Publications in peer reviewed journals

24 Publications found
  • Storage, landscape distribution, and burial history of soil organic matter in contrasting areas of continuous permafrost

    Palmtag J, Hugelius G, Lashchinskiy N, Tamstorf MP, Richter A, Elberling B, Kuhry P
    2015 - Arctic Antarctic and Alpine Research, 47: 71-88


    This study describes and compares soil organic matter (SOM) quantity and characteristics in two areas of continuous permafrost, a mountainous region in NE Greenland (Zackenberg study site) and a lowland region in NE Siberia (Cherskiy and Shalaurovo study sites). Our assessments are based on stratified-random landscape-level inventories of soil profiles down to 1 m depth, with physico-chemical, elemental, and radiocarbon-dating analyses. The estimated mean soil organic carbon (SOC) storage in the upper meter of soils in the NE Greenland site is 8.3 +/- 1.8 kg C m(-2) compared to 20.3 +/- 2.2 kg C m(-2) and 30.0 +/- 2.0 kg C m(-2) in the NE Siberian sites (95% confidence intervals). The lower SOC storage in the High Arctic site in NE Greenland can be largely explained by the fact that 59% of the study area is located at higher elevation with mostly barren ground and thus very low SOC contents. In addition, SOC-rich fens and bogs occupy a much smaller proportion of the landscape in NE Greenland (similar to 3%) than in NE Siberia (similar to 20%). The contribution of deeper buried C-enriched material in the mineral soil horizons to the total SOC storage is lower in the NE Greenland site (similar to 13%) compared to the NE Siberian sites (similar to 24%-30%). Buried SOM seems generally more decomposed in NE Greenland than in NE Siberia, which we relate to different burial mechanisms prevailing in these regions.

  • Social dynamics within decomposer communities lead to nitrogen retention and organic matter build-up in soils

    Kaiser C, Franklin O, Richter A, Dieckmann U
    2015 - Nature Communication, 6: 8960


    The chemical structure of organic matter has been shown to be only marginally important for its decomposability by microorganisms. The question of why organic matter does accumulate in the face of powerful microbial degraders is thus key for understanding terrestrial carbon and nitrogen cycling. Here we demonstrate, based on an individual-based microbial community model, that social dynamics among microbes producing extracellular enzymes (‘decomposers’) and microbes exploiting the catalytic activities of others (‘cheaters’) regulate organic matter turnover. We show that the presence of cheaters increases nitrogen retention and organic matter build-up by downregulating the ratio of extracellular enzymes to total microbial biomass, allowing nitrogen-rich microbial necromass to accumulate. Moreover, increasing catalytic efficiencies of enzymes are outbalanced by a strong negative feedback on enzyme producers, leading to less enzymes being produced at the community level. Our results thus reveal a possible control mechanism that may buffer soil CO2 emissions in a future climate.

  • Microbial nitrogen dynamics in organic and mineral soil horizons along a latitudinal transect in western Siberia

    Wild B, Schnecker J, Knoltsch A, Takriti M, Mooshammer M, Gentsch N, Mikutta R, Eloy Alves RJ, Gittel A, Lashchinskiy N, Richter A
    2015 - Global Biogeochemical Cycles, 29: 567-582


    Soil N availability is constrained by the breakdown of N-containing polymers such as proteins to oligopeptides and amino acids that can be taken up by plants and microorganisms. Excess N is released from microbial cells as ammonium (N mineralization), which in turn can serve as substrate for nitrification. According to stoichiometric theory, N mineralization and nitrification are expected to increase in relation to protein depolymerization with decreasing N limitation, and thus from higher to lower latitudes and from topsoils to subsoils. To test these hypotheses, we compared gross rates of protein depolymerization, N mineralization and nitrification (determined using N-15 pool dilution assays) in organic topsoil, mineral topsoil, and mineral subsoil of seven ecosystems along a latitudinal transect in western Siberia, from tundra (67 degrees N) to steppe (54 degrees N). The investigated ecosystems differed strongly in N transformation rates, with highest protein depolymerization and N mineralization rates in middle and southern taiga. All N transformation rates decreased with soil depth following the decrease in organic matter content. Related to protein depolymerization, N mineralization and nitrification were significantly higher in mineral than in organic horizons, supporting a decrease in microbial N limitation with depth. In contrast, we did not find indications for a decrease in microbial N limitation from arctic to temperate ecosystems along the transect. Our findings thus challenge the perception of ubiquitous N limitation at high latitudes, but suggest a transition from N to C limitation of microorganisms with soil depth, even in high-latitude systems such as tundra and boreal forest.

  • Properties and bioavailability of particulate and mineral-associated organic matter in Arctic permafrost soils, Lower Kolyma Region, Russia

    Gentsch N, Mikutta R, Shibistova O, Wild B, Schnecker J, Richter A, Urich T, Gittel A, Santruckova H, Barta J, Lashchinskiy N, Mueller CW, Fuß R, Guggenberger G
    2015 - European Journal of Soil Science, 66: 722-734


