Brassica napus L., cv. Mercedes) was field-grown under elevated soil temperature (+2.5 °C), reduced precipitation amount (−25%), reduced precipitation frequency (−50%) both separately and in combination in order to investigate effects on crop development, seed yield, and seed quality. Soil warming accelerated crop development during early plant growth and during spring. At maturity, however, plants in all treatments were similar in quantitative (aboveground biomass, seed yield) and qualitative (protein and oil content, amino acids, fatty acids) parameters. We observed the long-term effects of the precipitation manipulation on leaf size, leaf senescence and biomass allocation. Seed yield was not affected by the altered climatic factors, perhaps due to adaptation of soil microorganisms to permanent soil warming and to relatively wet conditions during the seed-filling period. Overall, OSR performed well under moderate changes in soil temperature and precipitation patterns; thus, we observed stable seed yield without negative impacts on nutritive seed quality.

Arbuscular mycorrhizal fungi (AMF) are important symbionts for plant nutrient uptake, but their exact role in plant nitrogen (N) nutrition is unclear. Protists on the other hand play an acknowledged role in plant N acquisition, and there is increasing evidence for a close interaction with AMF. In a split root set up, we investigated the distinct roles of mycorrhiza (Rhizophagus irregularis), protists (Acanthamoeba castellanii), and their interaction on plant N uptake, within-root system allocation patterns, and shoot-to-root ratio of winter wheat. In addition, we applied a quantitative metabolomics approach to characterize associated changes in soil microbial communities by microbial phospholipid fatty acid (PLFA) analysis from rhizosphere soil. AMF markedly altered plant shoot-to-root allometry by reducing root biomass of wheat, and mycorrhiza partly took over root system functioning. Protists promoted shoot and root growth, and improved plant N uptake by the release of N from consumed bacterial biomass, a mechanism known as microbial loop. The shoot system however responded little to these alterations of the root system and of the rhizosphere community composition, indicating that the plants optimized shoot growth despite varying investment into roots. Mycorrhiza reduced root biomass and plant N, especially in the combined treatments with protists by changing within root system allocation of N and root biomass. These systemic effects on root allocation pattern suggest that mycorrhiza also gained control over N provided by protist grazers. Protists and mycorrhiza altered rhizosphere bacterial communities in contrasting but consistent ways as shown by quantitative shifts in microbial PLFA profiles. Remarkably, the changes in bacterial community composition were systemically conveyed within the root system to the split-root chamber where the symbionts were lacking. Accordingly the synergistic effects of protists and mycorrhiza indicated systemic effects on nutrient- and on root-allocation within root systems as an emergent property that could not be predicted from single treatments with mycorrhiza or protists alone. The tight plant and microbial feed backs uncovered in this study have far reaching implications for understanding the assembly of plant microbiomes, and testify central roles of both protists and mycorrhizas in the assembly process.

Phosphorus (P) is an important nutrient, whose plant-available form phosphate is often low in natural forest ecosystems. Mycorrhizal fungi mine the soil for P and supply their host with this resource. It is unknown how ectomycorrhizal communities respond to changes in P availability. Here, we used young beech (Fagus sylvatica L.) trees in natural forest soil from a P-rich and P-poor site to investigate the impact of P amendment on soil microbes, mycorrhizas, beech P nutrition, and photosynthesis. We hypothesized that addition of P to forest soil increased P availability, thereby, leading to enhanced microbial biomass and mycorrhizal diversity in P-poor but not in P-rich soil. We expected that P amendment resulted in increased plant P uptake and enhanced photosynthesis in both soil types. Young beech trees with intact soil cores from a P-rich and a P-poor forest were kept in a common garden experiment and supplied once in fall with triple superphosphate. In the following summer, labile P in the organic layer, but not in the mineral top soil, was significantly increased in response to fertilizer treatment. P-rich soil contained higher microbial biomass than P-poor soil. P treatment had no effect on microbial biomass but influenced the mycorrhizal communities in P-poor soil and shifted their composition toward higher similarities to those in P-rich soil. Plant uptake efficiency was negatively correlated with the diversity of mycorrhizal communities and highest for trees in P-poor soil and lowest for fertilized trees. In both soil types, radioactive P tracing (H333PO4) revealed preferential aboveground allocation of new P in fertilized trees, resulting in increased bound P in xylem tissue and enhanced soluble P in bark, indicating increased storage and transport. Fertilized beeches from P-poor soil showed a strong increase in leaf P concentrations from deficient to luxurious conditions along with increased photosynthesis. Based on the divergent behavior of beech in P-poor and P-rich forest soil, we conclude that acclimation of beech to low P stocks involves dedicated mycorrhizal community structures, low P reserves in storage tissues and photosynthetic inhibition, while storage and aboveground allocation of additional P occurs regardless of the P nutritional status.

