Browsing by Person "Kandeler, Ellen"

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Biochar amendment for C sequestration in a temperate agroecosystem : implications for microbial C- and N-cycling
(2018) Bamminger, Chris; Kandeler, Ellen
Climate warming will have great impact on terrestrial ecosystems. Different soil properties such as temperature and moisture will be altered, thereby influencing C- and N-cycles, microbial activity as well as plant growth. This may contribute to the observed increase in soil greenhouse gas (GHG) emissions under climate change. Therefore, new options are needed to mitigate theses projected consequences. Biochar is primarily suggested to be effective in long-term C sequestration in agricultural soils due to its long-term stability. In addition, it could be applied to improve various soil properties, plant growth and to reduce soil GHG emissions. To date, knowledge about such beneficial biochar effects in soil under predicted warming climate is extremely scarce. In the first study, a slow-pyrolysis biochar from Miscanthus x giganteus feedstock (600 °C, 30 Min.) was incubated for short time (37d) under controlled laboratory conditions in agricultural soil in the presence of earthworms and N-rich litter (Phacelia tanacetifolia Benth.). Biochar increased microbial abundances and the fungal-to-bacterial PLFA ratio after 37 days in arable soil applied with litter suggesting improved living conditions for microorganisms with biochar. Fungi may benefit most from newly created habitats due to colonizable biochar pores and surfaces. Additionally, fungi could have also mineralized small amounts of recalcitrant biochar-C during plant litter decomposition. Without litter, biochar led to interactions between earthworms and soil microorganisms resulting in enhanced bacterial and fungal abundances. This indicates better growth habitats for soil microbes in earthworm casts containing biochar. Soil respiration and metabolic quotients (qCO2) and N2O emissions (in litter treatments) were decreased after biochar application suggesting a more efficient microbial community and underscoring the GHG mitigation potential of the used biochar. The field experiment, investigated in the second and third study, focused on the stability and long-term soil C sequestration potential of comparable Miscanthus biochar (850 °C, 30 Min.). Related effects on soil GHG emissions, physical, chemical and microbiological soil properties as well as plant growth were determined in an agroecosystem at year-round elevated soil temperature (+2.5 °C, since 2008). The second study investigated the short-term effects of biochar on microbial abundances and growth of winter rapeseed during the first year after field application to a warmed temperate arable soil. It was found that fungal biomass and the fungal-to-bacterial ratio were increased in the warmed biochar plots only after three months in the presence of spring barley litter from the previous growing season. The disappearance of this effect points to an overall high stability of the investigated biochar. Moreover, biochar proved to be effective in mitigating negative effects of seasonal dryness on microbial abundances and early plant growth in the dry spring period in 2014. However, biochar had no effect on final aboveground biomass of winter rapeseed at harvest in the first growing season. As shown in the third study, after two vegetation periods of winter rapeseed and spring wheat, the assumption that plant productivity in already fertile temperate arable soils is unlikely to be further enhanced with biochar amendment, was confirmed. Total CO2 emissions after two years were not reduced by biochar and remained unchanged even under warming suggesting a high degradation stability of the used biochar. N2O emissions were increased in biochar-amended soil at elevated soil temperature, presumably due to enhanced water and fertilizer retention with biochar. By using the global warming potential (GWP100) of total soil GHG emissions, the storage of biochar-C in soil was estimated to compensate warming-induced elevated soil GHG emissions for 20 years. To conclude, this thesis revealed that biochar may have only minor influence on soil microorganisms and crop growth in temperate, fertile arable field soils. However, it was shown that biochar could be a valuable tool for C sequestration in temperate arable soils, thus potentially offsetting a warming-induced increase in GHG emissions. In order to face climate change impacts, more long-term studies on microbiological effects and the C sequestration potential of biochar in cultivated soil under warming are urgently needed.
Biological regulation of subsoil C-cycling
(2019) Preußer, Sebastian; Kandeler, Ellen
Soils are the largest terrestrial reservoir of organic carbon (OC). Substantial proportions of the stored OC are found in stabilized form in deeper soil layers. Beside the quality and quantity of C input from plant biomass, C storage in soil is primarily controlled by the microbial decomposition capacity. Various physical, chemical and biological factors (e.g., substrate availability, temperature, water content, pH, texture) vary within soil profiles and directly or indirectly influence the abundance, composition and activity of microbial communities and thus the microbial C turnover. While soil microbiological research has so far focused mainly on processes in topsoil, the mechanisms of C storage and turnover in subsoil are largely unknown. The objective of the present thesis was therefore to investigate the specific influence of substrate availability and different environmental factors as well as their interactions on microbial communities and their regulatory function in subsoil C-cycling. This objective was addressed in three studies. In the first and second study, one-year field experiments were established in which microbial communities from different soil depths were exposed to altered habitat conditions to identify crucial factors influencing the spatial and temporal development of microbial abundance and substrate utilization within soil profiles. This was achieved by reciprocal translocation of soils between subsoil horizons (first study) and topsoil and subsoil horizons (second study) in combination with addition of 13C-labelled substrates and different sampling dates. In the third study, a flow cascade experiment with soil columns from topsoil and subsoil horizons and soil minerals (goethite) coated with 13C-labelled organic matter (OM) was established. This laboratory experiment investigated the importance of exchange processes of OM with reactive soil minerals for the quality and quantity of dissolved OM and the influence of these soil micro-habitats on microbial abundance and community composition with increasing soil depth. In the first study, the reciprocal translocation of subsoils from different soil depths revealed that due to comparable micro-climatic conditions and soil textures within the subsoil profile, no changes in microbial biomass, community composition and activity occurred. Moreover, increasing microbial substrate utilization in relation to the quantity of added substrate indicated that deep soil layers exhibit high potential for microbial C turnover. However, this potential was constrained by low soil moisture in interplay with the coarse soil texture and the resulting micro-scale fragmentation of the subsoil environment. The bacterial substrate utilization was more affected by this spatial separation between microorganisms and potentially available substrate than that of fungi, which was further confirmed by the translocation experiment with topsoil and subsoil in the second study. While the absolute substrate utilization capacity of bacteria decreased from the more moist topsoil to the drier subsoil, fungi were able to increase their substrate utilization and thus to partially compensate the decrease in C input from other sources. Furthermore, the addition of root litter as a preferential C source of fungal decomposer communities led to a pronounced fungal growth in subsoil. The third study demonstrated the high importance of reactive soil minerals both in topsoil and in subsoil for microbial growth due to extensive exchange processes of OM and the associated high availability of labile C. In particular copiotrophic bacteria such as Betaproteobacteria benefited from the increased C availability under non-limiting water conditions leading to a pronounced increase in bacterial dominance in the microbial communities of these soil micro-habitats. In conclusion, this thesis showed that subsoil exhibits great potential for both bacterial and fungal C turnover, albeit this potential is limited by various factors. This thesis, however, allowed to determine the specific effects of these factors on bacteria and fungi and their function in subsoil C-cycling and thus to identify those factors of critical importance. The micro-climate in subsoil, in particular soil moisture, was the primary factor limiting bacterial growth and activity, whereas fungi were more strongly restricted by substrate limitations.