    Permafrost degradation may cause strong feedbacks of arctic ecosystems to global warming, but this will depend on if, and to what extent, organic matter (OM) is protected against biodegradation by mechanisms other than freezing and anoxia. Here, we report on the amount, chemical composition and bioavailability of particulate (POM) and mineral-associated OM (MOM) in permafrost soils of the East Siberian Arctic. The average total organic carbon (OC) stock across all soils was 24.0 +/- 6.7 kg m(-2) within 100 cm soil depth. Density fractionation (density cut-off 1.6 g cm(-3)) revealed that 54 +/- 16% of the total soil OC and 64 +/- 18% of OC in subsoil horizons was bound to minerals. As well as sorption of OM to clay-sized minerals (R-2 = 0.80; P < 0.01), co-precipitation of OM with hydrolyzable metals may also transfer carbon into the mineral-bound fraction. Carbon:nitrogen ratios, stable carbon and nitrogen isotopes, C-13-NMR and X-ray photoelectron spectroscopy showed that OM is transformed in permafrost soils, which is a prerequisite for the formation of mineral-organic associations. Mineral-associated OM in deeper soil was enriched in C-13 and N-15, and had narrow C:N and large alkyl C:(O-/N-alkyl C) ratios, indicating an advanced stage of decomposition. Despite being up to several thousands of years old, when incubated under favourable conditions (60% water-holding capacity, 15 degrees C, adequate nutrients, 90 days), only 1.5-5% of the mineral-associated OC was released as CO2. In the topsoils, POM had the largest mineralization but was even less bioavailable than the MOM in subsoil horizons. Our results suggest that the formation of mineral-organic associations acts as an important additional factor in the stabilization of OM in permafrost soils. Although the majority of MOM was not prone to decomposition under favourable conditions, mineral-organic associations host a readily accessible carbon fraction, which may actively participate in ecosystem carbon exchange.

  • The application of ecological stoichiometry to plant-microbial-soil organic matter transformations

    Zechmeister-Boltenstern S, Keiblinger KM, Mooshammer M, Peñuelas J, Richter A, Sardans J, Wanek W
    2015 - Ecological Monographs, 85: 135-155


    Elemental stoichiometry constitutes an inherent link between biogeochemistry and the structure and processes within food webs, and thus is at the core of ecosystem functioning. Stoichiometry allows for spanning different levels of biological organization, from cellular metabolism to ecosystem structure and nutrient cycling, and is therefore particularly useful for establishing links between different ecosystem compartments. We review elemental carbon : nitrogen : phosphorus (C:N:P) ratios in terrestrial ecosystems (from vegetation, leaf litter, woody debris, and dead roots, to soil microbes and organic matter). While the stoichiometry of the plant, litter, and soil compartments of ecosystems is well understood, heterotrophic microbial communities, which dominate the soil food web and drive nutrient cycling, have received increasing interest in recent years. This review highlights the effects of resource stoichiometry on soil microorganisms and decomposition, specifically on the structure and function of heterotrophic microbial communities and suggests several general patterns. First, latitudinal gradients of soil and litter stoichiometry are reflected in microbial community structure and function. Second, resource stoichiometry may cause changes in microbial interactions and community dynamics that lead to feedbacks in nutrient availability. Third, global change alters the C:N, C:P, and N:P ratios of primary producers, with repercussions for microbial decomposer communities and critical ecosystem services such as soil fertility. We argue that ecological stoichiometry provides a framework to analyze and predict such global change effects at various scales.

  • Biological nitrogen fixation and biomass production stability in alfalfa (Medicago sativa L.) genotypes under organic management conditions

    Moghaddam A, Raza A, Vollmann, J, Ardakani MR, Wanek W, Gollner G, Friedel JK
    2015 - Biological Agriculture & Horticulture, 31: 177-192


    Assessments of the stability as well as the performance of plant genotypes across diverse environmental conditions is important for plant breeders as a tool for selecting superior cultivars for the target environments. Alfalfa is the best known fodder crop with a high ability for biological nitrogen fixation (BNF) and high drought tolerance, and it is well-known as an important component of organic farming systems especially in the dry, Pannonian region of east Austria. In a 2-year experiment (2007-2008), 18 alfalfa genotypes from different geographical origins were evaluated under irrigated and rainfed conditions in order to recognize high performance and stable genotypes based on biomass production and BNF in organically managed fields at the University of Natural Resources and Life Sciences, Vienna, Austria. The analysis of variance showed significant differences for the main factors namely year, location, genotype and their interactions in the studied traits. With regard to mean comparisons and stability analysis for shoot dry matter, total biomass yield and BNF, the cultivar Sitel was the best performing genotype followed by PlatoZS, Fix232, Vlasta and Gharghologh. Although additive main effects and multiplicative interaction analysis was found to be more informative in describing the adaptive response of the genotypes, the superiority measure P-i was the best stability parameter to select high yielding and stable genotypes, based on its simplicity of calculation and correlation with crop performance in this study.

  • Storage and transformation of organic matter fractions in cryoturbated permafrost soils across the Siberian Arctic

    Gentsch N, Mikutta R, Alves RJE, Barta J, Capek P, Gittel A, Hugelius G, Kuhry P, Lashchinskiy N, Palmtag J, Richter A, Santrukova H, Schnecker J, Shibistova O, Urich T, Wild B, Guggenberger G
    2015 - Biogeosciences, 12: 4525-4542


