Publications in peer reviewed journals

47 Publications found
  • Fine roots stimulate nutrient release during early stages of leaf litter decomposition in a Central Amazon rainforest

    Martins NP, Fuchslueger L, Fleischer K, Andersen KM, Assis RL, Baccaro FB, Camargo PB, Cordeiro AL, Grandis A, Hartley IP, Hofhansl F, Lugli LF, Lapola DM, Menezes JG, Norby RJ, Rammig A, Rosa JS, Schaap KJ, Takeshi B, Valverde-Barrantes OJ, Quesada CA
    2021 - Plant and soil, 469: 287-303



    Large parts of the Amazon rainforest grow on weathered soils depleted in phosphorus and rock-derived cations. We tested the hypothesis that in this ecosystem, fine roots stimulate decomposition and nutrient release from leaf litter biochemically by releasing enzymes, and by exuding labile carbon stimulating microbial decomposers.


    We monitored leaf litter decomposition in a Central Amazon tropical rainforest, where fine roots were either present or excluded, over 188 days and added labile carbon substrates (glucose and citric acid) in a fully factorial design. We tracked litter mass loss, remaining carbon, nitrogen, phosphorus and cation concentrations, extracellular enzyme activity and microbial carbon and nutrient concentrations.


    Fine root presence did not affect litter mass loss but significantly increased the loss of phosphorus and cations from leaf litter. In the presence of fine roots, acid phosphatase activity was 43.2% higher, while neither microbial stoichiometry, nor extracellular enzyme activities targeting carbon- and nitrogen-containing compounds changed. Glucose additions increased phosphorus loss from litter when fine roots were present, and enhanced phosphatase activity in root exclusions. Citric acid additions reduced litter mass loss, microbial biomass nitrogen and phosphorus, regardless of fine root presence or exclusion.


    We conclude that plant roots release significant amounts of acid phosphatases into the litter layer and mobilize phosphorus without affecting litter mass loss. Our results further indicate that added labile carbon inputs (i.e. glucose) can stimulate acid phosphatase production by microbial decomposers, highlighting the potential importance of plant-microbial feedbacks in tropical forest ecosystems.
  • Bypass and hyperbole in soil science: A perspective from the next generation of soil scientists

    Portell X,  Sauzet O,  Balseiro-Romero M,  Benard P,  Cardinael R,  Couradeau E,  Danra DD,  Evans DL,  Fry EL, Hammer E,  Mamba D,  Merino-Martín L,  Mueller CW, Paradelo M, Rees F,  Rossi M, Schmidt H,  Schnee LS,  Védère C, Vidal A
    2021 - European Journal of Soil Science, 72: 31-34
  • Cyanate is a low abundance but actively cycled nitrogen compound in soil

    Mooshammer M, Wanek W, Jones S, Richter A, Wagner W
    2021 - Communications Earth & Environment, 2: Article 161


    Cyanate can serve as a nitrogen and/or carbon source for different microorganisms and as an energy source for autotrophic ammonia oxidizers. However, the extent of cyanate availability and utilisation in terrestrial ecosystems and its role in biogeochemical cycles is poorly known. Here we analyse cyanate concentrations in soils across a range of soil types, land management practices and climates. Soil cyanate concentrations were three orders of magnitude lower than ammonium or nitrate. We determined cyanate consumption in a grassland and rice paddy soil using stable isotope tracer experiments. We find that cyanate turnover was rapid and dominated by biotic processes. We estimated that in-situ cyanate production rates were similar to those associated with urea fertilizer decomposition, a major source of cyanate in the environment. We provide evidence that cyanate is actively turned over in soils and represents a small but continuous nitrogen/energy source for soil microbes.

  • Arctic Tundra Land Cover Classification on the Beaufort Coast Using the Kennaugh Element Framework on Dual-Polarimetric TerraSAR-X Imagery

    A’Campo W, Bartsch A, Roth A, Wendleder A, Martin VS, Durstewitz L, Lodi R, Wagner J, Hugelius G
    2021 - Remote Sensing, 13: Article 4780


    Arctic tundra landscapes are highly complex and are rapidly changing due to the warming climate. Datasets that document the spatial and temporal variability of the landscape are needed to monitor the rapid changes. Synthetic Aperture Radar (SAR) imagery is specifically suitable for monitoring the Arctic, as SAR, unlike optical remote sensing, can provide time series regardless of weather and illumination conditions. This study examines the potential of seasonal backscatter mechanisms in Arctic tundra environments for improving land cover classification purposes by using a time series of HH/HV TerraSAR-X (TSX) imagery. A Random Forest (RF) classification was applied on multi-temporal Sigma Nought intensity and multi-temporal Kennaugh matrix element data. The backscatter analysis revealed clear differences in the polarimetric response of water, soil, and vegetation, while backscatter signal variations within different vegetation classes were more nuanced. The RF models showed that land cover classes could be distinguished with 92.4% accuracy for the Kennaugh element data, compared to 57.7% accuracy for the Sigma Nought intensity data. Texture predictors, while improving the classification accuracy on the one hand, degraded the spatial resolution of the land cover product. The Kennaugh elements derived from TSX winter acquisitions were most important for the RF model, followed by the Kennaugh elements derived from summer and autumn acquisitions. The results of this study demonstrate that multi-temporal Kennaugh elements derived from dual-polarized X-band imagery are a powerful tool for Arctic tundra land cover mapping.

  • The effect of salinity, light regime and food source on C and N uptake in a benthic foraminifera

    Lintner M, Lintner B, Wanek W, Keul N, Heinz P
    2021 - Biogeosciences, 18: 1395–1406


    Foraminifera are unicellular organisms that play an important role in marine organic matter cycles. Some species are able to isolate chloroplasts from their algal food source and incorporate them as kleptoplasts into their own metabolic pathways, a phenomenon known as kleptoplastidy. One species showing this ability is Elphidium excavatum, a common foraminifer in the Kiel Fjord, Germany. The Kiel Fjord is fed by several rivers and thus forms a habitat with strongly fluctuating salinity. Here, we tested the effects of the food source, salinity and light regime on the food uptake (via 15N and 13C algal uptake) in this kleptoplast-bearing foraminifer. In our study E. excavatum was cultured in the lab at three salinity levels (15, 20 and 25) and uptake of C and N from the food source Dunaliella tertiolecta (Chlorophyceae) and Leyanella arenaria (Bacillariophyceae) were measured over time (after 3, 5 and 7 d). The species was very well adapted to the current salinity of the sampling region, as both algal N and C uptake was highest at a salinity of 20. It seems that E. excavatum coped better with lower than with higher salinities. The amount of absorbed C from the green algae D. tertiolecta showed a tendency effect of salinity, peaking at a salinity of 20. Nitrogen uptake was also highest at a salinity of 20 and steadily increased with time. In contrast, C uptake from the diatom L. arenaria was highest at a salinity of 15 and decreased at higher salinities. We found no overall significant differences in C and N uptake from green algae vs. diatoms. Furthermore, the food uptake at a light–dark rhythm of 16:8h was compared to continuous darkness. Darkness had a negative influence on algal C and N uptake, and this effect increased with incubation time. Starving experiments showed a stimulation of food uptake after 7 d. In summary, it can be concluded that E. excavatum copes well with changes of salinity to a lower level. For changes in light regime, we showed that light reduction caused a decrease of C and N uptake by E. excavatum.

  • Increased microbial expression of organic nitrogen cycling genes in long-term warmed grassland soils

    Séneca J, Söllinger A, Herbold C, Pjevac P, Prommer J, Verbruggen E, Sigurdssaon BD, Peñuelas J, Janssens IA, Urich T, Tveit AT, Richter A
    2021 - ISME Communications, 1: Article 69


    Global warming increases soil temperatures and promotes faster growth and turnover of soil microbial communities. As microbial cell walls contain a high proportion of organic nitrogen, a higher turnover rate of microbes should also be reflected in an accelerated organic nitrogen cycling in soil. We used a metatranscriptomics and metagenomics approach to demonstrate that the relative transcription level of genes encoding enzymes involved in the extracellular depolymerization of high-molecular-weight organic nitrogen was higher in medium-term (8 years) and long-term (>50 years) warmed soils than in ambient soils. This was mainly driven by increased levels of transcripts coding for enzymes involved in the degradation of microbial cell walls and proteins. Additionally, higher transcription levels for chitin, nucleic acid, and peptidoglycan degrading enzymes were found in long-term warmed soils. We conclude that an acceleration in microbial turnover under warming is coupled to higher investments in N acquisition enzymes, particularly those involved in the breakdown and recycling of microbial residues, in comparison with ambient conditions.

  • Isotopic Elucidation of Microbial Nitrogen Transformations in Forest Soils

    Xu S-Q, Liu X-Y, Sun Z-C, Hu C-C, Wanek W, Koba K
    2021 - Global Biogeochemical Cycles, 35: Article e2021GB00707


    Soil nitrogen (N) transformations between labile N forms (extractable organic N [EON], ammonium [NH4+], and nitrate [NO3]) regulate soil N availability. However, it has long been difficult to quantify the transformations of total soil organic and labile N forms in soils, which has left large uncertainties in evaluating atmospheric N deposition effects on soil N dynamics. Based on concentrations and natural abundances of N isotopes of soil organic N, EON, NH4+, and NO3 across 11 forests with variant N deposition levels, we established a quantitative isotopic framework to estimate the fractions of soil N depolymerization (fD), mineralization (fM), nitrification (fN), and of NO3 losses (fL) via denitrification and leaching. Based on the fractions, the gross production and storage of corresponding soil labile N were estimated for forests of China and Japan. We found that fDfM, and fN increased, while fL decreased with the increase of N deposition among the study forests. And the contribution of denitrification (relative to the NO3 leaching) to total NO3 losses also increased with increasing N deposition. Our method provides new and straightforward insights into the present soil N transformations and allows to evaluate the soil N status. These findings are useful for modeling forest N cycles under different N deposition regimes.