Carbon and nitrogen transformations in alpine ecosystems of the Eastern Alps, Austria
(2007) Koch, Oliver; Kandeler, Ellen
This thesis investigated net CH4 and net CO2 emissions from sites in the alpine region of the Eastern Alps, Austria. Four mature alpine sites (one dry meadow and three fen sites) were selected and the influence of abiotic (radiation, temperature, soil water conditions) and biotic (above-ground standing plant biomass) environmental controls on diurnal and seasonal emission patterns were studied. For a better understanding of the response of soil C- and N pools to global warming, the temperature sensitivity of activities involved in C- and N cycling were determined. The first part of the thesis dealt with net methane fluxes measured over a period of 24 months. During snow-free periods, average methane emissions of the fen sites ranged between 19 and 116 mg CH4 m-2 d-1. Mean emissions during snow periods were much lower, being 18 to 59% of annual fluxes. The alpine dry meadow functioned as a small methane sink during snow-free periods (-2.1 mg CH4 m-2 d-1 (2003); -1.0 mg CH4 m-2 d-1 (2004)). The diurnal and seasonal methane uptake of the dry meadow was positively related to soil temperature and negatively related to water-filled pore space (wfps). In the fen, the seasonal methane fluxes were related to soil temperature and groundwater table. The live above-ground standing plant biomass contributed to net methane fluxes only at those sites with higher water table positions. This study provided evidence that alpine fens acted as methane sources throughout the year, whereas an alpine meadow site acted as a net methane sink during snow-free periods. In the second part of the thesis the CO2 balance was estimated based on diurnal flux measurements and on the influence of photosynthetic active radiation (PAR), plant green area index (GAI), soil temperature and wfps. The daylight net ecosystem CO2 emission rate was influenced by PAR and GAI throughout snow-free seasons. The seasonal net CO2 emission rate at night was positively related to soil temperature, while low wfps reduced flux rates at the meadow and at the driest fen study site but reinforced carbon loss at the wetter fen sites. The daily average ecosystem net CO2 gain during snow-free periods at the meadow was 3.5 g CO2 m-2 d-1 and at the fen sites between 1.5 and 3.4 g CO2 m-2 d-1. The mean average daily CO2 emission during snow periods was low, being -0.9 g CO2 m-2 d-1 for the meadow and between -0.2 and -0.7 g CO2 m-2 d-1 for all fen sites. All sites function as significant annual net carbon sinks, with a net carbon gain from 50 to 121 g C m-2 a-1 (averaged over both years), irrespective of water balance. The results indicate that alpine fen sites, that have built up a large carbon stock in the past, are not expected to gain a further carbon surplus compared with meadows under the current climate. Temperature is important for regulating biological activities. The third part of the thesis focused on temperature sensitivity of soil C mineralization, N mineralization and potential enzyme activities involved in the C- and N cycle (ß-glucosidase, ß-xylosidase, N-acetyl-ß-glucosaminidase, tyrosine aminopeptidase, leucine aminopeptidase) over a temperature range of 0-30°C. The objective was to calculate Q10 values and relative temperature sensitivities (RTS) and to quantify seasonal (summer, autumn, winter) and site-specific factors. The Q10 values of C mineralization were significantly higher (average 2.0) than for N mineralization (average 1.7). The Q10 values of both activities were significantly negatively related to soil organic matter quality. In contrast, the chemical soil properties, microbial biomass and sampling date did not influence Q10 values. Analysis of RTS showed that the temperature sensitivity increased with decreasing temperature. The C- and N mineralization and potential aminopeptidase activities (tyrosine, leucine) showed an almost constant temperature dependence over 0-30°C. In contrast, ß-glucosidase, ß-xylosidase and N-acetyl-ß-glucosaminidase showed a distinctive increase in temperature sensitivity with decreasing temperature. Low temperature at the winter sampling date caused a greater increase in the RTS of all activities than in autumn and summer. Our results indicate a disproportion of the RTS for potential enzyme activities of the C- and N cycle and a disproportion of the RTS for easily degradable C compounds (ß-glucose, ß-xylose) compared with the C mineralization of soil organic matter. Thus, temperature may play an important role in regulating the decay of different soil organic matter fractions.
Community Structure and Activity of Nitrate-Reducing Microorganisms in Soils under Global Climate Change
(2006) Deiglmayr, Kathrin; Kandeler, Ellen
Since the beginning of the Industrial Revolution, atmospheric carbon dioxide concentrations have been steadily increasing and, thus, contributed to a warming of the climate and altered biogeochemical cycles. To study the response of soil microorganisms to altered environmental conditions under global climate change, the nitrate-reducing community was regarded as a model community in the present thesis. This functional group, which performs the first step in the denitrification pathway, was selected because it is phylogenetically very diverse. In particular rising levels of atmospheric carbon dioxide as the most important catalyst of temperature rise and the retreat of glaciers in the Alps as one of the most evident consequences of climate change were investigated. The behaviour of nitrate reducers was investigated in a biphasic approach: (i) at the level of its enzyme activity of the nitrate reductase and (ii) at the level of community structure, which was characterised by RFLP (Restriction Fragment Length Polymorphism)-fingerprints using the functional gene narG. The effect of elevated atmospheric carbon dioxide concentrations on nitrate-reducing micro-organisms was studied in the Swiss FACE (Free Air Carbon dioxide Enrichment) experiment including the rhizosphere of two functional plant types (Lolium perenne and Trifolium repens), two N fertilisation levels and two sampling dates (June and October 2002). Whereas in June no significant treatment effect was observed, the nitrate reductase activity proved to be significantly reduced under elevated atmospheric carbon dioxide at the autumn sampling date. Simultaneously, elevated enzyme activities were recorded under Trifolium repens and high N fertilisation pointing to a control of nitrate reductase activity by nitrate availability at the time of sampling. The community structure of nitrate reducers, however, showed a different response pattern with sampling date and the strongly varying pH of the different experimental plots constituting the main driving factors. With respect to the three experimental factors atmospheric carbon dioxide, plant type and N fertilisation the composition of the nitrate reducers revealed a high stability. The microbial succession of nitrate-reducing microorganisms was studied in the rhizosphere of Poa alpina across the glacier foreland of the Rotmoosferner/Oetz valley. Sampling was performed in August and at the end of the short period of vegetation in September. The nitrate reductase activity increased significantly with progressing successional age, whereas organic carbon together with nitrate concentrations in the soils explained the major part of this effect. The microbial community of nitrate reducers revealed a significant shift across the glacier foreland, with pH and organic carbon representing the most important environmental factors inducing this shift. A detailed analysis of the clone libraries that were constructed for the youngest and the oldest site in the glacier foreland pointed to the tendency of lower diversity in the late succession compared to the young succession. Possibly an increasing selective pressure due to higher densities of microorganisms and, hence, a higher competition for limited resources contributed to the decline in diversity. In conclusion, the functional group of nitrate reducers responded to changing environmental conditions under global climate change particularly through altered enzyme activities. The amount and the direction of this response depended strongly on the nitrate availability and the organic carbon content in soils. The community structure of nitrate-reducing microorganisms, however, proved to be resilient towards short-term substrate fluctuations. This indicates that the genetic pool of this group of soil microorganisms possesses a high functional stability characterized by a relatively persistent composition and an independent modulation of enzyme activity.