    In permafrost soils, the temperature regime and the resulting cryogenic processes are important determinants of the storage of organic carbon (OC) and its small-scale spatial variability. For cryoturbated soils, there is a lack of research assessing pedon-scale heterogeneity in OC stocks and the transformation of functionally different organic matter (OM) fractions, such as particulate and mineral-associated OM. Therefore, pedons of 28 Turbels were sampled in 5m wide soil trenches across the Siberian Arctic to calculate OC and total nitrogen (TN) stocks based on digital profile mapping. Density fractionation of soil samples was performed to distinguish between particulate OM (light fraction, LF, < 1.6 g cm(-3)), mineral associated OM (heavy fraction, HF, > 1.6 g cm(-3)), and a mobilizable dissolved pool (mobilizable fraction, MoF). Across all investigated soil profiles, the total OC storage was 20.2 +/- 8.0 kgm(-2) (mean +/- SD) to 100 cm soil depth. Fifty-four percent of this OC was located in the horizons of the active layer (annual summer thawing layer), showing evidence of cryoturbation, and another 35% was present in the upper permafrost. The HF-OC dominated the overall OC stocks (55 %), followed by LF-OC (19% in mineral and 13% in organic horizons). During fractionation, approximately 13% of the OC was released as MoF, which likely represents a readily bioavailable OM pool. Cryogenic activity in combination with cold and wet conditions was the principle mechanism through which large OC stocks were sequestered in the subsoil (16.4 +/- 8.1 kgm(-2); all mineral B, C, and permafrost horizons). Approximately 22% of the subsoil OC stock can be attributed to LF material subducted by cryoturbation, whereas migration of soluble OM along freezing gradients appeared to be the principle source of the dominant HF (63 %) in the subsoil. Despite the unfavourable abiotic conditions, low C/N ratios and high delta C-13 values indicated substantial microbial OM transformation in the subsoil, but this was not reflected in altered LF and HF pool sizes. Partial least-squares regression analyses suggest that OC accumulates in the HF fraction due to co-precipitation with multivalent cations (Al, Fe) and association with poorly crystalline iron oxides and clay minerals. Our data show that, across all permafrost pedons, the mineral-associated OM represents the dominant OM fraction, suggesting that the HF-OC is the OM pool in permafrost soils on which changing soil conditions will have the largest impact.

  • The effect of warming on the vulnerability of subducted organic carbon in arctic soils

    Capek P, Diakova K, Dickopp JE, Barta J, Wild B, Schnecker J, Alves RJE, Aiglsdorfer S, Guggenberger G, Gentsch N, Hugelius G, Kuhry P, Lashchinsky N, Gittel A, Schleper C, Mikutta R, Palmtag J, Shibistova O, Urich T, Richter A, Santruckova H
    2015 - Soil Biology and Biochemistry, 90: 19-29


    Arctic permafrost soils contain large stocks of organic carbon (OC). Extensive cryogenic processes in these soils cause subduction of a significant part of OC-rich topsoil down into mineral soil through the process of cryoturbation. Currently, one-fourth of total permafrost OC is stored in subducted organic horizons. Predicted climate change is believed to reduce the amount of OC in permafrost soils as rising temperatures will increase decomposition of OC by soil microorganisms. To estimate the sensitivity of OC decomposition to soil temperature and oxygen levels we performed a 4-month incubation experiment in which we manipulated temperature (4-20 degrees C) and oxygen level of topsoil organic, subducted organic and mineral soil horizons. Carbon loss (C-LOSS) was monitored and its potential biotic and abiotic drivers, including concentrations of available nutrients, microbial activity, biomass and stoichiometry, and extracellular oxidative and hydrolytic enzyme pools, were measured. We found that independently of the incubation temperature, C-LOSS from subducted organic and mineral soil horizons was one to two orders of magnitude lower than in the organic topsoil horizon, both under aerobic and anaerobic conditions. This corresponds to the microbial biomass being lower by one to two orders of magnitude. We argue that enzymatic degradation of autochthonous subducted OC does not provide sufficient amounts of carbon and nutrients to sustain greater microbial biomass. The resident microbial biomass relies on allochthonous fluxes of nutrients, enzymes and carbon from the OC-rich topsoil. This results in a "negative priming effect", which protects autochthonous subducted OC from decomposition at present. The vulnerability of subducted organic carbon in cryoturbated arctic soils under future climate conditions will largely depend on the amount of allochthonous carbon and nutrient fluxes from the topsoil. (C) 2015 Elsevier Ltd. All rights reserved.

  • Non-structural carbohydrates in woody plants compared among laboratories

    Quentin AG, Pinkard EA, Ryan MG, Tissue DT, Baggett LS, Adams HD, Maillard P, Marchand J, Landhäusser SM, Lacointe A, Gibon Y, Anderegg WR, Asao S, Atkin OK, Bonhomme M, Claye C, Chow PS, Clément-Vidal A, Davies ND, Dickman LT, Dumbur R, Ellsworth DS, Falk K, Galiano L, Grünzweig JM, Hartmann H, Hoch G, Jones JE, Koike T, Kuhlmann I, Lloret F, Maestro M, Mansfield SD, Martínez-Vilalta J, Maucourt M, McDowell NG, Moing A, Muller B, Nebauer SG, Niinemets U, Palacio S, Piper F, Raveh E, Richter A, Rolland G, Rosas T, Saint Joanis B, Sala A, Smith RA, Sterck F, Stinziano JR, Tobias M, Unda F, Watanabe M, Way DA, Weerasinghe LK, Wild B, Wiley E, Woodruff DR
    2015 - Tree Physiology, 35: 1146-1165