  • Leaf trait co-variation and trade-offs in gallery forest C3 and CAM epiphytes

    Oliveira RdP, Zotz G, Wanek W, Franco AC
    2021 - Biotropica, 3: 520–535


    Despite their unique adaptations to thrive in canopy environments without access to soil resources, epiphytes are underrepresented in studies of functional traits and of functional composition of tropical plant communities. We investigated functional traits of spermatophytic (seed‐bearing) C3 and CAM epiphyte communities in flooded and non‐flooded gallery forests in Central Brazil. The two forest types differ in floristic, structure, microclimate, and edaphic conditions. We studied plant size, leaf thickness, leaf dry matter content (LDMC), leaf water content, leaf area (LA), specific leaf area (SLA), leaf C, N, P, K, Mg, and Ca, and stable isotope ratios (δ13C and δ15N). Because photosynthetic pathway (C3 or CAM) is an important aspect of ecological differentiation for spermatophytic epiphytes, we expected that functional trait syndromes in a multivariate space would be more associated with photosynthetic pathway than forest type, and changes in abundance of C3 and CAM epiphytes would drive functional trait composition at the community level. C3 and CAM epiphytes segregated in the multivariate trait space; however, more complex functional typologies were also evident. Despite lower light levels, CAM epiphytes were more abundant in the flooded gallery forest. There, they accounted for 80% of all individuals, whereas C3 epiphytes dominated in the non‐flooded forest. These large differences in the proportion of CAM and C3 epiphytes strongly affected functional trait values at the community level, despite very little intraspecific variation in trait values between forest types for species that occurred in both forests.

  • Evidence for seasonal changes in population structure of Reed Buntings (Emberiza schoeniclus) in the Lake Neusiedl Region

    Kofler B,  Wanek W,  Voigt CC, Schulze CH
    2021 - Journal of Ornithology, 162: 421-433


    Migrating species and populations are affected by conditions existing across large geographical scales. The degree to which populations mix at stop-over sites and wintering grounds is variable and can involve important fitness consequences. Thus, the link between breeding and wintering grounds and seasonal shifts in population structure are important to understand population demographics, local adaption and diversification patterns. This work aims to infer migration patterns and population connectivity in the Reed Bunting (Emberiza schoeniclus) community of a Ramsar-designated wetland, Lake Neusiedl, Seewinkel, Austria. Stable isotope markers (hydrogen, δ2H, carbon, δ13C) in feathers were analyzed to assign individuals to a specific breeding or natal area. Based on the geographic assignment, birds were compared with respect to their ecomorphological traits and their habitat selection (large reed bed versus more open habitat at shore of an alkaline lake) during winter to assess potential ecological differences and possibly associated morphological divergence. Stable isotope data revealed that the local Reed Bunting community undergoes seasonal changes in population structure. Three different populations that vary in their δ2H signature and to some extent in morphological characteristics as well as in δ13C values were identified. Indications for differential migration related to age and/or sex were additionally found, since males and females as well as first-year birds and adults showed remarkable differences in their traits and feather isotope values. δ2H values diverged significantly between these groups in both age and sex classes and birds with highest feather δ2H values were considered to represent the local breeding population at Lake Neusiedl and possibly also Reed Buntings from close surroundings. Moreover, this group had the highest δ13C values in first-year individuals as well as in adults. In both sex classes, the assignment of Reed Buntings to different populations was ultimately possible due to differences in bill shape with residents being characterized by more massive bills than individuals belonging to populations visiting the study area only during migration or for wintering. During winter, beak shape and feather δ2H and δ13C values differed significantly between birds caught in the reed stand of Lake Neusiedl and those captured near the salt pan, hence, indicating differential habitat selection among co-occurring populations during winter. These findings suggest that morphological attributes, such as bill shape and size as well as linked adaptations to different food sources play a decisive role for niche segregation among wintering populations. The results provide evidence that different populations mix during migration and during winter at Lake Neusiedl and the extent to which they are specialized to a specific habitat varied.

  • Prevalence of RT-qPCR-detected SARS-CoV-2 infection at schools: First results from the Austrian School-SARS-CoV-2 prospective cohort study

    Willeit P, Krause R, Lamprecht B, Berghold A, Hanson B, Stelzl E, Stoiber H, Zuber J, Heinen R, Köhler A, Bernhard D, Borena W, Doppler C, von Laer D, Schmidt H, Pröll J, Steinmetz I, Wagner M
    2021 - The Lancet Regional Health - Europe, 5: Article 100086


    Background The role of schools in the SARS-CoV-2 pandemic is much debated. We aimed to quantify reliably the prevalence of SARS-CoV-2 infections at schools detected with reverse-transcription quantitative polymerase-chain-reaction (RT-qPCR).

    Methods This nationwide prospective cohort study monitors a representative sample of pupils (grade 1-8) and teachers at Austrian schools throughout the school year 2020/2021. We repeatedly test participants for SARS-CoV-2 infection using a gargling solution and RT-qPCR. We herein report on the first two rounds of examinations. We used mixed-effect logistic regression to estimate odds ratios and robust 95% confidence intervals (95% CI).

    Findings We analysed data on 10734 participants from 245 schools (9465 pupils, 1269 teachers). Prevalence of SARS-CoV-2 infection increased from 0.39% at round 1 (95% CI 0.28-0·55%, 29 September-22 October 2020) to 1·39% at round 2 (95% CI 1·04-1·85%, 10-16 November). Odds ratios for SARS-CoV-2 infection were 2·26 (95% CI 1·25-4·12, P=0·007) in regions with >500 vs. ≤500 inhabitants/km2, 1·67 (95% CI 1·42-1·97, P<0·001) per two-fold higher regional 7-day incidence, and 2·78 (95% CI 1·73-4·48, P<0·001) in pupils at schools with high/very high vs. low/moderate social deprivation. Associations of community incidence and social deprivation persisted in a multivariable adjusted model. Prevalence did not differ by average number of pupils per class nor between age groups, sexes, pupils vs. teachers, or primary (grade 1-4) vs. secondary schools (grade 5-8).

    Interpretation This monitoring study in Austrian schools revealed SARS-CoV-2 infection in 0·39%-1·39% of participants and identified associations of regional community incidence and social deprivation with higher prevalence.

    Funding BMBWF Austria.

  • Editorial: Exchanges at the Root-Soil Interface: Resource Trading in the Rhizosphere That Drives Ecosystem Functioning

    Preece K, Canarin A, Verbruggen E, Fuchslueger L
    2021 - Frontiers in Forests and Global Change, 4: Article 747492
  • Litter inputs and phosphatase activity affect the temporal variability of organic phosphorus in a tropical forest soil in the Central Amazon

    Schaap KJ,  Fuchslueger L, Hoosbeek MR,  Hofhansl F, Martins NP, Valverde-Barrantes OJ, Hartley I, Lugli LF, Quesada CA
    2021 - Plant and soil, 469: 423-441



    The tropical phosphorus cycle and its relation to soil phosphorus (P) availability are a major uncertainty in projections of forest productivity. In highly weathered soils with low P concentrations, plant and microbial communities depend on abiotic and biotic processes to acquire P. We explored the seasonality and relative importance of drivers controlling the fluctuation of common P pools via processes such as litter production and decomposition, and soil phosphatase activity.


    We analyzed intra-annual variation of tropical soil phosphorus pools using a modified Hedley sequential fractionation scheme. In addition, we measured litterfall, the mobilization of P from litter and soil extracellular phosphatase enzyme activity and tested their relation to fluctuations in P- fractions.


    Our results showed clear patterns of seasonal variability of soil P fractions during the year. We found that modeled P released during litter decomposition was positively related to change in organic P fractions, while net change in organic P fractions was negatively related to phosphatase activities in the top 5 cm.


    We conclude that input of P by litter decomposition and potential soil extracellular phosphatase activity are the two main factors related to seasonal soil P fluctuations, and therefore the P economy in P impoverished soils. Organic soil P followed a clear seasonal pattern, indicating tight cycling of the nutrient, while reinforcing the importance of studying soil P as an integrated dynamic system in a tropical forest context.