Constraints on microbial pesticide degradation in soils
(2023) Wirsching, Johannes; Kandeler, Ellen
Pesticides are an essential component of intensified agriculture and have contributed significantly to the increase in food production observed in recent decades. Since 1960, pesticide use has increased by a factor of fifteen to twenty, representing a market value of $40 billion in 2016. Soil monitoring campaigns to track pesticide contamination of croplands across Europe are quantifying pesticide residues whose residence times in soils exceed expected values. Diffuse contamination by pesticide residues raises concerns about soil functions, soil biodiversity, and food safety, as well as the transport of contaminants by wind and water to surface waters or to adjacent, organically managed croplands. Data on the frequency of occurrence and concentrations of pesticide residues in soil demonstrate a discrepancy between the determination of persistence and subsequent approval and their actual fate in soil. This raises the question of whether degradability of individual organic compounds has been adequately studied. Microbiological degradation is the most important process for reducing pesticide loads in soils. A reliable estimate of pesticide residence time requires an expanded understanding of the factors limiting microbial degradation. Degradation of anthropogenic organic chemicals in soils occurs much more slowly than would be expected based on their physicochemical properties. While processes that determine the fate of pesticides in soil have often been studied at different spatial and temporal scales, reasons for discrepancies between the observed complete degradation of pesticides under laboratory conditions and their persistence in the field remain unclear. This thesis addresses this challenge by focusing on the central question of why inherently biodegradable compounds in soils display increased persistence under field conditions. Organic contaminants in low but detectable and environmentally significant concentrations could remain in the soil once available concentrations fall below a threshold where bioenergetic growth restrictions come into play. In addition, potential microbial and biophysical limitations and environmental factors such as soil temperature and soil moisture are often examined separately in current degradation studies. Combinations of temperature and soil moisture changes associated with different concentration levels have been less well examined, resulting in an incomplete understanding of the degradation process. Another key factor in the demonstrated discrepancy between predicted and actual persistence in the field could be due to laboratory experiments that cannot account for field-scale processes. Therefore, degradation rates determined in laboratory experiments cannot be confidently extrapolated to the field scale. . This thesis identified further important regulatory mechanisms for microbially mediated pesticide degradation. The previously unknown concentration-dependent degradation dynamics and the concentration-dependent influence of limiting environmental conditions on microbial degradation emphasize the importance of studies using a realistic concentration range. Evidence of deep transport of a highly sorptive pesticide such as glyphosate primarily via preferential flow pathways into the subsoil with lower degradation dynamics underscores the need to include processes that can only be verified in field studies as part of risk assessments. The results of this thesis suggest that the biodegradation rates of pesticides are not homogeneous at field scales and may account in part for the discrepancy between complete degradation of pesticides under laboratory conditions and their persistence in the field. Laboratory studies in which soil samples are pooled and mixed to obtain a single "representative" sample can provide a simplified understanding of the process, but the complexity, particularly that of soil heterogeneity, of pesticide distribution and microbial degradation associated with prevailing climatic conditions, requires calibration of previously used methods in field studies and possibly at landscape, watershed, or regional scales. The scale-dependent degradation aspect will become even more important in the future; as soil properties and processes that control the toxicological aspects of contaminants include temperature and moisture, and changes associated with climate change will lead to an increase in extreme precipitation, longer dry periods, and soil erosion.
Effect of reduced nitrogen deposition on microbial activity, abundance and diversity in forest soils
(2012) Enowashu, Esther Eneckeh; Kandeler, Ellen
The deposition of nitrogen has increased many-fold due to anthropogenic activities. Since forest ecosystems are often limited by N availability, elevated N inputs from the atmosphere can have a fertilization effect but in the long-term, excess N can influence above- and below-ground production. One of the consequences of N deposition and increased N inputs is a shift in microbial community structure and function as ecosystems move towards N saturation. Soil microorganisms through the action of enzymes play an important role in N dynamics. Thus, the availability and turnover of N depends strongly on microbial abundance, diversity and activity which are in turn influenced by soil properties. Studies on the effects of high nitrogen inputs and the response of forest ecosystems to nitrogen saturation are many and well understood. However, the reversibility of N-induced shifts in forest ecosystem processes is largely unknown. This thesis was therefore designed to study the response of soil microorganisms to reduced N deposition. A biphasic approach was employed to look into (i) the general microbial functional status of the Solling forest site as well as (ii) the microbial community structure which may be a key regulator of two important processes of N transformation: denitrification and proteolysis. The goal of the present thesis was addressed in three studies. Denitrification is considered sensitive to environmental changes and the response of nitrate-reducers and denitrifiers to reduced N deposition was determined in the first study. The goal of the second study was to investigate the overall microbial activity of the Solling forest profiles especially focussing on enzymes involved in the N cycle. This revealed a pronounced activity of peptidases whereby a set of novel pepN primers encoding alanine aminopeptidase enzyme was designed in the third study to determine the group of bacteria involved in proteolysis in forest as well as agricultural and grassland soils. The Solling experimental station was established more than two decades ago and it gave the opportunity to study the N cycle in a natural forest ecosystem at different sampling dates and depths. A combination of classical biological methods and modern molecular techniques were used in the studies. Soil physico-chemical parameters (OC, Nt, NO3-, NH4+, pH, % Water content) were analysed to gain more information on mineralization and immobilization of N in the soil profiles. The analysis of microbial biomass, ergosterol content and the activity of several enzymes of the N, C and P cycles as well as enzyme activity of nitrate reducers was determined in order to interpret microbial functions. The abundance of nitrate reducers and denitrifiers were determined by quantitative PCR of 16S rRNA, nitrate reductase (narG and napA) and denitrification (nirK, nirS and nosZ) genes. The diversity of peptide degrading bacteria was analysed by PCR, cloning and sequencing and the construction of pepN gene libraries. The results of the first study indicated that time and space were the main drivers influencing the abundance and activity of the nitrate reducers and denitrifier communities in the forest soil profiles. Reduced N deposition had a of minimal effect. Interestingly, the ratios of nosZ to16S rRNA gene and nosZ to nirK increased with soil depth thereby tempting to conclude that the size of denitriﬁers capable of reducing N2O into N2 might be bigger in the mineral horizons. In the second study, a stronger response of N cycling enzymes to reduced N deposition could be seen. However, these responses especially that of specific peptidases differed in magnitude which could be indicative of a modification of the reaction rates of the different N cycling enzymes. Correlation of nutrients (N, C, P) with microbial biomass and enzyme activities in the soil profiles revealed that substrate availability was the main factor influencing microbial activity. In the third study, analyses of gene libraries from extracted DNA from forest, agricultural and glacier soil samples revealed a high diversity of pepN sequences related to mainly α-Proteobacteria. A majority of the sequences showed similarity to published data revealing that the amplified region of pepN might be conserved. Linking diversity and enzymatic data, lowest diversity was observed in the agricultural soil where activity levels of alanine aminopeptidase were lowest indicating the importance of diversity studies for ecosystem functioning. In conclusion, this thesis offers valuable contributions to understanding the impact of N deposition. The approach used was suitable to assess the response of the different microbial communities to reduced N deposition. The magnitude of the response depended strongly on space, time and substrate availability in soils as well as their interactions.
Effectiveness of bio-effectors on maize, wheat and tomato performance and phosphorus acquisition from greenhouse to field scales in Europe and Israel: a meta-analysis
(2024) Nkebiwe, Peteh Mehdi; Stevens Lekfeldt, Jonas D.; Symanczik, Sarah; Thonar, Cécile; Mäder, Paul; Bar-Tal, Asher; Halpern, Moshe; Biró, Borbala; Bradáčová, Klára; Caniullan, Pedro C.; Choudhary, Krishna K.; Cozzolino, Vincenza; Di Stasio, Emilio; Dobczinski, Stefan; Geistlinger, Joerg; Lüthi, Angelika; Gómez-Muñoz, Beatriz; Kandeler, Ellen; Kolberg, Flora; Kotroczó, Zsolt; Kulhanek, Martin; Mercl, Filip; Tamir, Guy; Moradtalab, Narges; Piccolo, Alessandro; Maggio, Albino; Nassal, Dinah; Szalai, Magdolna Zita; Juhos, Katalin; Fora, Ciprian G.; Florea, Andreea; Poşta, Gheorghe; Lauer, Karl Fritz; Toth, Brigitta; Tlustoš, Pavel; Mpanga, Isaac K.; Weber, Nino; Weinmann, Markus; Yermiyahu, Uri; Magid, Jakob; Müller, Torsten; Neumann, Günter; Ludewig, Uwe; de Neergaard, Andreas
Biostimulants (Bio-effectors, BEs) comprise plant growth-promoting microorganisms and active natural substances that promote plant nutrient-acquisition, stress resilience, growth, crop quality and yield. Unfortunately, the effectiveness of BEs, particularly under field conditions, appears highly variable and poorly quantified. Using random model meta-analyses tools, we summarize the effects of 107 BE treatments on the performance of major crops, mainly conducted within the EU-funded project BIOFECTOR with a focus on phosphorus (P) nutrition, over five years. Our analyses comprised 94 controlled pot and 47 field experiments under different geoclimatic conditions, with variable stress levels across European countries and Israel. The results show an average growth/yield increase by 9.3% (n=945), with substantial differences between crops (tomato > maize > wheat) and growth conditions (controlled nursery + field (Seed germination and nursery under controlled conditions and young plants transplanted to the field) > controlled > field). Average crop growth responses were independent of BE type, P fertilizer type, soil pH and plant-available soil P (water-P, Olsen-P or Calcium acetate lactate-P). BE effectiveness profited from manure and other organic fertilizers, increasing soil pH and presence of abiotic stresses (cold, drought/heat or salinity). Systematic meta-studies based on published literature commonly face the inherent problem of publication bias where the most suspected form is the selective publication of statistically significant results. In this meta-analysis, however, the results obtained from all experiments within the project are included. Therefore, it is free of publication bias. In contrast to reviews of published literature, our unique study design is based on a common standardized protocol which applies to all experiments conducted within the project to reduce sources of variability. Based on data of crop growth, yield and P acquisition, we conclude that application of BEs can save fertilizer resources in the future, but the efficiency of BE application depends on cropping systems and environments.