    Non-structural carbohydrates (NSC) in plant tissue are frequently quantified to make inferences about plant responses to environmental conditions. Laboratories publishing estimates of NSC of woody plants use many different methods to evaluate NSC. We asked whether NSC estimates in the recent literature could be quantitatively compared among studies. We also asked whether any differences among laboratories were related to the extraction and quantification methods used to determine starch and sugar concentrations. These questions were addressed by sending sub-samples collected from five woody plant tissues, which varied in NSC content and chemical composition, to 29 laboratories. Each laboratory analyzed the samples with their laboratory-specific protocols, based on recent publications, to determine concentrations of soluble sugars, starch and their sum, total NSC. Laboratory estimates differed substantially for all samples. For example, estimates for Eucalyptus globulus leaves (EGL) varied from 23 to 116 (mean = 56) mg g(-1) for soluble sugars, 6-533 (mean = 94) mg g(-1) for starch and 53-649 (mean = 153) mg g(-1) for total NSC. Mixed model analysis of variance showed that much of the variability among laboratories was unrelated to the categories we used for extraction and quantification methods (method category R(2) = 0.05-0.12 for soluble sugars, 0.10-0.33 for starch and 0.01-0.09 for total NSC). For EGL, the difference between the highest and lowest least squares means for categories in the mixed model analysis was 33 mg g(-1) for total NSC, compared with the range of laboratory estimates of 596 mg g(-1). Laboratories were reasonably consistent in their ranks of estimates among tissues for starch (r = 0.41-0.91), but less so for total NSC (r = 0.45-0.84) and soluble sugars (r = 0.11-0.83). Our results show that NSC estimates for woody plant tissues cannot be compared among laboratories. The relative changes in NSC between treatments measured within a laboratory may be comparable within and between laboratories, especially for starch. To obtain comparable NSC estimates, we suggest that users can either adopt the reference method given in this publication, or report estimates for a portion of samples using the reference method, and report estimates for a standard reference material. Researchers interested in NSC estimates should work to identify and adopt standard methods. © The Author 2015. Published by Oxford University Press. All rights reserved. For Permissions, please email:

  • Contribution of carbonate weathering to the CO2 efflux from temperate forest soils

    Schindlbacher A, Borken W, Djukic I, Brandstätter C, Spötl C, Wanek W
    2015 - Biogeochemistry, 124: 273-290


    Temperate forests provide favorable conditions for carbonate bedrock weathering as the soil CO2 partial pressure is high and soil water is regularly available. As a result of weathering, abiotic CO2 can be released and contribute to the soil CO2 efflux. We used the distinct isotopic signature of the abiotic CO2 to estimate its contribution to the total soil CO2 efflux. Soil cores were sampled from forests on dolomite and limestone and were incubated under the exclusion of atmospheric CO2. Efflux and isotopic signatures of CO2 were repeatedly measured of cores containing the whole mineral soil and bedrock material (heterotrophic respiration + CO2 from weathering) and of cores containing only the mineral top-soil layer (A-horizon; heterotrophic respiration). An aliquot of the cores were let dry out during incubation to assess effects of soil moisture. Although the delta C-13 values of the CO2 efflux from the dolomite soil cores were within a narrow range (A-horizon -26.2 +/- A 0.1 aEuro degrees; whole soil profile wet -25.8 +/- A 0.1 aEuro degrees; whole soil profile dry -25.5 +/- A 0.1 aEuro degrees) the CO2 efflux from the separated A-horizons was significantly depleted in C-13 when compared to the whole soil profiles (p = 0.015). The abiotic contribution to the total CO2 efflux from the dolomite soil cores was 2.0 +/- A 0.5 % under wet and 3.4 +/- A 0.5 % under dry conditions. No abiotic CO2 efflux was traceable from the limestone soil cores. An overall low contribution of CO2 from weathering was affirmed by the amount and C-13 signature of the leached dissolved inorganic carbon (DIC) and the radiocarbon signature of the soil CO2 efflux in the field. Together, our data point towards no more than 1-2 % contribution of abiotic CO2 to the growing season soil CO2 efflux in the field.

  • Landscape-Scale Controls on Aboveground Forest Carbon Stocks on the Osa Peninsula, Costa Rica

    Taylor P, Asner G, Dahlin K, Anderson C, Knapp D, Martin R, Mascaro J, Chazdon R, Cole R, Wanek W, Hofhansl F, Vilchez-Alvaeado B, Townsend A
    2015 - PLoS One, 10: in press


    Tropical forests store large amounts of carbon in tree biomass, although the environmental controls on forest carbon stocks remain poorly resolved. Emerging airborne remote sensing techniques offer a powerful approach to understand how aboveground carbon density (ACD) varies across tropical landscapes. In this study, we evaluate the accuracy of the Carnegie Airborne Observatory (CAO) Light Detection and Ranging (LiDAR) system to detect top-of-canopy tree height (TCH) and ACD across the Osa Peninsula, Costa Rica. LiDAR and field-estimated TCH and ACD were highly correlated across a wide range of forest ages and types. Top-of-canopy height (TCH) reached 67 m, and ACD surpassed 225 Mg C ha-1, indicating both that airborne CAO LiDAR-based estimates of ACD are accurate in tall, high-biomass forests and that the Osa Peninsula harbors some of the most carbon-rich forests in the Neotropics. We also examined the relative influence of lithologic, topoedaphic and climatic factors on regional patterns in ACD, which are known to influence ACD by regulating forest productivity and turnover. Analyses revealed a spatially nested set of factors controlling ACD patterns, with geologic variation explaining up to 16% of the mapped ACD variation at the regional scale, while local variation in topographic slope explained an additional 18%. Lithologic and topoedaphic factors also explained more ACD variation at 30-m than at 100-m spatial resolution, suggesting that environmental filtering depends on the spatial scale of terrain variation. Our result indicate that patterns in ACD are partially controlled by spatial variation in geologic history and geomorphic processes underpinning topographic diversity across landscapes. ACD also exhibited spatial autocorrelation, which may reflect biological processes that influence ACD, such as the assembly of species or phenotypes across the landscape, but additional research is needed to resolve how abiotic and biotic factors contribute to ACD variation across high biomass, high diversity tropical landscapes.