  • Impact of Nutrient Additions on Free-Living Nitrogen Fixation in Litter and Soil of Two French-Guianese Lowland Tropical Forests

    Van Langenhove L, Depaepe T, Verryckt LT, Vallicrosa H, Fuchslueger L, Lugli LF, Bréchet L, Ogaya R, Llusià J, Urbina I, Gargallo-Garriga A, Grau O, Richter A, Peñuelas J, Van Der Straeten D, Janssens IA
    2021 - Journal of Geophysical Research: Biogeosciences, 126: Article e2020JG00602


    Lakes receive and transform significant amounts of terrestrial carbon and are often considered a source of atmospheric carbon dioxide (CO2). Yet, continuous direct measurements of lake-atmosphere CO2 exchange with high temporal resolution are sparse. In this study, we measured the CO2 exchange of a mountain lake in the eastern Austrian Alps continuously for one year using the eddy covariance (EC) and the boundary layer model (BLM) approaches. Results from both the EC and the BLM methods indicated the lake to be a small source of atmospheric CO2 with highest emissions in fall. EC flux measurements were affected by low-frequency contributions especially during low wind conditions. The CO2 concentration gradient at the air-water interface decreased during night-time due to an increase in atmospheric CO2 above the lake, likely caused by cold and CO2-rich air draining from the surrounding land. Consequently, BLM fluxes were lower during night-time than during daytime. This diel pattern was lacking in the EC flux measurements because the EC instruments deployed at the shore of the lake did not capture low nocturnal lake CO2 fluxes due to the local wind regime. Overall, this study illustrates the effect of the surrounding landscape on lake-atmosphere flux measurements. We conclude that estimating CO2 evasion from lakes situated in complex topography needs to explicitly account for biases in EC flux measurements caused by low-frequency contributions and local wind regimes.

  • Genomic insights into diverse bacterial taxa that degrade extracellular DNA in marine sediments 

    Wasmund K, Pelikan C, Schintlmeister A, Wagner M, Watzka M, Richter A, Bhatnagar S, Noel A, Hubert CRJ, Rattei T, Hofmann T, Hausmann B, Herbold CW, Loy A
    2021 - Nature Microbiology, 6: 885-898


    While best known as the code for genetic information, DNA is also a nutrient for specialised microbes. An international team of researchers led by Ken and Alex from DOME has discovered several bacteria in sediment samples from the Atlantic Ocean that use DNA as a food source. One bacterium newly named by the team in fact is a true expert in degrading DNA. The study provides new insights into the diversity and ecological function of the still largely undescribed microscopic world in the oceans.

  • Tradeoffs and synergies in tropical root traits linked to nutrient and water acquisition

    Cusack D, Addo-Danso SD, Agee EA, Andersen KM, Arnaud M, Batterman SA, Brearley F, Ciochina MI, Cordeiro AL, Diaz-Toribio MH, Dietterich LH, Fisher JB, Fleischer K, Fortunel C, Fuchslueger L, Guerrero-Ramírez NR, Kotowska M, Lugli LF, Marín C, McCulloch LA, Maeght JL, Metcalf D, Norby RJ, Oliveira R, Powers JS, Reichert T, Smith SW, Smith-Martin CM, Soper FM, Toro L, Umana MN, Vlaverde-Barrantes OJ, Weemstra M, Werden LK, Wong M, Wright SJ, Yaffar D
    2021 - Frontiers in Global Change – Forest soils, 4: Article 704469


    Vegetation processes are fundamentally limited by nutrient and water availability, the uptake of which is mediated by plant roots in terrestrial ecosystems. While tropical forests play a central role in global water, carbon, and nutrient cycling, we know very little about tradeoffs and synergies in root traits that respond to resource scarcity. Tropical trees face a unique set of resource limitations, with rock-derived nutrients and moisture seasonality governing many ecosystem functions, and nutrient versus water availability often separated spatially and temporally. Root traits that characterize biomass, depth distributions, production and phenology, morphology, physiology, chemistry, and symbiotic relationships can be predictive of plants’ capacities to access and acquire nutrients and water, with links to aboveground processes like transpiration, wood productivity, and leaf phenology. In this review, we identify an emerging trend in the literature that tropical fine root biomass and production in surface soils are greatest in infertile or sufficiently moist soils. We also identify interesting paradoxes in tropical forest root responses to changing resources that merit further exploration. For example, specific root length, which typically increases under resource scarcity to expand the volume of soil explored, instead can increase with greater base cation availability, both across natural tropical forest gradients and in fertilization experiments. Also, nutrient additions, rather than reducing mycorrhizal colonization of fine roots as might be expected, increased colonization rates under scenarios of water scarcity in some forests. Efforts to include fine root traits and functions in vegetation models have grown more sophisticated over time, yet there is a disconnect between the emphasis in models characterizing nutrient and water uptake rates and carbon costs versus the emphasis in field experiments on measuring root biomass, production, and morphology in response to changes in resource availability. Closer integration of field and modeling efforts could connect mechanistic investigation of fine-root dynamics to ecosystem-scale understanding of nutrient and water cycling, allowing us to better predict tropical forest-climate feedbacks.

  • Glacier forelands reveal fundamental plant and microbial controls on short-term ecosystem nitrogen retention

    de Vries F, Thion C, Bahn M, Pinto BB, Cecillon S, Frey B, Grant H, Nicol G, Wanek W, Prosser J, Bardgett R
    2021 - Journal of Ecology, 109: 3710-3723



    1. Human activities have massively increased the amount of reactive nitrogen in the biosphere, which is leading to increased nitrogen (N) inputs in terrestrial ecosystems. The retention of N is a crucial ecosystem function of both managed and natural ecosystems, and there is a long history of experimental, observational, and conceptual studies identifying its major controls. Yet, the plant and soil microbial controls on the retention of added N remain elusive.
    2. Here, we used three ecosystem chronosequences in front of retreating glaciers in the European Alps to test our hypothesis that the retention of added reactive 15N increases as succession proceeds, and to identify the plant and microbial controls on ecosystem N retention.
    3. We found that the uptake and retention of N did not change during succession, despite consistent changes in plant, soil, and microbial properties with increasing time since deglaciation. Instead, we found that plant and microbial properties that remained constant during succession controlled 15N uptake and retention: low root and microbial C/N ratios, as well as high root biomass, increased plant and microbial uptake of added N. In addition, high soil concentrations of nitrate and ammonium reduced the uptake of N in microbes and roots, respectively.
    4. Synthesis. Our results demonstrate that plant and microbial N demand, as well as soil N availability, drive the short-term retention of added N during succession in glacier forelands. This finding represents an advance in our understanding of the fundamental controls on ecosystem N retention and the role of plant-microbial interactions in this process. Such understanding is crucial for predicting and mitigating the response of terrestrial ecosystems to the ever-increasing amounts of reactive N in the biosphere.


    1. Human activities have massively increased the amount of reactive nitrogen in the biosphere, which is leading to increased nitrogen (N) inputs in terrestrial ecosystems. The retention of N is a crucial ecosystem function of both managed and natural ecosystems, and there is a long history of experimental, observational, and conceptual studies identifying its major controls. Yet, the plant and soil microbial controls on the retention of added N remain elusive.
    2. Here, we used three ecosystem chronosequences in front of retreating glaciers in the European Alps to test our hypothesis that the retention of added reactive 15N increases as succession proceeds, and to identify the plant and microbial controls on ecosystem N retention.
    3. We found that the uptake and retention of N did not change during succession, despite consistent changes in plant, soil, and microbial properties with increasing time since deglaciation. Instead, we found that plant and microbial properties that remained constant during succession controlled 15N uptake and retention: low root and microbial C/N ratios, as well as high root biomass, increased plant and microbial uptake of added N. In addition, high soil concentrations of nitrate and ammonium reduced the uptake of N in microbes and roots, respectively.
    4. Synthesis. Our results demonstrate that plant and microbial N demand, as well as soil N availability, drive the short-term retention of added N during succession in glacier forelands. This finding represents an advance in our understanding of the fundamental controls on ecosystem N retention and the role of plant-microbial interactions in this process. Such understanding is crucial for predicting and mitigating the response of terrestrial ecosystems to the ever-increasing amounts of reactive N in the biosphere.
  • Denitrification is the major nitrous acid production pathway in boreal agricultural soils

    Bhattarai HR, Wanek W, Siljanen H, Ronkainen J, Liimatainen M, Hu Y, Nykänen H, Biasi C, Maljanen M
    2021 - Communications Earth and Environment, 2: 54


    Nitrous acid (HONO) photolysis produces hydroxyl radicals—a key atmospheric oxidant. Soils are strong HONO emitters, yet HONO production pathways in soils and their relative contributions are poorly constrained. Here, we conduct 15N tracer experiments and isotope pool dilution assays on two types of agricultural soils in Finland to determine HONO emission fluxes and pathways. We show that microbial processes are more important than abiotic processes for HONO emissions. Microbial nitrate reduction (denitrification) considerably exceeded ammonium oxidation as a source of nitrite—a central nitrogen pool connected with HONO emissions. Denitrification contributed 97% and 62% of total HONO fluxes in low and high organic matter soil, respectively. Microbial ammonium oxidation only produced HONO in high organic matter soil (10%). Our findings indicate that microbial nitrate reduction is an important HONO production pathway in aerobic soils, suggesting that terrestrial ecosystems favouring it could be HONO emission hotspots, thereby influencing atmospheric chemistry.