Effects of elevated soil temperature and altered precipitation patterns on N-cycling and production of N2O and CO2 in an agricultural soil
(2016) Latt, Yadana Khin; Kandeler, Ellen
Both temperature and precipitation regimes are expected to change with climate change and are, at the same time, major environmental factors regulating biogeochemical cycles in terrestrial ecosystems. Therefore, crop water availability, soil nitrogen transformations, losses, and uptake by plants as well as CO2 emissions from soil are likely to be changed by climate change. Agriculture is known to be one of the most important human activities for releasing significant amounts of N2O and CO2 to the atmosphere. Due to global concern about the changing climate, there has been a great interest in reducing emissions of N2O and CO2 from agricultural soils. CO2 and N2O are produced in soil primarily by microbial processes. Their production and emissions from the soil are controlled by a number of environmental variables including inorganic N availability, soil temperature and water content. Agricultural management practices, such as irrigation, affect these environmental variables and thus have the potential to dramatically alter N2O and CO2 emissions from the soil. The present study is titled "Effects of elevated soil temperature and altered precipitation patterns on N cycling and production of N2O and CO2 in an agricultural soil". The objectives of this study were: to determine the effects of elevated soil temperature on N cycling in a winter wheat cropping system, to investigate the short-term response of N2O and CO2 fluxes during rewetting of soils after extended dry periods in summer, and to determine the effects of different degrees of rewetting on the CO2 emission peaks after rewetting in laboratory incubations. In the 1st experiment, we used the Hohenheim Climate Change (HoCC) experiment in Stuttgart, Germany, to test the hypothesis that elevated soil temperature will increase microbial N cycling, plant N uptake and wheat growth. In the HoCC experiment, soil temperature is elevated by 2.5°C at 4 cm depth. This experiment was conducted at non-roofed plots (1m x 1m) with ambient (Ta) and elevated (Te) soil temperature and with ambient precipitation. In 2012, winter wheat (Triticum aestivum) was planted. C and N concentrations in soil and aboveground plant fractions, soil microbial biomass C and N (Cmic and Nmic), mineral N content (NH4+ - N and NO3- - N), potential nitrification and enzymes involved in nitrogen cycling were analyzed at soil depths of 0-15 and 15-30 cm at five sampling dates. The plants were rated weekly for their phenological development and senescence behavior. We found that an increase in soil temperature by 2.5oC did not have a persistent effect on mineral N content and the activity of potential nitrification within the soil. Plant growth development also did not respond to increased soil temperature. However microbial biomass C and N, and some enzyme activities involved in N-cycling, tended to increase under elevated soil temperature. Overall, the results of this study suggested that soil warming by 2.5oC slightly stimulates soil N cycling but does not alter plant growth development. In the 2nd experiment, in 2013, the effects of a change in the amount and frequency of precipitation patterns on N2O and CO2 emissions were studied after the two dry periods in summer in the HoCC experiment. N2O and CO2 gas samples were taken from four subplots (1m x 1m) of each roofed plot exposed to ambient (Ta) or elevated (Te) soil temperature and four precipitation manipulations (ambient plot, reduced precipitation amount, reduced precipitation frequency, and reduced precipitation amount and frequency). We found that CO2 emissions were affected only by temperature, but not by precipitation pattern. It can be said that N2O and CO2 emissions after rewetting of dry soil were not altered by changing precipitation patterns during dry periods in summer. In the year 2014, using laboratory incubations, we also measured the short-term response of CO2 production to a rewetting of dry soil to different volumetric water contents for 24 hours. This study was conducted by manipulating microcosms with agricultural soil from the HoCC experimental site, which had been exposed to severe drought conditions of three months duration for each of the last six years. The results showed that CO2 production increased with increases in the water content of soils by rewetting at 5%, 15%, 25%, 35% and 45% VWC. With increasing water additions more peaks in CO2 production were detected and different temporal patterns of CO2 emission were affected by adding different amounts of water. It might be due to the fact that with greater water additions successively larger pore sizes were water filled and therefore different bacterial groups located in different pore size classes might have contributed to CO2 production. In summary, the results from field study suggested that climate warming will affect N cycling in soils in an agricultural cropping system. The results from both field and microcosm rewetting experiments contribute to a better understanding of C and N dynamics in soil by investigating the effect of varying soil water content on the emission of N2O and CO2.
Effects of resource availability and quality on soil microorganisms and their carbon assimilation
(2014) Kramer, Susanne; Kandeler, Ellen
Soil microorganisms play a pivotal role in decomposition processes and therefore influence nutrient cycling and ecosystem function. Availability and quality of resources determines activity, growth and identity of substrate users. In agricultural systems, availability of resources is dependent on, for example, crop type, management, season, and depth. At depth substrate availability and microbial biomass decrease. However, there remain gaps in our understanding of C turnover in subsoil and how processes in the topsoil may influence abundance, activity, and function of microorganisms in deeper soil layers. With respect to substrate quality it is thought that bacteria are the dominant users of high quality substrates and more labile components whereas fungi are more important for the degradation of low quality and more recalcitrant substrates (i.e. cellulose, lignin). Therefore, this thesis was designed to increase our understanding of C turnover and the influence of both availability and quality of substrates on microorganisms in an agricultural soil. In the first and second studies, a recently established C3-C4 plant exchange field experiment was used to investigate the C flow from belowground (root) and aboveground (shoot litter) resources into the belowground food web. Maize plants were cultivated to introduce a C4 signal into the soil both by plant growth (belowground / root channel) and also by applying shoot litter (aboveground litter channel). To separate C flow from the shoot litter versus the root channel, maize litter was applied on wheat cultivated plots, while on half of the maize planted plots no maize litter was returned. Wheat cultivated plots without additional maize litter application served as a reference for the calculation of incorporated maize-C into different soil pools. Soil samplings took place in two consecutive years in summer, autumn and winter. Three depths were considered (0-10 cm: topsoil, 40-50 cm: rooted zone beneath the plough layer, 60-70 cm: unrooted zone). In the third study a microcosm experiment with substrates of different recalcitrance and complexity was carried out to identify primary decomposers of different plant litter materials (leaves and roots) during early stages of decomposition (duration of 32 days) and to follow the C flow into the next higher trophic level (protozoa).