  • Microbial physiology and soil CO2 efflux after 9 years of soil warming in a temperate forest - no indications for thermal adaptations

    Schindlbacher A, Schnecker J, Takriti M, Borken W, Wanek W
    2015 - Global Change Biology, 21: 4265-4277


    Thermal adaptations of soil microorganisms could mitigate or facilitate global warming effects on soil organic matter (SOM) decomposition and soil CO2 efflux. We incubated soil from warmed and control subplots of a forest soil warming experiment to assess whether 9 years of soil warming affected the rates and the temperature sensitivity of the soil CO2 efflux, extracellular enzyme activities, microbial efficiency, and gross N mineralization. Mineral soil (0-10 cm depth) was incubated at temperatures ranging from 3 to 23 °C. No adaptations to long-term warming were observed regarding the heterotrophic soil CO2 efflux (R10 warmed: 2.31 ± 0.15 μmol m(-2) s(-1) , control: 2.34 ± 0.29 μmol m(-2) s(-1) ; Q10 warmed: 2.45 ± 0.06, control: 2.45 ± 0.04). Potential enzyme activities increased with incubation temperature, but the temperature sensitivity of the enzymes did not differ between the warmed and the control soils. The ratio of C : N acquiring enzyme activities was significantly higher in the warmed soil. Microbial biomass-specific respiration rates increased with incubation temperature, but the rates and the temperature sensitivity (Q10 warmed: 2.54 ± 0.23, control 2.75 ± 0.17) did not differ between warmed and control soils. Microbial substrate use efficiency (SUE) declined with increasing incubation temperature in both, warmed and control, soils. SUE and its temperature sensitivity (Q10 warmed: 0.84 ± 0.03, control: 0.88 ± 0.01) did not differ between warmed and control soils either. Gross N mineralization was invariant to incubation temperature and was not affected by long-term soil warming. Our results indicate that thermal adaptations of the microbial decomposer community are unlikely to occur in C-rich calcareous temperate forest soils. © 2015 The Authors. Global Change Biology published by John Wiley & Sons Ltd.

  • Vulnerability of native savanna trees and exotic Khaya senegalensis to seasonal drought

    Arndt SK, Sanders GJ, Bristow M, Hutley LB, Beringer J, Livesley SJ
    2015 - Tree Physiology, 35: 783-791


    Seasonally dry ecosystems present a challenge to plants to maintain water relations. While native vegetation in seasonally dry ecosystems have evolved specific adaptations to the long dry season, there are risks to introduced exotic species. African mahogany, Khaya senegalensis Desr. (A. Juss.), is an exotic plantation species that has been introduced widely in Asia and northern Australia, but it is unknown if it has the physiological or phenotypic plasticity to cope with the strongly seasonal patterns of water availability in the tropical savanna climate of northern Australia. We investigated the gas exchange and water relations traits and adjustments to seasonal drought in K. senegalensis and native eucalypts (Eucalyptus tetrodonta F. Muell. and Corymbia latifolia F. Muell.) in a savanna ecosystem in northern Australia. The native eucalypts did not exhibit any signs of drought stress after 3 months of no rainfall and probably had access to deeper soil moisture late into the dry season. Leaf water potential, stomatal conductance, transpiration and photosynthesis all remained high in the dry season but osmotic adjustment was not observed. Overstorey leaf area index (LAI) was 0.6 in the native eucalypt savanna and did not change between wet and dry seasons. In contrast, the K. senegalensis plantation in the wet season was characterized by a high water potential, high stomatal conductance and transpiration and a high LAI of 2.4. In the dry season, K. senegalensis experienced mild drought stress with a predawn water potential -0.6 MPa. Overstorey LAI was halved, and stomatal conductance and transpiration drastically reduced, while minimum leaf water potentials did not change (-2 MPa) and no osmotic adjustment occurred. Khaya senegalensis exhibited an isohydric behaviour and also had a lower hydraulic vulnerability to cavitation in leaves, with a P50 of -2.3 MPa. The native eucalypts had twice the maximum leaf hydraulic conductance but a much higher P50 of -1.5 MPa. Khaya senegalensis has evolved in a wet-dry tropical climate in West Africa (600-800 mm) and appears to be well suited to the seasonal savanna climate of northern Australia. The species exhibited a large phenotypic plasticity through leaf area adjustments and conservative isohydric behaviour in the 6 months dry season while operating well above its critical hydraulic threshold. © The Author 2015. Published by Oxford University Press. All rights reserved. For Permissions, please email:

  • Microbial community composition shapes enzyme patterns in topsoil and subsoil horizons along a latitudinal transect in Western Siberia

    Schnecker J, Wild B, Takriti M, Eloy Alves RJ, Gentsch N, Gittel A, Hofer A, Klaus K, Knoltsch A, Lashchinskiy N, Mikutta R, Richter A
    2015 - Soil Biology and Biochemistry, 83: 106-115