  • Responses of grassland soil CO2 production and fluxes to drought are shifted in a warmer climate under elevated CO2

    Reinthaler D, Harris E, Richter A, Herndl M, Pötsch E, Wachter H, Bahn M
    2021 - Soil Biology and Biochemistry, 163: Article 108436


    As the climate warms, drought events are expected to increase in intensity and frequency, with consequences for the carbon cycleSoil respiration (Rs) accounts for the largest flux of CO2 from terrestrial ecosystems to the atmosphere. While the drought responses of Rs have been well studied, it is uncertain how they will be modified in a future world, when higher temperatures will occur in combination with elevated atmospheric CO2 concentrations. In a global change experiment in a managed temperate grassland, we studied drought and post-drought responses of Rs dynamics under current versus likely future conditions (+3°, +300 ppm CO2). Furthermore, to understand the soil CO2 production (Ps) and transport dynamics underlying Rs fluxes we continuously monitored in-situ soil CO2 concentrations across the soil profile. Our results show that Rs was higher and that drought-induced reductions in Rs were delayed under future compared to current conditions. Peak drought reductions and post-drought pulses of Rs were more pronounced in the future scenario. Annual Rs was reduced by drought only under current but not under future conditions. An in-depth analysis of soil CO2 gradients and fluxes across the soil profile showed that elevated CO2 stimulated Ps primarily in the main rooting horizon and that warming affected Ps also in deeper soil layers. We found that both in the current and the future scenario drought led to the strongest reductions of Ps in the most productive soil layers, which also exhibited the largest depletion of soil moisture. We conclude that a future warmer climate under elevated CO2 amplifies soil CO2 production and efflux and their peak drought and post-drought responses, but delays the onset of the drought responses and thereby eliminates the overall drought effect on annual soil CO2 emissions.

  • Retaining eucalyptus harvest residues promotes different pathways for particulate and mineral-associated organic matter

    Ferreira GWD, Oliveira FCC, Soares EMB, Schnecker J, Silva IR, Grandy AS
    2021 - Ecosphere, 7: Article e03439


    Eucalyptus plantations have replaced other (agro)ecosystems over 5.6 Mha in Brazil. While these plantations rapidly accumulate carbon (C) in their biomass, the C storage in living forest biomass is transient, and thus, longer‐term sustainability relies on sustaining soil organic matter (SOM) stocks. A significant amount of harvest residues (HR) is generated every rotation and can yield SOM if retained in the field. Yet, there is little information on how managing eucalyptus HR changes SOM dynamics. We used isotopic and molecular approaches in a 3‐yr field decomposition experiment where a native grassland has been replaced by eucalyptus plantations to assess how HR management practices influence content and chemistry of two distinct SOM fractions [particulate (POM) and mineral‐associated organic matter (MAOM)] at two soil depths (0–1 and 1–5 cm). The management practices investigated were HR removal (−R), only bark removal (−B), and retention of all HR (including bark, +B), combined with two levels of nitrogen (N) fertilization [0 (−N) and 200 (+N) kg/ha]. N fertilization inhibited HR decomposition (P = 0.0409), while bark retention had little effect (P = 0.1164). Retaining HR, especially with bark, increased POM‐C and MAOM‐C content (2.1‐ and 1.2‐fold, respectively), decreased POM‐δ13C (1.2‐fold), and increased inorganic N retention (1.7‐fold) compared with plots where HR had been removed. Inorganic N applications, however, diminished the positive impacts of bark retention. Although the influence of HR management was most pronounced in POM, retaining HR reduced potential soil C mineralization by up to 20%. POM and MAOM chemistry shifted over time and revealed distinct influence of HR on the formation of these fractions. We demonstrate that HR management alters SOM dynamics and that retaining HR, particularly including bark, enhances SOM retention. With continuing conversion of native grassland ecosystems to eucalyptus, long‐term sustainability will require careful HR and fertilizer management to balance total biomass harvest with sustaining belowground SOM concentrations.

  • Permafrost causes unique fine-scale spatial variability across tundra soils

    Siewert MB., Lantuit H, Richter A, Hugelius G
    2021 - Global Biogeochemical Cycles, 35: e2020GB006659


    Spatial analysis in earth sciences is often based on the concept of spatial autocorrelation, expressed by W. Tobler as the first law of geography: “everything is related to everything else, but near things are more related than distant things." Here, we show that subsurface soil properties in permafrost tundra terrain exhibit tremendous spatial variability. We describe the subsurface variability of soil organic carbon (SOC) and ground ice content from the centimeter to the landscape scale in three typical tundra terrain types common across the Arctic region. At the soil pedon scale, that is, from centimeters to 1–2 m, variability is caused by cryoturbation and affected by tussocks, hummocks and nonsorted circles. At the terrain scale, from meters to tens of meters, variability is caused by different generations of ice‐wedges. Variability at the landscape scale, that is, ranging hundreds of meters, is associated with geomorphic disturbances and catenary shifts. The co‐occurrence and overlap of different processes and landforms creates a spatial structure unique to permafrost environments. The coefficient of variation of SOC at the pedon scale (21%–73%) exceeds that found at terrain (17%–66%) and even landscape scale (24%–67%). Such high values for spatial variation are otherwise found at regional to continental scale. Clearly, permafrost soils do not conform to Tobler's law, but are among the most variable soils on Earth. This needs to be accounted for in mapping and predictions of the permafrost carbon feedbacks through various ecosystem processes. We conclude that scale deserves special attention in permafrost regions.

  • Microbial activity responses to water stress in agricultural soils from simple and complex crop rotations

    Schnecker J, Meeden DB, Calderon F, Cavigelli M, Lehman RM, Tiemann LK, Grandy AS
    2021 - Soil, 547-561


    Increasing climatic pressures such as drought and flooding challenge agricultural systems and their management globally. How agricultural soils respond to soil water extremes will influence biogeochemical cycles of carbon and nitrogen in these systems. We investigated the response of soils from long-term agricultural field sites under varying crop rotational complexity to either drought or flooding stress. Focusing on these contrasting stressors separately, we investigated soil heterotrophic respiration during single and repeated stress cycles in soils from four different sites along a precipitation gradient (Colorado, MAP 421 mm; South Dakota, MAP 580 mm; Michigan, MAP 893 mm; Maryland, MAP 1192 mm); each site had two crop rotational complexity treatments. At the driest (Colorado) and wettest (Maryland) of these sites, we also analyzed microbial biomass, six potential enzyme activities, and N2O production during and after individual and repeated stress cycles. In general, we found site specific responses to soil water extremes, irrespective of crop rotational complexity and precipitation history. Drought usually caused more severe changes in respiration rates and potential enzyme activities than flooding. All soils returned to control levels for most measured parameters as soon as soils returned to control water levels following drought or flood stress, suggesting that the investigated soils were highly resilient to the applied stresses. The lack of sustained responses following the removal of the stressors may be because they are well in the range of natural in situ soil water fluctuations at the investigated sites. Without the inclusion of plants in our experiment, we found that irrespective of crop rotation complexity, soil and microbial properties in the investigated agricultural soils were more resistant to flooding but highly resilient to drought and flooding during single or repeated stress pulses.

  • Phytodetrital quality (C:N ratio) and temperature changes affect C and N cycling of the intertidal mixotrophic foraminifer Haynesina germanica

    Wukovits J,  Enge A,  Bukenberger P,  Wanek W,  Watzka M, Heinz P
    2021 - Aquatic Biology, 30: 119-132


    The combination of lower diet quality and increased metabolic rates is assumed to cause cascading effects on organismic C cycling. Future changes in CO2 levels or terrestrial nutrient discharges in marine ecosystems can lead to increased phytoplankton C:N ratios relative to consumer C:N ratios, lowering the quality of the food source. In this study, we compared the single and interactive effects of diet quality and temperature on the feeding behavior and C and N intake and release of a common and abundant intertidal mixotrophic protist, the foraminifer Haynesina germanica. Two batches of artificially produced and dual isotope-labeled (13C/15N) chlorophyte detritus with different C:N ratios (5.6 and 7.1) were fed to the foraminifer at 3 different temperatures (15, 20, 25°C). We observed a strong interactive effect of temperature and diet. A very strong increase in feeding rates was observed at 20°C for the low-quality food source. Respiration rates of carbon derived from the low-quality diet (C:N ratio of 7.1) were lower than those of the high-quality diets and increased at 25°C. This indicates that a high C content of the diet might be of advantage in calcifying mixotrophs, since respired excess C could be advantageous for test calcification. Additionally, respired excess C could be a useful resource of CO2 for kleptoplast photosynthesis and functionality in the mixotrophic lifestyle of H. germanica. Further, the observed effects of diet and temperature could impact nutrient fluxes in the habitat of H. germanica, possibly leading to food-web shifts in the future.

  • Microbial responses to herbivory-induced vegetation changes in a high-Arctic peatland

    Bender KM, Svenning MM, Hu Y, Richter A, Schückel J, Liebner S, Tveit AT
    2021 - Polar Biology, 44: 899-911


    Herbivory by barnacle geese (Branta leucopsis) alters the vegetation cover and reduces ecosystem productivity in high-Arctic peatlands, limiting the carbon sink strength of these ecosystems. Here we investigate how herbivory-induced vegetation changes affect the activities of peat soil microbiota using metagenomics, metatranscriptomics and targeted metabolomics in a comparison of fenced exclosures and nearby grazed sites. Our results show that a different vegetation with a high proportion of vascular plants developed due to reduced herbivory, resulting in a larger and more diverse input of polysaccharides to the soil at exclosed study sites. This coincided with higher sugar and amino acid concentrations in the soil at this site as well as the establishment of a more abundant and active microbiota, including saprotrophic fungi with broad substrate ranges, like Helotiales (Ascomycota) and Agaricales (Basidiomycota). A detailed description of fungal transcriptional profiles revealed higher gene expression for cellulose, hemicellulose, pectin, lignin and chitin degradation at herbivory-exclosed sites. Furthermore, we observed an increase in the number of genes and transcripts for predatory eukaryotes such as Entomobryomorpha (Arthropoda). We conclude that in the absence of herbivory, the development of a vascular vegetation alters the soil polysaccharide composition and supports larger and more active populations of fungi and predatory eukaryotes.