Fate of microbial carbon derived from biogas residues applied to arable soil
(2015) Coban, Halil; Kandeler, Ellen
Soil organic matter (SOM) is the major determinant of soil fertility as it has a number of positive impacts such as improving soil physical parameters, providing nutrients for crops, and supplying energy for the microbial biomass activity in soil. Loss of organic matter is a soil threat observed worldwide. Also, bioenergy crop cultivation may accelerate SOM loss due to higher biomass harvesting compared to food crops. It is necessary to supply adequate organic matter input to arable soils in order to maintain sustainable food and biofuel production. Biogas residues (BGRs), the side-products of biogas production, are rich in microbial and plant biomass; they thus can be used as a soil conditioner and contribute to replenishing the carbon (C) pool in soil. However, our knowledge on the contribution of BGRs particularly the microbial residues present in it to SOM formation is limited, even though scientific interest on SOM formation via microbial inputs is growing. Therefore, the objective of this thesis were i) developing an approach to label microbial biomass of biogas residues, ii) tracing the fate of labelled BGRs in arable soil, iii) determining the C flux within microbial food web, and iv) determining the impacts of other soil conditioners on the mineralization of BGRs. In the first study a method was developed to label the autotrophic microorganisms in a biogas reactor using KH13CO3-amended cow manure as substrate. Analyses of phospholipid fatty acids (PLFA) and ether lipids confirmed the successful labelling of microorganisms, especially Gram-positive bacteria and methanogenic archaea. After removal of unused labelled carbonates by an acid fumigation approach, the labelled BGRs were incubated in soil for 378 days. The fate of 13C was traced in CO2 and in bulk soil with a mass balance having 93% mean recovery. Results showed that about 40% of the C derived from BGRs was rapidly mineralized within the first seven days, and mineralization reached 65% at the end of experiment. The data could be fitted to a two-pool exponential degradation model assuming two C pools each decaying exponentially. The proportions of readily degrading and stable C pools were determined to be 51% and 49%, respectively, with half-lives of 3 days and 1.9 years, respectively. The long half-life of the stable C pool in BGRs may indicate a mid-term contribution to SOM. In addition, the mineralization of SOM was enhanced by BGR-application, i.e. priming effects were detected, thus their extensive application should be avoided. A differential fatty acid approach was used in the second study for the separation of C input from BGRs to living biomass and non-living SOM. Phospholipid fatty acids (PLFA) as indicators of living biomass were compared with total fatty acids (t-FA), which are found also in necromass. Using PLFA as biomarkers of specific microbial groups, C redistribution within the microbial food web was determined. Results showed that BGRs increased the microbial biomass in soil. The sum of 13C-labelled PLFA and t-FA decreased during incubation to 60% and to 33%, respectively. The level of enrichment was different for the individual PLFA and indicated that Gram-negative bacteria were predating on Gram-positive bacteria. A contribution of ether lipids was also detected indicating C flow from decaying methanogens. This study confirmed that microbial biomass in BGRs applied to arable soil significantly contributes to SOM formation. After determining the fate of microbial C derived from BGRs in arable soil, the impacts of other soil conditioners on the mineralization of BGRs were tested in the third study. For this, labelled BGRs were incubated in soil both alone and together with compost, biochar and untreated manure. The amount of C mineralized to CO2 and the degradation rate constant of stable C pool were not affected by any of the co-amendments. However, manure resulted in a higher mineralization rate constant of the readily degrading C pool. C flow within microbial food web was from Gram-positive bacteria and methanogenic archaea to mainly Gram-negative bacteria and slightly to fungi in all treatments. This study showed that co-amending BGRs with other soil conditioners brings neither benefits nor harms in terms of the formation or the mineralization of soil organic matter. The proposed labelling approach using KH13CO3 may be useful for tracing the fate of BGRs. The enrichment in both bacteria and archaea were sufficient to be measured in an incubation experiment lasting for more than one year. However, there are disadvantages of the proposed approach such as presence of highly enriched residual carbonates. The fumigation method should be optimized for a complete removal of the highly labelled residual carbonates which will increase the precision of the overall approach.
Formation of mineral‐associated organic matter in temperate soils is primarily controlled by mineral type and modified by land use and management intensity
(2023) Bramble, De Shorn E.; Ulrich, Susanne; Schöning, Ingo; Mikutta, Robert; Brandt, Luise; Poll, Christian; Kandeler, Ellen; Mikutta, Christian; Konrad, Alexander; Siemens, Jan; Yang, Yang; Polle, Andrea; Schall, Peter; Ammer, Christian; Kaiser, Klaus; Schrumpf, Marion
Formation of mineral-associated organic matter (MAOM) supports the accumulation and stabilization of carbon (C) in soil, and thus, is a key factor in the global C cycle. Little is known about the interplay of mineral type, land use and management intensity in MAOM formation, especially on subdecadal time scales. We exposed mineral containers with goethite or illite, the most abundant iron oxide and phyllosilicate clay in temperate soils, for 5 years in topsoils of 150 forest and 150 grassland sites in three regions across Germany. Results show that irrespective of land use and management intensity, more C accumulated on goethite than illite (on average 0.23 ± 0.10 and 0.06 ± 0.03 mg m−2 mineral surface respectively). Carbon accumulation across regions was consistently higher in coniferous forests than in deciduous forests and grasslands. Structural equation models further showed that thinning and harvesting reduced MAOM formation in forests. Formation of MAOM in grasslands was not affected by grazing. Fertilization had opposite effects on MAOM formation, with the positive effect being mediated by enhanced plant productivity and the negative effect by reduced plant species richness. This highlights the caveat of applying fertilizers as a strategy to increase soil C stocks in temperate grasslands. Overall, we demonstrate that the rate and amount of MAOM formation in soil is primarily driven by mineral type, and can be modulated by land use and management intensity even on subdecadal time scales. Our results suggest that temperate soils dominated by oxides have a higher capacity to accumulate and store C than those dominated by phyllosilicate clays, even under circumneutral pH conditions. Therefore, adopting land use and management practices that increase C inputs into oxide-rich soils that are under their capacity to store C may offer great potential to enhance near-term soil C sequestration.