    Soil horizons below 30 cm depth contain about 60% of the organic carbon stored in soils. Although insight into the physical and chemical stabilization of soil organic matter (SOM) and into microbial community composition in these horizons is being gained, information on microbial functions of subsoil microbial communities and on associated microbially-mediated processes remains sparse. To identify possible controls on enzyme patterns, we correlated enzyme patterns with biotic and abiotic soil parameters, as well as with microbial community composition, estimated using phospholipid fatty acid profiles. Enzyme patterns (i.e. distance-matrixes calculated from these enzyme activities) were calculated from the activities of six extracellular enzymes (cellobiohydrolase, leucine-amino-peptidase, N-acetylglucosaminidase, chitotriosidase, phosphatase and phenoloxidase), which had been measured in soil samples from organic topsoil horizons, mineral topsoil horizons, and mineral subsoil horizons from seven ecosystems along a 1500 km latitudinal transect in Western Siberia. We found that hydrolytic enzyme activities decreased rapidly with depth, whereas oxidative enzyme activities in mineral horizons were as high as, or higher than in organic topsoil horizons. Enzyme patterns varied more strongly between ecosystems in mineral subsoil horizons than in organic topsoils. The enzyme patterns in topsoil horizons were correlated with SOM content (i.e., C and N content) and microbial community composition. In contrast, the enzyme patterns in mineral subsoil horizons were related to water content, soil pH and microbial community composition. The lack of correlation between enzyme patterns and SOM quantity in the mineral subsoils suggests that SOM chemistry, spatial separation or physical stabilization of SOM rather than SOM content might determine substrate availability for enzymatic breakdown. The correlation of microbial community composition and enzyme patterns in all horizons, suggests that microbial community composition shapes enzyme patterns and might act as a modifier for the usual dependency of decomposition rates on SOM content or C/N ratios.

  • Metatranscriptomic census of active protists in soils

    Geisen S, Tveit AT, Clark IM, Richter A, Svenning MM, Bonkowski M, Urich T
    2015 - The ISME Journal, 9: 2178-2190


    The high numbers and diversity of protists in soil systems have long been presumed, but their true diversity and community composition have remained largely concealed. Traditional cultivation-based methods miss a majority of taxa, whereas molecular barcoding approaches employing PCR introduce significant biases in reported community composition of soil protists. Here, we applied a metatranscriptomic approach to assess the protist community in 12 mineral and organic soil samples from different vegetation types and climatic zones using small subunit ribosomal RNA transcripts as marker. We detected a broad diversity of soil protists spanning across all known eukaryotic supergroups and revealed a strikingly different community composition than shown before. Protist communities differed strongly between sites, with Rhizaria and Amoebozoa dominating in forest and grassland soils, while Alveolata were most abundant in peat soils. The Amoebozoa were comprised of Tubulinea, followed with decreasing abundance by Discosea, Variosea and Mycetozoa. Transcripts of Oomycetes, Apicomplexa and Ichthyosporea suggest soil as reservoir of parasitic protist taxa. Further, Foraminifera and Choanoflagellida were ubiquitously detected, showing that these typically marine and freshwater protists are autochthonous members of the soil microbiota. To the best of our knowledge, this metatranscriptomic study provides the most comprehensive picture of active protist communities in soils to date, which is essential to target the ecological roles of protists in the complex soil system.

  • Irrigation water quality influences heavy metal uptake by willows in biosolids

    Laidlaw WS, Baker AJ, Gregory D, Arndt SK
    2015 - Journal of Environmental Management, 155: 31-39


    Phytoextraction is an effective method to remediate heavy metal contaminated landscapes but is often applied for single metal contaminants. Plants used for phytoextraction may not always be able to grow in drier environments without irrigation. This study investigated if willows (Salix x reichardtii A. Kerner) can be used for phytoextraction of multiple metals in biosolids, an end-product of the wastewater treatment process, and if irrigation with reclaimed and freshwater influences the extraction process. A plantation of willows was established directly onto a tilled stockpile of metal-contaminated biosolids and irrigated with slightly saline reclaimed water (EC ∼2 dS/cm) at a wastewater processing plant in Victoria, Australia. Biomass was harvested annually and analysed for heavy metal content. Phytoextraction of cadmium, copper, nickel and zinc was benchmarked against freshwater irrigated willows. The minimum irrigation rate of 700 mm per growing season was sufficient for willows to grow and extract metals. Increasing irrigation rates produced no differences in total biomass and also no differences in the extraction of heavy metals. The reclaimed water reduced both the salinity and the acidity of the biosolids significantly within the first 12 months after irrigation commenced and after three seasons the salinity of the biosolids had dropped to <15% of initial values. A flushing treatment to remove excess salts was therefore not necessary. Irrigation had an impact on biosolids attributes such as salinity and pH, and that this had an influence on metal extraction. Reclaimed water irrigation reduced the biosolid pH and this was associated with reductions of the extraction of Ni and Zn, it did not influence the extraction of Cu and enhanced the phytoextraction of Cd, which was probably related to the high chloride content of the reclaimed water. Our results demonstrate that flood-irrigation with reclaimed water was a successful treatment to grow willows in a dry climate. However, the reclaimed water can also change biosolids properties, which will influence the effectiveness of willows to extract different metals. Copyright © 2015 Elsevier Ltd. All rights reserved.

  • Convergence of soil nitrogen isotopes across global climate gradients

    Craine JM, Elmore AJ, Wang L, Augusto L, Baisden WT, Brookshire EN, Cramer MD, Hasselquist NJ, Hobbie EA, Kahmen A; Koba K, Kranabetter JM, Mack MC, Marin-Spiotta E, Mayor JR, McLauchlan KK, Michelsen A, Nardoto GB, Oliveira RS, Perakis SS, Peri PL, Quesada CA, Richter A, Schipper LA, Stevenson BA, Turner BL, Viani RA, Wanek W, Zeller B
    2015 - Scientific Reports, 5: 8


    Quantifying global patterns of terrestrial nitrogen (N) cycling is central to predicting future patterns of primary productivity, carbon sequestration, nutrient fluxes to aquatic systems, and climate forcing. With limited direct measures of soil N cycling at the global scale, syntheses of the (15)N:(14)N ratio of soil organic matter across climate gradients provide key insights into understanding global patterns of N cycling. In synthesizing data from over 6000 soil samples, we show strong global relationships among soil N isotopes, mean annual temperature (MAT), mean annual precipitation (MAP), and the concentrations of organic carbon and clay in soil. In both hot ecosystems and dry ecosystems, soil organic matter was more enriched in (15)N than in corresponding cold ecosystems or wet ecosystems. Below a MAT of 9.8°C, soil δ(15)N was invariant with MAT. At the global scale, soil organic C concentrations also declined with increasing MAT and decreasing MAP. After standardizing for variation among mineral soils in soil C and clay concentrations, soil δ(15)N showed no consistent trends across global climate and latitudinal gradients. Our analyses could place new constraints on interpretations of patterns of ecosystem N cycling and global budgets of gaseous N loss.