  • Assimilation of Particular Organic Matter and Dissolved Organic or Inorganic Compounds by Cribroelphidium selseyense (Foraminifera)

    Lintner M, Lintner B,  Wanek W, Keul N,  Heinz P, Schmidt S
    2021 - Frontiers in Marine Science, 8: Article 778148


    Marine carbon and nitrogen processing through microorganisms’ metabolism is an important aspect of the global element cycles. For that purpose, we used foraminifera to analyze the element turnover with different algae food sources. In the Baltic Sea, benthic foraminifera are quite common and therefore it is important to understand their metabolism. Especially, Cribroelphidium selseyense, also occurring in the Baltic Sea, has often been used for laboratory feeding experiments to test their effect on carbon or nitrogen turnover. Therefore, foraminifera were collected from the Kiel Fjord and fed with six different algal species in two qualities (freeze-dried algae vs. fresh algae, all 13C- and 15N-labeled). Also, labeled dissolved inorganic C and N compounds and glucose were offered to the foraminifera to test direct assimilation of dissolved compounds (carbon and nitrogen) from the water column. Our experiments showed that after 15 days of incubation, there were highly significant differences in isotope labeling in foraminifera fed with fresh algae and dry algae, depending on algal species. Further, different algal species led to different 13C and 15N enrichment in the studied foraminifera, highlighting a feeding preference for one diatom species and an Eustigmatophyte. A significant carbon assimilation from HCO3 was observed after 7 days of incubation. The N assimilation from NH4+ was significantly higher than for NO3 as an inorganic N source. The uptake of glucose showed a lag phase, which was often observed during past experiments, where foraminifera were in a steady state and showed no food uptake at regular intervals. These results highlight the importance of food quality on the feeding behavior and metabolic pathways for further studies of foraminiferal nutrition and nutrient cycling.

  • Recovery of aboveground biomass, species richness and composition in tropical secondary forests in SW Costa Rica

    Oberleitner F, Egger C, Oberdorfer S, Dullinger S, Wanek W, Hietz P
    2021 - Forest Ecology and Management, 479: Article 118580


    Tropical secondary forests comprise about half of the world’s tropical forests and are important as carbon sinks and to conserve biodiversity. Their rate of recovery varies widely; however, particularly older secondary forests are difficult to date so that the recovery rate is uncertain. As a consequence, factors affecting recovery are difficult to analyse. We used aerial surveys going back to 1968 to date 12 secondary forests in the wet tropics of SW Costa Rica and evaluated the recovery of aboveground biomass, tree species richness and tree species composition in relation to nearby old-growth forests and previous land use. To confirm the validity of the space-for-time substitution, the plots were re-censused after four years. We found fast rates of aboveground biomass accumulation, especially in the first years of succession. After 20 years AGB had reached c. 164 Mg/ha equivalent to 52% of the biomass in old-growth forests in the region. Species richness increased at a slower pace and had reached c. 31% of old-growth forests after 20 years. Recovery rates differed substantially among forests, with biomass at least initially recovering faster in forests after clearcuts whereas species numbers increased faster in forests recovering from pastures. Biomass recovery was positively related to the forest cover in the vicinity and negatively to species richness, whereas species richness was related to soil parameters. The change during the four years between the censuses is broadly in line with the initial chronosequence. While the recovery of tropical secondary forests has been studied in many places, our study shows that various environmental parameters affect the speed of recovery, which is important to include in efforts to manage and restore tropical landscapes.

  • Nitrogen Kinetic Isotope Effects of Nitrification by the Complete Ammonia Oxidizer Nitrospira inopinata

    Liua S, Jung MY, zhang S, Wagner M, Daims H, Wanek W
    2021 - mSphere, 6: Article e00634-21


    Analysis of nitrogen isotope fractionation effects is useful for tracing biogeochemical nitrogen cycle processes. Nitrification can cause large nitrogen isotope effects through the enzymatic oxidation of ammonia (NH3) via nitrite (NO2) to nitrate (NO3) (15εNH4+→NO2- and 15εNO2-→NO3-). The isotope effects of ammonia-oxidizing bacteria (AOB) and archaea (AOA) and of nitrite-oxidizing bacteria (NOB) have been analyzed previously. Here, we studied the nitrogen isotope effects of the complete ammonia oxidizer (comammox) Nitrospira inopinata that oxidizes NH3 to NO3. At high ammonium (NH4+) availability (1 mM) and pH between 6.5 and 8.5, its 15εNH4+→NO2- ranged from −33.1 to −27.1‰ based on substrate consumption (residual substrate isotopic composition) and −35.5 to −31.2‰ based on product formation (cumulative product isotopic composition), while the 15εNO2-→NO3- ranged from 6.5 to 11.1‰ based on substrate consumption. These values resemble isotope effects of AOB and AOA and of NOB in the genus Nitrospira, suggesting the absence of fundamental mechanistic differences between key enzymes for ammonia and nitrite oxidation in comammox and canonical nitrifiers. However, ambient pH and initial NH4+ concentrations influenced the isotope effects in N. inopinata. The 15εNH4+→NO2- based on product formation was smaller at pH 6.5 (−31.2‰) compared to pH 7.5 (−35.5‰) and pH 8.5 (−34.9‰), while 15εNO2-→NO3- was smaller at pH 8.5 (6.5‰) compared to pH 7.5 (8.8‰) and pH 6.5 (11.1‰). Isotopic fractionation via 15εNH4+→NO2- and 15εNO2-→NO3- was smaller at 0.1 mM NH4+ compared to 0.5 to 1.0 mM NH4+. Environmental factors, such as pH and NH4+ availability, therefore need to be considered when using isotope effects in 15N isotope fractionation models of nitrification.

  • A critical perspective on interpreting amplicon sequencing data in soil ecological research

    Alteio L, Séneca J, Canarini A, Angel R, Jansa J, Guseva K, Kaiser C, Richter A, Schmidt H
    2021 - Soil Biology and Biochemistry, 160: Article 108357


    Microbial community analysis via marker gene amplicon sequencing has become a routine method in the field of soil research. In this perspective, we discuss technical challenges and limitations of amplicon sequencing and present statistical and experimental approaches that can help addressing the spatio-temporal complexity of soil and the high diversity of organisms therein. We illustrate the impact of compositionality on the interpretation of relative abundance data and discuss effects of sample replication on the statistical power in soil community analysis. Additionally, we argue for the need of increased study reproducibility and data availability, as well as complementary techniques for generating deeper ecological insights into microbial roles and our understanding thereof in soil ecosystems. At this stage, we call upon researchers and specialized soil journals to consider the current state of data analysis, interpretation, and availability to improve the rigor of future studies.

  • An unexpected source of nitrogen for root uptake: positively charged amino acids dominate soil diffusive nitrogen fluxes. Commentary.

    Inselbacher E, Wanek W
    2021 - New Phytologist, 231: 2104-2106


    This article is a Commentary on Homyak et al. (2021), 231: 2162–2173.


    Soils typically contain a large variety of nitrogen (N) forms, including inorganic N and a range of organic N compounds of varying molecular size (Warren, 2013). Inorganic N was long been considered to constitute the main source of N for plants, but this view has changed considerably since plants were shown to be capable of directly taking up and metabolizing organic N forms, including amino acids, peptides, proteins and quaternary ammonium compounds (Näsholm et al., 2009; Warren, 2013). Amino acid uptake especially has been demonstrated in every plant species studied thus far and the underlying uptake mechanisms have been investigated extensively (Näsholm et al., 2009; Narcy et al., 2013). Yet even if plants have the potential to take up amino acids, those N forms first have to be bioavailable and have to be consistently replenished at root surfaces. However, reliably estimating such N availability is challenging due to the sheer complexity of soils and plant root systems.

  • Functional traits of a rainforest vascular epiphyte community: trait covariation and indications for host specificity

    Wagner K, Wanek W, Zotz G
    2021 - Diversity, 13: 97


    Trait matching between interacting species may foster diversity. Thus, high epiphyte diversity in tropical forests may be partly due to the high diversity of trees and some degree of host specificity. However, possible trait matching between epiphyte and host is basically unexplored. Since the epiphytic habitat poses particular challenges to plants, their trait correlations should differ from terrestrial plants, but to what extent is unclear as epiphytes are underrepresented or missing in the large trait databases. We quantified 28 traits of 99 species of vascular epiphytes in a lowland forest in Panama that were related to plant size, leaf, stem, and root morphology; photosynthetic mode; and nutrient concentrations. We analyzed trait covariation, community weighted means, and functional diversity for assemblages on stems and in crowns of four tree species. We found intriguing differences between epiphytes and terrestrial plants regarding trait covariation in trait relations between plant maximal height, stem specific density, specific root length, and root tissue den-sity, i.e., stem and root economic spectra. Regarding host specificity, we found strong evidence for environmental filtering of epiphyte traits, but only in tree crowns. On stems, community weighted means differed in only one case, whereas > 2/3 of all traits differed in tree crowns. Although we were only partly able to interpret these differences in the light of tree trait differences, these findings mark an important step towards a functional understanding of epiphyte host specificity.