Soil food web structure and function is primarily determined by the major basal resources, which are living plant tissue, root exudates and dead organic matter. A field experiment was performed to disentangle the interlinkage of the root-and detritus-based soil food chains. An arable site was cropped either with maize, amended with maize shoot litter or remained bare soil, representing food webs depending on roots, aboveground litter and soil organic matter as predominant resource, respectively. The soil micro-food web, i.e. microorganisms and nematodes, was investigated in two successive years along a depth transect. The community composition of nematodes was used as model to determine the changes in the rhizosphere, detritusphere and bulk soil food web. In the first growing season the impact of treatments on the soil micro-food web was minor. In the second year plant-feeding nematodes increased under maize, whereas after harvest the Channel Index assigned promotion of the detritivore food chain, reflecting decomposition of root residues. The amendment with litter did not foster microorganisms, instead biomass of Gram-positive and Gram-negative bacteria as well as that of fungi declined in the rooted zone. Likely higher grazing pressure by nematodes reduced microbial standing crop as bacterial and fungal feeders increased. However, populations at higher trophic levels were not promoted, indicating limited flux of litter resources along the food chain. After two years of bare soil microbial biomass and nematode density remained stable, pointing to soil organic matter-based resources that allow bridging periods with deprivation. Nematode communities were dominated by opportunistic taxa that are competitive at moderate resource supply. In sum, removal of plants from the system had less severe effects than expected, suggesting considerable food web resilience to the disruption of both the root and detrital carbon channel, pointing to a legacy of organic matter resources in arable soils.

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.

The benefit of plant growth-promoting microorganisms (PGPMs) as plant inoculants is influenced by a wide range of environmental factors. Therefore, microbial consortia products (MCPs) based on multiple PGPM strains with complementary functions, have been proposed as superior, particularly under challenging environmental conditions and for restoration of beneficial microbial communities in disturbed soil environments. To test this hypothesis, the performance of a commercial MCP inoculant based on 22 PGPM strains was investigated in greenhouse experiments with maize on three soils with contrasting pH, organic matter content and microbial activity, under different P and N fertilization regimes. Interestingly, the MCP inoculant stimulated root and shoot growth and improved the acquisition of macronutrients only on a freshly collected field soil with high organic matter content, exclusively in combination with stabilized ammonium fertilization. This was associated with transiently increased expression of AuxIAA5 in the root tissue, a gene responsive to exogenous auxin supply, suggesting root growth promotion by microbial auxin production as a major mode of action of the MCP inoculant. High microbial activity was indicated by intense expression of soil enzyme activities involved in C, N and P cycling in the rhizosphere (cellulase, leucine peptidase, alkaline and acid phosphatases) but without MCP effects. By contrast, the MCP inoculation did not affect maize biomass production or nutrient acquisition on soils with very little Corg and low microbial activity, although moderate stimulation of rhizosphere enzymes involved in N and P cycling was recorded. There was also no indication for MCP-induced solubilization of Ca-phosphates on a calcareous sub-soil fertilized with rock-phosphate. The results demonstrate that the combination of multiple PGPM strains with complementary properties as MCP inoculants does not necessarily translate into plant benefits in challenging environments. Thus, a better understanding of the conditions determining successful MCP application is mandatory.

Many arable lands have accumulated large reserves of residual phosphorus (P) and a relatively large proportion of soil P is less available for uptake by plants. Root released organic anions are widely documented as a key physiological strategy to enhance P availability, while limited information has been generated on the contribution of rhizosphere organic anions to P utilization by crops grown in agricultural soils that are low in available P and high in extractable Ca, Al and Fe. We studied the role of rhizosphere organic anions in P uptake from residual P in four common crops Triticum aestivum, Avena sativa, Solanum tuberosum and Brassica napus in low- and high-P availability agricultural soils from long-term fertilization field trials in a mini-rhizotron experiment with four replications. Malate was generally the dominant organic anion. More rhizosphere citrate was detected in low P soils than in high P soil. Brassica napus showed 74-103% increase of malate in low P loam, compared with clay loam. Avena sativa had the greatest rhizosphere citrate concentration in all soils (5.3-15.2 mol g-1 root DW). Avena sativa also showed the highest level of root colonization by arbuscular mycorrhizal fungi (36% and 40%), the greatest root mass ratio (0.51 and 0.66) in the low-P clay loam and loam respectively, and the greatest total P uptake (5.92 mg P/mini-rhizotron) in the low-P loam. Brassica napus had 15-44% more rhizosphere APase activity, ~0.1-0.4 units lower rhizosphere pH than other species, the greatest increase in rhizosphere water-soluble P in the low-P soils, and the greatest total P uptake in the low-P clay loam. Shoot P content was mainly explained by rhizosphere APase activity, water-soluble P and pH within low P soils across species. Within species, P uptake was mainly linked to rhizosphere water soluble P, APase and pH in low P soils. The effects of rhizosphere organic anions varied among species and they appeared to play minor roles in improving P availability and uptake.