  • A pan-Arctic synthesis of CH4 and CO2 production from anoxic soil incubations

    Treat C, Natali S, Ernakovich J, Iversen CM, Lupascu M, McGuire AD, Norby RJ, Roy Chowdhury T, Richter A, Santruckova H, Schädel C, Schuur EA, Sloan VL, Turetsky MR, Waldrop MP
    2015 - Global Change Biology, 21: 2787-2803


    Permafrost thaw can alter the soil environment through changes in soil moisture, frequently resulting in soil saturation, a shift to anaerobic decomposition, and changes in the plant community. These changes, along with thawing of previously frozen organic material, can alter the form and magnitude of greenhouse gas production from permafrost ecosystems. We synthesized existing methane (CH4 ) and carbon dioxide (CO2 ) production measurements from anaerobic incubations of boreal and tundra soils from the geographic permafrost region to evaluate large-scale controls of anaerobic CO2 and CH4 production and compare the relative importance of landscape-level factors (e.g., vegetation type and landscape position), soil properties (e.g., pH, depth, and soil type), and soil environmental conditions (e.g., temperature and relative water table position). We found fivefold higher maximum CH4 production per gram soil carbon from organic soils than mineral soils. Maximum CH4 production from soils in the active layer (ground that thaws and refreezes annually) was nearly four times that of permafrost per gram soil carbon, and CH4 production per gram soil carbon was two times greater from sites without permafrost than sites with permafrost. Maximum CH4 and median anaerobic CO2 production decreased with depth, while CO2 :CH4 production increased with depth. Maximum CH4 production was highest in soils with herbaceous vegetation and soils that were either consistently or periodically inundated. This synthesis identifies the need to consider biome, landscape position, and vascular/moss vegetation types when modeling CH4 production in permafrost ecosystems and suggests the need for longer-term anaerobic incubations to fully capture CH4 dynamics. Our results demonstrate that as climate warms in arctic and boreal regions, rates of anaerobic CO2 and CH4 production will increase, not only as a result of increased temperature, but also from shifts in vegetation and increased ground saturation that will accompany permafrost thaw. © 2015 John Wiley & Sons Ltd.

  • Storage management influences greenhouse gas emissions from biosolids

    Majumder R, Livesley SJ, Gregory D, Arndt SK
    2015 - Journal of Environmental Management, 151: 361-368


    Biosolids produced by wastewater treatment plants are often stored in stockpiles and can be a significant source of greenhouse gases (GHG). Growing trees in shallow stockpiled biosolids may remove nutrients, keep the biosolids drier and offset GHG emissions through C sequestration. We directly measured methane (CH4), carbon dioxide (CO2) and nitrous oxide (N2O) flux from a large biosolid stockpile and two shallow stockpiles, one planted with Salix reichardtii (willow) trees, from December 2009 to January 2011. All stockpiles emitted large annual amounts of GHG ranging from 38 kg CO2-e Mg(-1) dry biosolid for the large stockpile, to 65 kg CO2-e Mg(-1) for the unplanted shallow stockpile, probably due to the greater surface area to volume ratio. GHG emissions were dominated by N2O and CO2 whilst CH4 emissions were negligible (<2%) from the large stockpile and the shallow stockpiles were actually a CH4 sink. Annual willow tree growth was 12 Mg dry biomass ha(-1), but this only offset 8% of the GHG emissions from the shallow planted stockpile. Our data highlight that biosolid stockpiles are significant sources for GHG emissions but alternate management options such as shallow stockpiles or planting for biomass production will not lead to GHG emission reductions. Copyright © 2015 Elsevier Ltd. All rights reserved.

  • Moss δ(13) C: an accurate proxy for past water environments in polar regions

    Bramley-Alves J, Wanek W, French K, Robinson SA
    2015 - Global Change Biology, 21: 2454-2464


    Increased aridity is of global concern. Polar regions provide an opportunity to monitor changes in bioavailable water free of local anthropogenic influences. However, sophisticated proxy measures are needed. We explored the possibility of using stable carbon isotopes in segments of moss as a fine-scale proxy for past bioavailable water. Variation in δ(13) C with water availability was measured in three species across three peninsulas in the Windmill Islands, East Antarctica and verified using controlled chamber experiments. The δ(13) C from Antarctic mosses accurately recorded long-term variations in water availability in the field, regardless of location, but significant disparities in δ(13) C between species indicated some make more sensitive proxies. δ(13) CSUGAR derived from living tissues can change significantly within the span of an Antarctic season (5 weeks) in chambers, but under field conditions, slow growth means that this technique likely represents multiple seasons. δ(13) CCELLULOSE provides a precise and direct proxy for bioavailable water, allowing reconstructions for coastal Antarctica and potentially other cold regions over past centuries. © 2014 John Wiley & Sons Ltd.