  • Tree Species and Epiphyte Taxa Determine the “Metabolomic niche” of Canopy Suspended Soils in a Species-Rich Lowland Tropical Rainforest

    Gargallo-Garriga A, Sardans J, Alrefaei AF, Klem K, Fuchslueger L, Ramírez-Rojas I, Donald J, Leroy C, Van Langenhove L, Verbruggen E, Janssens IA, Urban O, Peñuelas J
    2021 - Metabolites, 11: Article 718


    Tropical forests are biodiversity hotspots, but it is not well understood how this diversity is structured and maintained. One hypothesis rests on the generation of a range of metabolic niches, with varied composition, supporting a high species diversity. Characterizing soil metabolomes can reveal fine-scale differences in composition and potentially help explain variation across these habitats. In particular, little is known about canopy soils, which are unique habitats that are likely to be sources of additional biodiversity and biogeochemical cycling in tropical forests. We studied the effects of diverse tree species and epiphytes on soil metabolomic profiles of forest floor and canopy suspended soils in a French Guianese rainforest. We found that the metabolomic profiles of canopy suspended soils were distinct from those of forest floor soils, differing between epiphyte-associated and non-epiphyte suspended soils, and the metabolomic profiles of suspended soils varied with host tree species, regardless of association with epiphyte. Thus, tree species is a key driver of rainforest suspended soil metabolomics. We found greater abundance of metabolites in suspended soils, particularly in groups associated with plants, such as phenolic compounds, and with metabolic pathways related to amino acids, nucleotides, and energy metabolism, due to the greater relative proportion of tree and epiphyte organic material derived from litter and root exudates, indicating a strong legacy of parent biological material. Our study provides evidence for the role of tree and epiphyte species in canopy soil metabolomic composition and in maintaining the high levels of soil metabolome diversity in this tropical rainforest. It is likely that a wide array of canopy microsite-level environmental conditions, which reflect interactions between trees and epiphytes, increase the microscale diversity in suspended soil metabolomes.

  • No effect of long-term soil warming on diffusive soil inorganic and organic nitrogen fluxes in a temperate forest soil

    Heinzle J, Wanek W, Tian Y, Kwatcho-Kengdo S, Borken W, Schindlbacher A, Inselsbacher E
    2021 - Soil Biology and Biochemistry, 158: Article 108261


    Climate warming affects nitrogen (N) cycling in forest soils, but implications for plant available N have remained unclear. We estimated in situ diffusive fluxes of amino acids and inorganic N in a temperate forest soil after 14 years of soil warming. Results from four sampling campaigns (n = 1152 microdialysis samples) during the growing season showed no effect of warming on diffusive N fluxes. Diffusive NH4+ fluxes increased from spring towards autumn while NO3 fluxes followed an opposite trend. Overall, the proportion of amino acids in the total diffusive N flux was low (13–30%) in this carbonate soil compared to other temperate and boreal forest soils.

  • Consistent shift in nutritional ecology of ants reveals trophic flexibility across alpine tree-line ecotones

    Guariento E,  Wanek W, Fiedler
    2021 - Ecological Entomology, 46: 1082-1092


    1. Studying the feeding ecology of ants can reveal their trophic position and allow inferences on interactions with other organisms. We investigated the nutritional ecology of ants at the Alpine tree line (from subalpine forests to alpine grasslands), testing the hypothesis that changing food availability reflects upon ant feeding preference and trophic position. Five slopes with five sites each were sampled using a combined experimental (baiting) and chemical (stable isotope) approach.

    2. Sugar resources were most preferred by the whole ant community in the alpine environment and on the tree line and were therefore likely limiting in these habitats. This shift was not detected in the two dominant ant taxa occurring over the investigated gradient, viz. the slave ant Formica lemani and mound-building wood ants (Formica s. str.).

    3. Yet, stable carbon (C) and nitrogen (N) isotope signatures of both these ant species likewise revealed a shift in resource use over the ecotone, from a lower trophic position in subalpine forests to a more carnivore-dominated lifestyle in alpine grassland. Moreover, wood ants were found to occupy a higher trophic position compared to F. lemani.

    4. In conclusion, energy resources turned out to be more limiting for ants in the alpine environment, but less so in subalpine forests. The low trophic position of these ant species in the forest is considered to be a result of abundant trophobiotic associations with honeydew-producing homopterans that occur in far larger numbers on conifer trees in the forest.

  • Spatiotemporal Dynamics of Maize (Zea mays L.) Root Growth and Its Potential Consequences for the Assembly of the Rhizosphere Microbiota

    Bonkowski M, Tarkka M, Razavi BS, Schmidt H, Blagodatskaya E, Koller R, Yu P, Knief C, Hochholdinger F, Vetterlein D
    2021 - Frontiers in microbiology, 12: Article 619499


    Numerous studies have shown that plants selectively recruit microbes from the soil to establish a complex, yet stable and quite predictable microbial community on their roots – their “microbiome” (Berg and Smalla, 2009Hartmann et al., 2009Weinert et al., 2010). Microbiome assembly is considered as a key process in the self-organization of root systems (Vetterlein et al., 2020). Better control of microbiome assembly would improve plant health and fitness by promoting beneficial microbial traits (Friesen et al., 2011Oyserman et al., 2018Wille et al., 2019). A fundamental question for understanding plant-microbe relationships is where a predictable microbiome is formed along the root axis and through which microbial dynamics the stable formation of a microbiome is challenged. Theoretically, community assembly begins with random, unregulated colonization of taxa from nearby sites (i.e., neutral processes), a process that continues throughout the lifetime of roots; while ordered dynamics (microbiome assembly) occur through selection (i.e., niche-based processes) when (i) exudates promote fast-growing copiotrophic taxa, (ii) root signals attract specific symbionts or pathogens, (iii) increased competition due to limited resource availability leads to species sorting, and (iv) predation selects for specific microbial traits among members of the microbiome (Vellend, 2010Hardoim et al., 2011Ho et al., 2017Kudjordjie et al., 2019Amacker et al., 2020Chen et al., 2020). These microbial assembly processes again are embedded in plant-driven spatiotemporal dynamics at small and large scales, caused by differences in the quality and quantity of rhizodeposition: (i) along the root axis, (ii) during diurnal cycles, (iii) on different root types, and (iv) during plant development. Emphasizing maize as a model species for which numerous data on dynamic root traits are available, this mini-review aims to give an integrative overview on the dynamic nature of root growth and its consequences for microbiome assembly based on theoretical considerations from microbial community ecology.

  • Effects of heavy elements (Pb, Cu, Zn) on algal food uptake by Elphidium excavatum (Foraminifera)

    Lintner M,  Biedrawa B,  Wanek W, Keul N,  von der Kammer F,  Hofmann T, Heinz P
    2021 - Heliyon, 18: Article e08427


    Foraminifera are unicellular organisms and play a pivotal role in the marine material cycles. Past observations have shown that the species Elphidium excavatum is the most common foraminifera in the Baltic Sea. Feeding experiments showed that the food uptake and thus the turnover of organic matter are influenced by changes of physical parameters (e.g., temperature, salinity). Since many areas of the Baltic Sea are strongly affected by anthropogenic activity and are strongly contaminated by heavy elements from shipping in the past, this study examined the effect of heavy elements pollution on the food uptake of the most common foraminiferal species of the Baltic Sea, E. excavatum which was a subject of several previous studies. Therefore, Baltic Sea seawater was enriched with metals at various levels above normal seawater levels and the uptake of 13C- and 15N-labelled phytodetritus was measured by isotope ratio mass spectrometry. For each combination of metal type, concentration and time point 20 individuals of E. excavatum (three replicates) were fed with the green algae Dunaliella tertiolecta. The effect of dose parameters was measured in a two-way analysis of variance. Significant differences of food uptake were observable at different types and levels of heavy elements in sea water. Even a 557-fold increase in the Pb concentration did not affect food uptake, whereas strong negative effects were found for higher levels of Zn (144 and 1044-fold) and especially for Cu (5.6 and 24.3-fold). In summary it can be stated, that an increase in the heavy elements pollution in the Kiel Fjord will lead to a significant reduction in the turnover of organic matter by foraminifera such as E. excavatum.

  • Ecological memory of recurrent drought modifies soil processes via changes in soil microbial community

    Canarini A, Schmidt H, Fuchslueger L, Martin V, Herbold CW, Zezula D, Gündler P, Hasibeder R, Jecmenica M, Bahn M, Richter A
    2021 - Nature Communications, 12: Article 5308


    Climate change is altering the frequency and severity of drought events. Recent evidence indicates that drought may produce legacy effects on soil microbial communities. However, it is unclear whether precedent drought events lead to ecological memory formation, i.e., the capacity of past events to influence current ecosystem response trajectories. Here, we utilize a long-term field experiment in a mountain grassland in central Austria with an experimental layout comparing 10 years of recurrent drought events to a single drought event and ambient conditions. We show that recurrent droughts increase the dissimilarity of microbial communities compared to control and single drought events, and enhance soil multifunctionality during drought (calculated via measurements of potential enzymatic activities, soil nutrients, microbial biomass stoichiometry and belowground net primary productivity). Our results indicate that soil microbial community composition changes in concert with its functioning, with consequences for soil processes. The formation of ecological memory in soil under recurrent drought may enhance the resilience of ecosystem functioning against future drought events.