Function and composition of the soil microbial community in calcareous grassland exposed to elevated atmospheric carbon dioxide
(2003) Ebersberger, Diana; Kandeler, Ellen
Terrestrial ecosystems generally respond to rising atmospheric carbon dioxide (CO2) concentrations with increased net primary productivity and increased water use efficiency. This may change the amount and quality of organic substances entering the soil and fuelling microbial metabolism. Soil microorganisms and their activity might also be affected by increased soil moisture at elevated CO2. This thesis was designed to analyse the response of the soil microbial community in a species-rich calcareous grassland in the Swiss Jura Mountains, which had been exposed to ambient and elevated CO2 concentrations (365 and 600 ppm) for six growing seasons. In the first study, laboratory incubation experiments were conducted to explore the relationship between litter quality under elevated carbon dioxide and enzymes involved in carbon cycling. Naturally senescent, mixed litter from the long-term field experiment was incubated with soil material for 10, 30 and 60 days. Soil samples were then obtained close to the litter layer using a microtome cutting device. Litter and soil samples were analysed for invertase and xylanase activity. The lower litter quality produced under elevated CO2, i.e. wider C/N ratio, yielded lower invertase and xylanase activities of litter. Litter addition stimulated activities in adjacent soil. Invertase activities of adjacent soil were not affected by litter quality, while soil xylanase activity was higher in soil compartments adjacent to litter from elevated CO2 plots. The reduced enzyme activities of litter produced under elevated CO2 can slow decomposition, at least during the initial stages. Since the effects of litter quality on enzyme activities in adjacent soil were small, we conclude that CO2-induced belowground C-inputs (e.g. increased root mass) and altered moisture conditions are more important controls of enzyme activities than altered litter quality. In the second study, functional diversity of the soil microbial community was assessed by analysing N-mineralisation and activities of enzymes of the C-, N-, P- and S-cycle of soil samples taken in spring and summer 1999, in the 6th season of CO2 exposure. In spring, N-mineralisation increased significantly by 30% at elevated CO2, while there was no significant difference between treatments in summer. The response of soil enzymes to CO2 enrichment was also more pronounced in spring, when alkaline phosphatase and urease activities were increased most strongly, by 32% and 21%, respectively. In summer, activity differences between CO2 treatments were greatest in the case of urease and protease (+21% and +17% at elevated CO2). The significant stimulation of N-mineralisation and enzyme activities at elevated CO2 was probably caused by higher soil moisture and/or increased root biomass. In the third study, soil microbial community structure of soil samples taken in spring and summer 1999 was analysed by means of PLFA profiles and 16S rDNA fingerprints obtained by PCR-DGGE. PLFA profiles were not affected by elevated CO2. Ordination analysis of DNA fingerprints revealed a significant relation between CO2 enrichment and variation in DNA fingerprints. This variation must be attributed to low intensity bands because dominant bands did not differ between treatments. Diversity of the bacterial community (number of bands in DNA fingerprints and Shannon indices) was not affected. The observed minute, but significant changes in the structure of the soil bacterial community might be caused by changes in the quality of rhizodeposits at elevated CO2. These could either result from altered rhizodeposition of individual plants or from altered species composition of the calcareous grassland.The 4th part of the thesis compiles data on soil microorganisms, soil fauna, soil structure and nitrogen cycle of calcareous grassland after CO2 exposure for six growing seasons. Microbial biomass, soil basal respiration and the metabolic quotient were not altered significantly. PLFA analysis revealed no significant shift in the ratio of fungi to bacteria. Protozoans, bacterivorous and fungivorous nematodes, acarians, collembolans, and root-feeding nematodes were not affected by elevated CO2. Total nematode numbers averaged slightly lower (-16%) and nematode mass was significantly reduced (by 43%) due to fewer large-diameter nematodes classified as omnivorous and predacious. CO2 exposure resulted in a shift towards smaller aggregate sizes; this was caused by higher soil moisture. Reduced aggregate sizes result in reduced pore neck diameters. This can confine the locomotion of large-diameter nematodes and possibly accounts for their decrease. The CO2 enrichment also affected the nitrogen cycle. N stocks in living plants and surface litter increased, but N in soil organic matter and microorganisms remained unaltered. N mineralisation increased considerably, but microbial N did not differ between treatments, indicating that net N immobilization rates were unaltered.
Hidden miners – the roles of cover crops and soil microorganisms in phosphorus cycling through agroecosystems
(2019) Hallama, Moritz; Pekrun, Carola; Lambers, Hans; Kandeler, Ellen
Background Phosphorus (P) is a limiting nutrient in many agroecosystems and costly fertilizer inputs can cause negative environmental impacts. Cover crops constitute a promising management option for sustainable intensification of agriculture. However, their interactions with the soil microbial community, which is a key driver of P cycling, and their effects on the following crop, have not yet been systematically assessed. Scope We conducted a meta-analysis of published field studies on cover crops and P cycling, focusing on plant-microbe interactions. Conclusions We describe several distinct, simultaneous mechanisms of P benefits for the main crop. Decomposition dynamics, governed by P concentration, are critical for the transfer of P from cover crop residues to the main crop. Cover crops may enhance the soil microbial community by providing a legacy of increased mycorrhizal abundance, microbial biomass P, and phosphatase activity. Cover crops are generally most effective in systems low in available P, and may access ‘unavailable’ P pools. However, their effects on P availability are difficult to detect by standard soil P tests, except for increases after the use of Lupinus sp. Agricultural management (i.e. cover crop species selection, tillage, fertilization) can improve cover crop effects. In summary, cover cropping has the potential to tighten nutrient cycling in agricultural systems under different conditions, increasing crop P nutrition and yield.
Increasing plant species richness by seeding has marginal effects on ecosystem functioning in agricultural grasslands
(2023) Freitag, Martin; Hölzel, Norbert; Neuenkamp, Lena; van der Plas, Fons; Manning, Peter; Abrahão, Anna; Bergmann, Joana; Boeddinghaus, Runa; Bolliger, Ralph; Hamer, Ute; Kandeler, Ellen; Kleinebecker, Till; Knorr, Klaus‐Holger; Marhan, Sven; Neyret, Margot; Prati, Daniel; Le Provost, Gaëtane; Saiz, Hugo; van Kleunen, Mark; Schäfer, Deborah; Klaus, Valentin H.
Experimental evidence shows that grassland plant diversity enhances ecosystem functioning. Yet, the transfer of results from controlled biodiversity experiments to naturally assembled ‘real world’ ecosystems remains challenging due to environmental variation among sites, confounding biodiversity ecosystem functioning relations in observational studies. To bridge the gap between classical biodiversity‐ecosystem functioning experiments and observational studies of naturally assembled and managed ecosystems, we created regionally replicated, within‐site gradients of species richness by seeding across agricultural grasslands differing in land‐use intensity (LUI) and abiotic site conditions. Within each of 73 grassland sites, we established a full‐factorial experiment with high‐diversity seeding and topsoil disturbance and measured 12 ecosystem functions related to productivity, and carbon and nutrient cycling after 4 years. We then analysed the effects of plant diversity (seeded richness as well as realized richness), functional community composition, land use and abiotic conditions on the ecosystem functions within (local scale) as well as among grassland sites (landscape scale). Despite the successful creation of a within‐site gradient in plant diversity (average increase in species richness in seeding treatments by 10%–35%), we found that only one to two of the 12 ecosystem functions responded to realized species richness, resulting in more closed nitrogen cycles in more diverse plant communities. Similar results were found when analysing the effect of the seeding treatment instead of realized species richness. Among sites, ecosystem functioning was mostly driven by environmental conditions and LUI. Also here, the only functions related to plant species richness were those associated with a more closed nitrogen cycle under increased diversity. The minor effects of species enrichment we found suggest that the functionally‐relevant niche space is largely saturated in naturally assembled grasslands, and that competitive, high‐functioning species are already present. Synthesis: While nature conservation and cultural ecosystem services can certainly benefit from plant species enrichment, our study indicates that restoration of plant diversity in naturally assembled communities may deliver only relatively weak increases in ecosystem functioning, such as a more closed nitrogen cycle, within the extensively to moderate intensively managed agricultural grasslands of our study.