  • Host tree phenology affects vascular epiphytes at the physiological, demographic and community level

    Heinzmann HJR, Beyschlag J, Hofhansl F, Wanek W, Zotz G
    2015 - AoB Plants, 7: 16


    he processes that govern diverse tropical plant communities have rarely been studied in life forms other than trees. Structurally dependent vascular epiphytes, a major part of tropical biodiversity, grow in a three-dimensional matrix defined by their hosts, but trees differ in their architecture, bark structure/chemistry and leaf phenology. We hypothesized that the resulting seasonal differences in microclimatic conditions in evergreen vs. deciduous trees would affect epiphytes at different levels, from organ physiology to community structure. We studied the influence of tree leaf phenology on vascular epiphytes on the Island of Barro Colorado, Panama. Five tree species were selected, which were deciduous, semi-deciduous or evergreen. The crowns of drought-deciduous trees, characterized by sunnier and drier microclimates, hosted fewer individuals and less diverse epiphyte assemblages. Differences were also observed at a functional level, e.g. epiphyte assemblages in deciduous trees had larger proportions of Crassulacean acid metabolism species and individuals. At the population level a drier microclimate was associated with lower individual growth and survival in a xerophytic fern. Some species also showed, as expected, lower specific leaf area and higher δ(13)C values when growing in deciduous trees compared with evergreen trees. As hypothesized, host tree leaf phenology influences vascular epiphytes at different levels. Our results suggest a cascading effect of tree composition and associated differences in tree phenology on the diversity and functioning of epiphyte communities in tropical lowland forests. Published by Oxford University Press on behalf of the Annals of Botany Company.

  • Summer drought alters carbon allocation to roots and root respiration in mountain grassland

    Hasibeder R, Fuchslueger L, Richter A, Bahn M
    2015 - New Phytologist, 3: 1117-1127


    Drought affects the carbon (C) source and sink activities of plant organs, with potential consequences for belowground C allocation, a key process of the terrestrial C cycle. The responses of belowground C allocation dynamics to drought are so far poorly understood. We combined experimental rain exclusion with (13)C pulse labelling in a mountain meadow to analyse the effects of summer drought on the dynamics of belowground allocation of recently assimilated C and how it is partitioned among different carbohydrate pools and root respiration. Severe soil moisture deficit decreased the ecosystem C uptake and the amounts and velocity of C allocated from shoots to roots. However, the proportion of recently assimilated C translocated belowground remained unaffected by drought. Reduced root respiration, reflecting reduced C demand under drought, was increasingly sustained by C reserves, whilst recent assimilates were preferentially allocated to root storage and an enlarged pool of osmotically active compounds. Our results indicate that under drought conditions the usage of recent photosynthates is shifted from metabolic activity to osmotic adjustment and storage compounds. © 2014 The Authors. New Phytologist © 2014 New Phytologist Trust.

  • Exploring the transfer of recent plant photosynthates to soil microbes: mycorrhizal pathway vs direct root exudation

    Kaiser C, Kilburn MR, Clode PL, Fuchslueger L, Koranda M, Cliff JB, Solaiman ZM, Murphy DV
    2015 - New Phytologist, 205: 1537-1551


    Plants rapidly release photoassimilated carbon (C) to the soil via direct root exudation and associated mycorrhizal fungi, with both pathways promoting plant nutrient availability. This study aimed to explore these pathways from the root's vascular bundle to soil microbial communities. Using nanoscale secondary ion mass spectrometry (NanoSIMS) imaging and (13) C-phospho- and neutral lipid fatty acids, we traced in-situ flows of recently photoassimilated C of (13) CO2 -exposed wheat (Triticum aestivum) through arbuscular mycorrhiza (AM) into root- and hyphae-associated soil microbial communities. Intraradical hyphae of AM fungi were significantly (13) C-enriched compared to other root-cortex areas after 8 h of labelling. Immature fine root areas close to the root tip, where AM features were absent, showed signs of passive C loss and co-location of photoassimilates with nitrogen taken up from the soil solution. A significant and exclusively fresh proportion of (13) C-photosynthates was delivered through the AM pathway and was utilised by different microbial groups compared to C directly released by roots. Our results indicate that a major release of recent photosynthates into soil leave plant roots via AM intraradical hyphae already upstream of passive root exudations. AM fungi may act as a rapid hub for translocating fresh plant C to soil microbes. © 2014 The Authors New Phytologist © 2014 New Phytologist Trust.

  • New insights into mechanisms driving carbon allocation in tropical forests

    Hofhansl F, Schnecker J, Singer G, Wanek W
    2015 - New Phytologist, 205: 137-146


    The proportion of carbon allocated to wood production is an important determinant of the carbon sink strength of global forest ecosystems. Understanding the mechanisms controlling wood production and its responses to environmental drivers is essential for parameterization of global vegetation models and to accurately predict future responses of tropical forests in terms of carbon sequestration. Here, we synthesize data from 105 pantropical old-growth rainforests to investigate environmental controls on the partitioning of net primary production to wood production (%WP) using structural equation modeling. Our results reveal that %WP is governed by two independent pathways of direct and indirect environmental controls. While temperature and soil phosphorus availability indirectly affected %WP via increasing productivity, precipitation and dry season length both directly increased %WP via tradeoffs along the plant economics spectrum. We provide new insights into the mechanisms driving %WP, allowing us to conclude that projected climate change could enhance %WP in less productive tropical forests, thus increasing carbon sequestration in montane forests, but adversely affecting lowland forests. © 2014 The Authors. New Phytologist © 2014 New Phytologist Trust.

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