  • Sensitivity and specificity of the antigen-based anterior nasal self-testing programme for detecting SARS-CoV-2 infection in schools, Austria, March 2021

    Willeit P, Bernar B, Zurl C, Al-Rawi M, Berghold A, Bernhard D, Borena W, Doppler C, Kerbl R, Köhler A, Krause R, Lamprecht B, Pröll J, Schmidt H, Steinmetz I, Stelzl E, Stoiber H, von Laer D, Zuber J, Müller T, Strenger V, Wagner M
    2021 - Euro Surveill., 26: Article 2100797


    This study evaluates the performance of the antigen-based anterior nasal screening programme implemented in all Austrian schools to detect SARS-CoV-2 infections. We combined nationwide antigen-based screening data obtained in March 2021 from 5,370 schools (Grade 1-8) with an RT-qPCR-based prospective cohort study comprising a representative sample of 244 schools. Considering a range of assumptions, only a subset of infected individuals are detected with the programme (low to moderate sensitivity) and non-infected individuals mainly tested negative (very high specificity).

  • How can fertilization regimes and durations shape earthworm gut microbiota in a long-term field experiment?

    Bi Q-F, Jin B-J, Zhu D, Jiang Y-G, Zheng B-X, O’Connor P, Yang X-R, Richter A, Lin X-Y, Zhu Y-H 
    2021 - Ecotoxicology and Environmental Safety, 224: Article 112643


    The positive roles of earthworms on soil functionality has been extensively documented. The capacity of the earthworm gut microbiota on decomposition and nutrient cycling under long-term fertilization in field conditions has rarely been studied. Here, we report the structural, taxonomic, and functional responses of Eisenia foetida and Pheretima guillelmi gut microbiota to different fertilization regimes and durations using 16S rRNA gene-based Illumina sequencing and high-throughput quantitative PCR techniques. Our results revealed that the core gut microbiota, especially the fermentative bacteria were mainly sourced from the soil, but strongly stimulated with species-specificity, potential benefits for the host and soil health. The functional compositions of gut microbiota were altered by fertilization with fertilization duration being more influential than fertilization regimes. Moreover, the combination of organic and inorganic fertilization with the longer duration resulted in a higher richness and connectivity in the gut microbiota, and also their functional potential related to carbon (C), nitrogen, and phosphorus cycling, particularly the labile C decomposition, denitrification, and phosphate mobilization. We also found that long-term inorganic fertilization increased the abundance of pathogenic bacteria in the P. guillelmi gut. This study demonstrates that understanding earthworm gut microbiota can provide insights into how agricultural practices can potentially alte

  • The effect of global change on soil phosphatase activity

    Margalef O, Sardans J, Maspons J, Molowny-Horas R, Fernández-Martínez M, Janssens IA, Ciais P, Richter A, Obersteiner M, Peñuelas J
    2021 - Global Change Biology, 27: 5681-6003


    Soil phosphatase enzymes are produced by plant roots and microorganisms and play a key role in the cycling of phosphorus (P), an often-limiting element in terrestrial ecosystems. The production of these enzymes in soil is the most important biological strategy for acquiring phosphate ions from organic molecules. Previous works showed how soil potential phosphatase activity is mainly driven by climatic conditions and soil nitrogen (N) and carbon. Nonetheless, future trends of the activity of these enzymes under global change remain little known. We investigated the influence of some of the main drivers of change on soil phosphatase activity using a meta-analysis of results from 97 published studies. Our database included a compilation of N and P fertilization experiments, manipulation experiments with increased atmospheric CO2 concentration, warming, and drought, and studies comparing invaded and non-invaded ecosystems. Our results indicate that N fertilization leads to higher phosphatase activity, whereas P fertilization has the opposite effect. The rise of atmospheric CO2 levels or the arrival of invasive species also exhibits positive response ratios on the activity of soil phosphatases. However, the occurrence of recurrent drought episodes decreases the activity of soil phosphatases. Our analysis did not reveal statistically significant effects of warming on soil phosphatase activity. In general, soil enzymatic changes in the reviewed experiments depended on the initial nutrient and water status of the ecosystems. The observed patterns evidence that future soil phosphatase activity will not only depend on present-day soil conditions but also on potential compensations or amplifications among the different drivers of global change. The responses of soil phosphatases to the global change drivers reported in this study and the consideration of cost–benefit approaches based on the connection of the P and N cycle will be useful for a better estimation of phosphatase production in carbon (C)–N–P models.

  • Shifts in the abundances of saprotrophic and ectomycorrhizal fungi at altered leaf litter inputs

    Marañon-Jimenez S, Radujkovic D, Verbruggen E, Grau O, Cuntz M, Peñuelas J, Richter A, Schrumpf M, Rebmann C
    2021 - Frontiers in Plant Science, 12: Article 682142


    Ectomycorrhizal (EcM) and saprotrophic fungi interact in the breakdown of organic matter, but the mechanisms underlying the EcM role on organic matter decomposition are not totally clear. We hypothesized that the ecological relations between EcM and saprotroph fungi are modulated by resources availability and accessibility, determining decomposition rates. We manipulated the amount of leaf litter inputs (No-Litter, Control Litter, Doubled Litter) on Trenched (root exclusion) and Non-Trenched plots (with roots) in a temperate deciduous forest of EcM-associated trees. Resultant shifts in soil fungal communities were determined by phospholipid fatty acids and DNA sequencing after 3 years, and CO2 fluxes were measured throughout this period. Different levels of leaf litter inputs generated a gradient of organic substrate availability and accessibility, altering the composition and ecological relations between EcM and saprotroph fungal communities. EcM fungi dominated at low levels of fresh organic substrates and lower organic matter quality, where short-distances exploration types seem to be better competitors, whereas saprotrophs and longer exploration types of EcM fungi tended to dominate at high levels of leaf litter inputs, where labile organic substrates were easily accessible. We were, however, not able to detect unequivocal signs of competition between these fungal groups for common resources. These results point to the relevance of substrate quality and availability as key factors determining the role of EcM and saprotroph fungi on litter and soil organic matter decay and represent a path forward on the capacity of organic matter decomposition of different exploration types of EcM fungi.

  • Recently photoassimilated carbon and fungus-deliverd nitrogen are spatially correlated in the ectomycorrhizal tissue of Fagus sylvatica

    Mayerhofer W, Schintelmeister A, Dietrich M, Gorka S, Wiesenbauer J, Martin V, Gabriel R, Reipert S, Weidinger M, Clode P, Wagner M, Wöbken D, Richter A, Kaiser C
    2021 - New Phytologist, 232: 2457-2474


    Ectomycorrhizal plants trade plant-assimilated carbon for soil nutrients with their fungal partners. The underlying mechanisms, however, are not fully understood. Here we investigate the exchange of carbon for nitrogen in the ectomycorrhizal symbiosis of Fagus sylvatica across different spatial scales from the root system to the cellular level. We provided 15 N-labelled nitrogen to mycorrhizal hyphae associated with one half of the root system of young beech trees, while exposing plants to a 13 CO2 atmosphere. We analysed the short-term distribution of 13 C and 15 N in the root system with isotope-ratio mass spectrometry, and at the cellular scale within a mycorrhizal root tip with nanoscale secondary ion mass spectrometry (NanoSIMS). At the root system scale, plants did not allocate more 13 C to root parts that received more 15 N. Nanoscale secondary ion mass spectrometry imaging, however, revealed a highly heterogenous, and spatially significantly correlated distribution of 13 C and 15 N at the cellular scale. Our results indicate that, on a coarse scale, plants do not allocate a larger proportion of photoassimilated C to root parts associated with N-delivering ectomycorrhizal fungi. Within the ectomycorrhizal tissue, however, recently plant-assimilated C and fungus-delivered N were spatially strongly coupled. Here, NanoSIMS visualisation provides an initial insight into the regulation of ectomycorrhizal C and N exchange at the microscale.

  • Warming and elevated CO2 intensify drought and recovery responses of grassland carbon allocation to soil respiration

    Meeran K, Ingrisch J, Reinthaler D, Canarini A, Müller L, Pötsch E, Richter A, Wanek W, Bahn M
    2021 - Global Change Biology, 27: 3230-3243


    Photosynthesis and soil respiration represent the two largest fluxes of CO2 in terrestrial ecosystems and are tightly linked through belowground carbon (C) allocation. Drought has been suggested to impact the allocation of recently assimilated C to soil respiration; however, it is largely unknown how drought effects are altered by a future warmer climate under elevated atmospheric CO2 (eT_eCO2). In a multifactor experiment on managed C3 grassland, we studied the individual and interactive effects of drought and eT_eCO2 (drought, eT_eCO2, drought × eT_eCO2) on ecosystem C dynamics. We performed two in situ 13CO2 pulse-labeling campaigns to trace the fate of recent C during peak drought and recovery. eT_eCO2 increased soil respiration and the fraction of recently assimilated C in soil respiration. During drought, plant C uptake was reduced by c. 50% in both ambient and eT_eCO2 conditions. Soil respiration and the amount and proportion of 13C respired from soil were reduced (by 32%, 70% and 30%, respectively), the effect being more pronounced under eT_eCO2 (50%, 84%, 70%). Under drought, the diel coupling of photosynthesis and SR persisted only in the eT_eCO2 scenario, likely caused by dynamic shifts in the use of freshly assimilated C between storage and respiration. Drought did not affect the fraction of recent C remaining in plant biomass under ambient and eT_eCO2, but reduced the small fraction remaining in soil under eT_eCO2. After rewetting, C uptake and the proportion of recent C in soil respiration recovered more rapidly under eT_eCO2 compared to ambient conditions. Overall, our findings suggest that in a warmer climate under elevated CO2 drought effects on the fate of recent C will be amplified and the coupling of photosynthesis and soil respiration will be sustained. To predict the future dynamics of terrestrial C cycling, such interactive effects of multiple global change factors should be considered.