Influence of land use on abundance, function and spatial distribution of N-cycling microorganisms in grassland soils
(2015) Keil, Daniel; Kandeler, Ellen
This thesis focuses on the influence of land use on the abundance, function and spatial distribution of N-cycling microorganisms in grassland soils, but also on soil biogeochemical properties, as well as on enzyme activities involved in the carbon-, nitrogen-, and phosphorous cycle. The objective of this thesis was tackled in three studies. All study sites that were investigated as part of this thesis were preselected and assigned according to study region and land use within the framework of the “Exploratories for Functional Biodiversity Research – The Biodiversity Exploratories” of the Deutsche Forschungsgemeinschaft priority program 1374. The first study addressed the question whether land-use intensity influences soil biogeochemical properties, as well as the abundance and spatial distributions of ammonia-oxidizing and denitrifying microorganisms in grasslands of the Schwäbische Alb. To this end, a geostatistical approach on replicated grassland sites (10 m × 10 m), belonging to either unfertilized pastures (n = 3) or fertilized mown meadows (n = 3), representing low and high land-use intensity, was applied. Results of this study revealed that land-use intensity changed spatial patterns of both soil biogeochemical properties and N-cycling microorganisms at the plot scale. For soil biogeochemical properties, spatial heterogeneity decreased with higher land-use intensity, but increased for ammonia oxidizers and nirS-type denitrifiers. This suggests that other factors, both biotic and abiotic than those measured, are driving the spatial distribution of these microorganisms at the plot scale. Furterhmore, the geostatistical analysis indicated spatial coexistence for ammonia oxidizers (amoA ammonia-oxidizing archaea and amoA ammonia-oxidizing bacteria) and nitrate reducers (napA and narG), but niche partitioning between nirK- and nirS-type denitrifiers. The second study aimed at whether land-use intensity contributes to spatial variation in microbial abundance and function in grassland ecosystems of the Schwäbische Alb assigned to either low (unfertilized pastures, n = 3), intermediate (fertilized mown pastures, n = 3), or high (fertilized mown meadows, n = 3) land-use intensity. Plot-scale (10 m × 10 m) spatial heterogeneity and autocorrelation of soil biogeochemical properties, microbial biomass and enzymes involved in C, N, and P cycle were investigated using a geostatistical approach. Geostatistics revealed spatial autocorrelations (p-Range) of chemical soil properties within the maximum sampling distance of the investigated plots, while greater variations of p-Ranges of soil microbiological properties indicated spatial heterogeneity at multiple scales. An expected decrease in small-scale spatial heterogeneity in high land-use intensity could not be confirmed for microbiological soil properties. Finding smaller spatial autocorrelations for most of the investigated properties indicated increased habitat heterogeneity at smaller scales under high land-use intensity. In the third study, the effects of warming and drought on the abundance of denitrifier marker genes, the potential denitrification activity and the N2O emission potential from grassland ecosystems located in the Schwäbische Alb, the Hainich, and the Schorfheide region were investigated. Land use was defined individually for each grassland site by a land-use index that integrated mowing, grazing and fertilization at the sites over the last three years before sampling of the soil. It was tested if the microbial community response to warming and drought depended on more static site properties (soil organic carbon, water holding capacity, pH) in interaction with land use, the study region and the climate change treatment. It was further tested to which extent the N2O emission potential was influenced by more dynamic properties, e.g. the actual water content, the availability of organic carbon and nitrate, or the size of the denitrifier community. Warming effects in enhanced the potential denitrification of denitrifying microorganisms. While differences among the study regions were mainly related to soil chemical and physical properties, the land-use index was a stronger driver for potential denitrification, and grasslands with higher land use also had greater potentials for N2O emissions. The total bacterial community did not respond to experimental treatments, displaying resilience to minor and short-term effects of climate change. In contrast, the denitrifier community tended to be influenced by the experimental treatments and particularly the nosZ abundance was influenced by drought. The results indicate that warming and drought affected the denitrifying communities and the potential denitrification, but these effects are overruled by study region and site-specific land-use index. This thesis gives novel insights into the performance of N-cycling microorganisms in grassland ecosystems. The spatial distribution of soil biogeochemical properties is strongly dependent on land-use intensity, as in return is the spatial distribution of nitrifying and denitrifying microorganisms and the ecosystem services they perform. Yet, future work will be necessary to fully understand the interrelating factors and seasonal variability, which influence the ecosystem functioning and ecosystem services that are provided by N-cycling soil microorganisms at multiple scales.
Linking microbial abundance and function to understand nitrogen cycling in grassland soils
(2017) Regan, Kathleen Marie; Kandeler, Ellen
This thesis characterized spatial and temporal relationships of the soil microbial community, the nitrogen cycling microbial community, and a subset of the nitrogen cycling community with soil abiotic properties and plant growth stages in an unfertilized temperate grassland. Unfertilized perennial grasslands depend solely on soil-available nitrogen and in these environments nitrogen cycling is considered to be both highly efficient and tightly coupled to plant growth. Unfertilized perennial grasslands with high plant diversity, such as ours, have also been shown to have higher soil organic carbon, total nitrogen, and microbial carbon; greater food web complexity; and more complex biological communities than more intensively managed grasslands or croplands. This made the choice of study plot especially well-suited for characterizing the relationships we sought to identify, and made it possible to detect spatial and temporal patterns at a scale that has heretofore been under-examined. The first study used a combination of abiotic, plant functional group, and PLFA measurements together with spatial statistics to interpret spatial and temporal changes in the microbial community over a season. We found that its overall structure was strongly related to the abiotic environment throughout the sampling period. The strength of that relationship varied, however, indicating that it was not constant over time and that other factors also influenced microbial community composition. PLFA analysis combined with principal components analysis made it possible to discern changes in abundances and spatial distributions among Gram-positive and Gram-negative bacteria as well as saprotrophic fungi. Modeled variograms and kriged maps of the changes in distributions of exemplary lipids of both bacterial groups also showed distinct differences in their distributions on the plot, especially at stages of most rapid plant growth. Although environmental properties were identified as the main structuring agents of the microbial community, components of those environmental properties varied over the season, suggesting that plant growth stage had an indirect influence, providing evidence of the complexity and dynamic nature of the microbial community in a grassland soil. The second study took the same analytical approach, this time applying it to abundances of key members of the soil nitrogen cycling community. Marker genes for total archaea and bacteria, nitrogen fixing bacteria, ammonia oxidizing archaea and bacteria, and denitrifying bacteria were quantified by qPCR. Potential nitrification activity and denitrifying enzyme activity were also determined. We found clear seasonal changes in the patterns of abundance of the measured genes and could associate these with changes in substrate availability related to plant growth stages. Most strikingly, we saw that small and ephemeral changes in soil environmental conditions resulted in changes in these microbial communities, while at the same time, process rates of their respective potential enzyme activities remained relatively stable. This suggests both short term niche-partitioning and functional redundancy within the nitrogen cycling microbial community. The seasonal changes in abundances we observed also provided additional evidence of a dynamic relationship between microorganisms and plants, an important mechanism controlling ecosystem nitrogen cycling. The third study determined spatial and temporal interactions between AOA, AOB and NOB. These steps are related in both space and time, as the ammonia-oxidizers provide the necessary substrate for nitrite-oxidizers. Using a combination of spatial statistics and phylogenetic analysis, our data indicated seasonally varying patterns of niche differentiation between the two bacterial groups, Nitrospira and Nitrobacter in April, but more homogeneous patterns by August which may have been due to different strategies for adapting to changes in substrate concentrations resulting from competition with plants. We then asked a further question: was the microbial structure at sampling sites with high NS gene abundances fundamentally different from those with low NS gene abundances? Using a phylogenetic approach, the operational taxonomic unit composition of NS was analyzed. Community composition did not change over the first half of the season, but by the second half, the relative proportion of a particular OTU had increased significantly. This suggested an intraspecific competition within the NS and the possible importance of OTU 03 in nitrite oxidation at a specific period of time. Observed positive correlations between AOA and Nitrospira further suggested that in this unfertilized grassland plot, the nitrification process may be predominantly performed by these groups, but is restricted to a limited timeframe.