The flow of plant-derived carbon in soil is a key component of global carbon cycling. Conceptual models of trophic carbon fluxes in soil have assumed separate bacterial and fungal energy channels in the detritusphere, controlled by both substrate complexity and recalcitrance. However, detailed understanding of the key populations involved and niche-partitioning between them is limited. Here, a microcosm experiment was performed to trace the flow of detritusphere C from substrate analogues (glucose, cellulose) and plant biomass amendments (maize leaves, roots) in an agricultural soil. Carbon flow was traced by rRNA stable isotope probing and amplicon sequencing across three microbial kingdoms. Distinct lineages within the Actinobacteria, Bacteroidetes, Gammaproteobacteria, Basidiomycota, Ascomycota as well as Peronosporomycetes were identified as important primary substrate consumers. A dynamic succession of primary consumers was observed especially in the cellulose treatments, but also in plant amendments over time. While intra-kingdom niche partitioning was clearly observed, distinct bacterial and fungal energy channels were not apparent. Furthermore, while the diversity of primary substrate consumers did not notably increase with substrate complexity, consumer succession and secondary trophic links to bacterivorous and fungivorous microbes resulted in increased food web complexity in the more recalcitrant substrates. This suggests that rather than substrate-defined energy channels, consumer succession as well as intra- and inter-kingdom cross-feeding should be considered as mechanisms supporting food web complexity in the detritusphere.

Understanding factors driving the ecology of N cycling microbial communities is of central importance for sustainable land use. In this study we report changes of abundance of denitrifiers, nitrifiers and nitrogen-fixing microorganisms (based on qPCR data for selected functional genes) in response to different land use intensity levels and the consequences for potential turnover rates. We investigated selected grassland sites being comparable with respect to soil type and climatic conditions, which have been continuously treated for many years as intensely used meadows (IM), intensely used mown pastures (IP) and extensively used pastures (EP), respectively. The obtained data were linked to above ground biodiversity pattern as well as water extractable fractions of nitrogen and carbon in soil. Shifts in land use intensity changed plant community composition from systems dominated by s-strategists in extensive managed grasslands to c-strategist dominated communities in intensive managed grasslands. Along the different types of land use intensity, the availability of inorganic nitrogen regulated the abundance of bacterial and archaeal ammonia oxidizers. In contrast, the amount of dissolved organic nitrogen determined the abundance of denitrifiers (nirS and nirK). The high abundance of nifH carrying bacteria at intensive managed sites gave evidence that the amounts of substrates as energy source outcompete the high availability of inorganic nitrogen in these sites. Overall, we revealed that abundance and function of microorganisms involved in key processes of inorganic N cycling (nitrification, denitrification and N fixation) might be independently regulated by different abiotic and biotic factors in response to land use intensity.

Very few principles have been unraveled that explain the relationship between soil properties and soil biota across large spatial scales and different land-use types. Here, we seek these general relationships using data from 52 differently managed grassland and forest soils in three study regions spanning a latitudinal gradient in Germany. We hypothesize that, after extraction of variation that is explained by location and land-use type, soil properties still explain significant proportions of variation in the abundance and diversity of soil biota. If the relationships between predictors and soil organisms were analyzed individually for each predictor group, soil properties explained the highest amount of variation in soil biota abundance and diversity, followed by land-use type and sampling location. After extraction of variation that originated from location or land-use, abiotic soil properties explained significant amounts of variation in fungal, meso- and macrofauna, but not in yeast or bacterial biomass or diversity. Nitrate or nitrogen concentration and fungal biomass were positively related, but nitrate concentration was negatively related to the abundances of Collembola and mites and to the myriapod species richness across a range of forest and grassland soils. The species richness of earthworms was positively correlated with clay content of soils independent of sample location and land-use type. Our study indicates that after accounting for heterogeneity resulting from large scale differences among sampling locations and land-use types, soil properties still explain significant proportions of variation in fungal and soil fauna abundance or diversity. However, soil biota was also related to processes that act at larger spatial scales and bacteria or soil yeasts only showed weak relationships to soil properties. We therefore argue that more general relationships between soil properties and soil biota can only be derived from future studies that consider larger spatial scales and different land-use types.