  • Mosses reduce soil nitrogen availability in subarctic birch forest via effects on soil thermal regime and sequestration of deposited nitrogen

    Koranda M, Michelsen A
    2021 - Journal of Ecology, 109: 1424-1438


    In high-latitude ecosystems bryophytes are important drivers of ecosystem functions. Alterations in abundance of mosses due to global change may thus strongly influence carbon (C) and nitrogen (N) cycling and hence cause feedback on climate. The effects of mosses on soil microbial activity are, however, still poorly understood. Our study aims at elucidating how and by which mechanisms bryophytes influence microbial decomposition processes of soil organic matter and thus soil nutrient availability.We present results from a field experiment in a subarctic birch forest in northern Sweden, where we partly removed the moss cover and replaced it with an artificial soil cover for simulating moss effects on soil temperature and moisture. We combined this with a fertilization experiment with 15N-labelled N for analysing the effects of moss N sequestration on soil processes.Our results demonstrate the capacity of mosses to reduce soil N availability and retard N cycling. The comparison with artificial soil cover plots suggests that the effect of mosses on N cycling is linked to the thermal insulation capacity of mosses causing low average soil temperature in summer and strongly reduced soil temperature fluctuations, the latter also leading to a decreased frequency of freeze-thaw events in autumn and spring. Our results also showed, however, that the negative temperature effect of mosses on soil microbial activity was in part compensated by stimulatory effects of the moss layer, possibly linked to leaching of labile substrates from the moss. Furthermore, our results revealed that bryophytes efficiently sequester added N from wet deposition and thus prevent effects of increased atmospheric N deposition on soil N availability and soil processes. Synthesis. Our study emphasizes the important role of mosses in carbon and nutrient cycling in high-latitude ecosystems and the potential strong impacts of reductions in moss abundance on microbial decomposition processes and nutrient availability in subarctic and boreal forests.

  • Ethanol production from wheat stray hydrolysate by Issatchenkia orientalis isolated from waste cooking oil

    Zwirzitz A, Alteio L, Sulzenbacher D, Atanasoff M, Selg M
    2021 - J Fungi, 7: Article 121


    The interest in using non-conventional yeasts to produce value-added compounds from low cost substrates, such as lignocellulosic materials, has increased in recent years. Setting out to discover novel microbial strains that can be used in biorefineries, an Issatchenkia orientalis strain was isolated from waste cooking oil (WCO) and its capability to produce ethanol from wheat straw hydrolysate (WSHL) was analyzed. As with previously isolated I. orientalis strains, WCO-isolated I. orientalis KJ27-7 is thermotolerant. It grows well at elevated temperatures up to 42 °C. Furthermore, spot drop tests showed that it is tolerant to various chemical fermentation inhibitors that are derived from the pre-treatment of lignocellulosic materials. I. orientalis KJ27-7 is particularly tolerant to acetic acid (up to 75 mM) and tolerates 10 mM formic acid, 5 mM furfural and 10 mM hydroxymethylfurfural. Important for biotechnological cellulosic ethanol production, I. orientalis KJ27-7 grows well on plates containing up to 10% ethanol and media containing up to 90% WSHL. As observed in shake flask fermentations, the specific ethanol productivity correlates with WSHL concentrations. In 90% WSHL media, I. orientalis KJ27-7 produced 10.3 g L-1 ethanol within 24 h. This corresponds to a product yield of 0.50 g g-1 glucose (97% of the theoretical maximum) and a volumetric productivity of 0.43 g L-1 h-1. Therefore, I. orientalis KJ27-7 is an efficient producer of lignocellulosic ethanol from WSHL.

  • Empirical support for the biogeochemical niche hypothesis in forest trees

    Sardans J, Vallicrosa H, Zuccarini P, Farré-Armengol G, Fernández-Martínez M, Guille P, Gargallo-Garriga A, Ciais P, Janssens IA, Obersteiner M, Richter A, Peñuelas J
    2021 - Nature Ecology & Evolution, 5: 184-194


    The possibility of using the elemental compositions of species as a tool to identify species/genotype niche remains to be tested at a global scale. We investigated relationships between the foliar elemental compositions (elementomes) of trees at a global scale with phylogeny, climate, N deposition and soil traits. We analysed foliar N, P, K, Ca, Mg and S concentrations in 23,962 trees of 227 species. Shared ancestry explained 60–94% of the total variance in foliar nutrient concentrations and ratios whereas current climate, atmospheric N deposition and soil type together explained 1–7%, consistent with the biogeochemical niche hypothesis which predicts that each species will have a specific need for and use of each bio-element. The remaining variance was explained by the avoidance of nutritional competition with other species and natural variability within species. The biogeochemical niche hypothesis is thus able to quantify species-specific tree niches and their shifts in response to environmental changes.

  • Rapid Responses of root traits and productivity to phosphorus and cation additions in a tropical lowland forest in Amazonia

    Lugli LF, Rosa JS, Andersen KM, Di Ponzio R, Almeida RV, Pires M, Cordeiro AL, Cunha HFV, Martins NP, Assis RL, Moraes ACM, Souza ST, Arag˜ao LEOC, Camargo JL, Fuchslueger L, Schaap KJ, Valverde-Barrantes OJ, Meir P, Quesada CA, Mercado LM, Hartley IP
    2021 - New Phytologist, 230: 116-128


    Soil nutrient availability can strongly affect root traits. In tropical forests, phosphorus (P) is often considered the main limiting nutrient for plants. However, support for the P paradigm is limited, and N and cations might also control tropical forests functioning. We used a large-scale experiment to determine how the factorial addition of nitrogen (N), P and cations affected root productivity and traits related to nutrient acquisition strategies (morphological traits, phosphatase activity, arbuscular mycorrhizal colonisation and nutrient contents) in a primary rainforest growing on low-fertility soils in Central Amazonia after 1 yr of fertilisation. Multiple root traits and productivity were affected. Phosphorus additions increased annual root productivity and root diameter, but decreased root phosphatase activity. Cation additions increased root productivity at certain times of year, also increasing root diameter and mycorrhizal colonisation. P and cation additions increased their element concentrations in root tissues. No responses were detected with N addition. Here we showed that rock-derived nutrients determined root functioning in low-fertility Amazonian soils, demonstrating not only the hypothesised importance of P, but also highlighting the role of cations. The changes in fine root traits and productivity indicated that even slow-growing tropical rainforests can respond rapidly to changes in resource availability.

  • Comparable canopy and soil free-living nitrogen fixation rates in a lowland tropical forest

    Van Langenhove L, Depaepe T, Verryckt LT, Fuchslueger L, Leroy JDC, Moorthy SMK, Gargallo-Garriga A, Ellwood MDF, Verbeeck H, Van Der Straeten D, Peñuelas J, Janssens IA
    2021 - Science of The Total Environment, 754: Article 142202


    Biological nitrogen fixation (BNF) is a fundamental part of nitrogen cycling in tropical forests, yet little is known about the contribution made by free-living nitrogen fixers inhabiting the often-extensive forest canopy. We used the acetylene reduction assay, calibrated with 15N2, to measure free-living BNF on forest canopy leaves, vascular epiphytes, bryophytes and canopy soil, as well as on the forest floor in leaf litter and soil. We used a combination of calculated and published component densities to upscale free-living BNF rates to the forest level. We found that bryophytes and leaves situated in the canopy in particular displayed high mass-based rates of free-living BNF. Additionally, we calculated that nearly 2 kg of nitrogen enters the forest ecosystem through free-living BNF every year, 40% of which was fixed by the various canopy components. Our results reveal that in the studied tropical lowland forest a large part of the nitrogen input through free-living BNF stems from the canopy, but also that the total nitrogen inputs by free-living BNF are lower than previously thought and comparable to the inputs of reactive nitrogen by atmospheric deposition.

  • Acidobacteria are active and abundant members of diverse atmospheric H2-oxidizing communities detected in temperate soils

    Eichorst S, Giguere A, Meier D, Herbold C, Richter A, Greening C, Woebken D
    2021 - ISME Journal, 15: 363-376


    Significant rates of atmospheric dihydrogen (H2) consumption have been observed in temperate soils due to the activity of high-affinity enzymes, such as the group 1h [NiFe]-hydrogenase. We designed broadly inclusive primers targeting the large subunit gene (hhyL) of group 1h [NiFe]-hydrogenases for long-read sequencing to explore its taxonomic distribution across soils. This approach revealed a diverse collection of microorganisms harboring hhyL, including previously unknown groups and taxonomically not assignable sequences. Acidobacterial group 1h [NiFe]-hydrogenase genes were abundant and expressed in temperate soils. To support the participation of acidobacteria in H2 consumption, we studied two representative mesophilic soil acidobacteria, which expressed group 1h [NiFe]-hydrogenases and consumed atmospheric H2 during carbon starvation. This is the first time mesophilic acidobacteria, which are abundant in ubiquitous temperate soils, have been shown to oxidize H2 down to below atmospheric concentrations. As this physiology allows bacteria to survive periods of carbon starvation, it could explain the success of soil acidobacteria. With our long-read sequencing approach of group 1h [NiFe]-hydrogenase genes, we show that the ability to oxidize atmospheric levels of H2 is more widely distributed among soil bacteria than previously recognized and could represent a common mechanism enabling bacteria to persist during periods of carbon deprivation.

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