Microbial community structure and function is shaped by microhabitat characteristics in soil
(2016) Ditterich, Franziska; Kandeler, Ellen
Soil microorganisms play a key role in degradation processes in soil, such as organic matter decomposition and degradation of xenobiotics. Microbial growth and activity and therefore degradation processes are influenced by different ecological factors, such as substrate availability, pH and temperature. During soil development different microhabitats are formed which differ in their physiochemical properties. There is some evidence that mineral composition is a driver for specific microbial colonization. Thereby, the heterogeneity of soils with differences in mineral composition and substrate availability can lead to a spatial distribution of soil microorganisms. At the soil-litter interface, a biogeochemical hot spot in soil, the abundance and activity of soil microorganisms increases due to high substrate availability, and degradation processes such as pesticide degradation are enhanced. This thesis aimed to clarify the influence of habitat properties on the structure and function of the microbial community in soil. In particular, focus was on mineral-microbe interactions that result from the mineral composition and substrate availability in an artificial soils system. Furthermore this thesis was designed to increase our understanding of the bacterial and fungal roles in pesticide degradation at the soil-litter interface using 4-chloro-2-methylphenoxyacetic acid (MCPA) as a model xenobiotic. These two aspects of the thesis were examined in three studies. The first study focused on the succession of microbial communities and enzyme activities in an artificial soils system with varying mineral composition and substrate availability over a period of 18 months. In the second study a microcosm experiment was used to study the bacterial pathway of MCPA degradation at the soil-litter interface. Over a period of 27 days the succession of bacterial degraders was followed. The third study focused on the degradation of MCPA in soil by nonspecific fungal enzymes, through the addition of fungal laccases as well as litter during 42 days of incubation. Both studies indicated the involvement of fungi in MCPA degradation and the importance of the ecological behavior of different degraders as a function of substrate availability. Results of the first study indicated that the microbial community was affected by mineral properties under high substrate availability and by the availability of beneficial nutrients at the end of incubation when substrate had become limited. The measured enzyme activities provided clear evidence that microbial community structure was driven by nutrient limitation during incubation. In the presence of easily available organic substrates at the beginning of the experiment, the soil microbial community was dominated by copiotrophic bacteria (e.g. Betaproteobacteria), whereas under substrate limitation at the end of incubation, more recalcitrant compounds became important to oligotrophic bacteria (e.g. Acidobacteria), which then became dominant. The results of the second study indicated that the contribution of the potential degraders to degradation of MCPA differed, and this was also seen in the succession of specific bacterial MCPA degraders. Added litter stimulated MCPA degradation due to the availability of litter-derived carbon and induced a two-phase response of fungi. This was seen in the development of pioneer and late stage fungal communities. Both fungal communities were probably involved in MCPA degradation. Therefore, the third study focused on the fungal pathway. These results indicated that the fungal laccases used had no direct influence on degradation and were as efficient as litter in providing additional nutrient sources, increasing MCPA degradation by bacteria and fungi. The observed differences between litter and enzyme addition underscored the observation that the enzyme effect was short-lived and that substrate quality is an important factor in degradation processes. In conclusion, this thesis demonstrated that soil microbial communities and therefore degradation processes are driven by mineral composition as well as substrate availability and quality. In addition, this thesis extends our understanding of degradation processes such as the degradation of xenobiotics, with MCPA as model compound, in soil. The combined insights from all three studies suggest that the use of a simple system such as the artificial soil system can increase our understanding of complex mechanisms such as degradation of pesticides.
Microbial drivers of plant richness and productivity in a grassland restoration experiment along a gradient of land‐use intensity
(2022) Abrahão, Anna; Marhan, Sven; Boeddinghaus, Runa S.; Nawaz, Ali; Wubet, Tesfaye; Hölzel, Norbert; Klaus, Valentin H.; Kleinebecker, Till; Freitag, Martin; Hamer, Ute; Oliveira, Rafael S.; Lambers, Hans; Kandeler, Ellen
Plant–soil feedbacks (PSFs) underlying grassland plant richness and productivity are typically coupled with nutrient availability; however, we lack understanding of how restoration measures to increase plant diversity might affect PSFs. We examined the roles of sward disturbance, seed addition and land‐use intensity (LUI) on PSFs. We conducted a disturbance and seed addition experiment in 10 grasslands along a LUI gradient and characterized plant biomass and richness, soil microbial biomass, community composition and enzyme activities. Greater plant biomass at high LUI was related to a decrease in the fungal to bacterial ratios, indicating highly productive grasslands to be dominated by bacteria. Lower enzyme activity per microbial biomass at high plant species richness indicated a slower carbon (C) cycling. The relative abundance of fungal saprotrophs decreased, while pathogens increased with LUI and disturbance. Both fungal guilds were negatively associated with plant richness, indicating the mechanisms underlying PSFs depended on LUI. We show that LUI and disturbance affect fungal functional composition, which may feedback on plant species richness by impeding the establishment of pathogen‐sensitive species. Therefore, we highlight the need to integrate LUI including its effects on PSFs when planning for practices that aim to optimize plant diversity and productivity.
Microbial regulation of soil organic matter decomposition at the regional scale
(2018) Ali, Rana Shahbaz; Kandeler, Ellen
The fate of soil organic carbon (SOC) is one of the greatest uncertainties in predicting future climate. Soil microorganisms, as primary decomposers of SOC, control C storage in terrestrial ecosystems by mediating feedbacks to climate change. Even small changes in microbial SOC decomposition rates at the regional scale have the potential to alter land-atmospheric feedbacks at the global scale. Despite their critical role, the ways in which soil microorganisms may change their abundances and functions in response to the climate change drivers of soil temperature and moisture is unclear. Additionally, most existing C models do not consider soil microorganisms explicitly as drivers of decomposition, one consequence of which is large variability in predicted SOC stock projections. This demonstrates the need for a better mechanistic understanding of microbial SOC decomposition at large scales. This thesis was designed to clarify the role of microbial SOC decomposition dynamics in response to climate change factors in two geographically distinct areas and land-use types. The hypothesis was that microbial communities would be adapted to climatic and edaphic conditions specific to each area and to the SOC organic quality in each land-use and would therefore exhibit distinct responses to soil temperature and moisture variations. Three studies were performed to address the goals of this thesis. The first study aimed to clarify temporal patterns of degradation in C pools that varied in complexity by modelling in situ potentials of microbially produced extracellular enzymes. Temperature and moisture sensitivity patterns of C cycling enzymes were followed over a period of thirteen months. The second study investigated group-specific temperature responses of bacteria and fungi to substrate quality variations through an additional incubation experiment. Here, complex environments were mimicked in order to determine the dependence of microbial responses not only on environmental conditions, but also under conditions of inter- and intra-specific community competition. Changes in microbial community composition, abundance, and function were determined at coarse (phospholipid fatty acid – PLFA, ergosterol) and relatively fine resolutions (16S rRNA, taxa-specific quantitative PCR, fungal ITS fragment). A third study investigated 1) the spatial variability of temperature sensitivity of microbial processes, and 2) the scale-specificity and relative significance of their biotic and physicochemical controls at landscape (two individual areas, each ca. 27 km2) and regional scales (pooled data of two areas). Strong seasonal dependency was observed in the temperature sensitivities (Q10) of hydrolytic and oxidative enzymes, whereas moisture sensitivity of β-glucosidase activities remained stable over the year. The range of measured enzyme Q10 values was similar irrespective of spatial scale, indicating a consistency of temperature sensitivities of these enzymes at large scales. Enzymes catalyzing the recalcitrant SOC pool exhibited higher temperature sensitivities than enzymes catalyzing the labile pool; because the recalcitrant C pool is relatively large, this could be important for understanding SOC sensitivity to predicted global warming. Response functions were used to model temperature-based and temperature and moisture-based in situ enzyme potentials to characterize seasonal variations in SOC decomposition. In situ enzyme potential explained measured soil respiration fluxes more efficiently than the commonly used temperature-respiration function, supporting the validity of our chosen modelling approach. As shown in the incubation experiment, increasing temperature stimulated respiration but decreased the total biomass of bacteria and fungi irrespective of substrate complexity, indicating strong stress responses by both over short time scales. This response did not differ between study areas and land-uses, indicating a dominant role of temperature and substrate quality in controlling microbial SOC decomposition. Temperature strongly influenced the responses of microbial groups exhibiting different life strategies under varying substrate quality availability; with soil warming, the abundance of oligotrophs (fungi and gram-positive bacteria) decreased, whereas copiotrophs (gram-negative) increased under labile C substrate conditions. Such an interactive effect of soil temperature and substrate quality was also visible at the taxon level, where copiotrophic bacteria were associated with labile C substrates and oligotrophic bacteria with recalcitrant substrates. Which physicochemical and biological factors might explain the observed alterations in microbial communities and their functions in response to climate change drivers at the regional scale was investigated in the third study. Here, it was shown that the soil C:N ratio exerted scale-dependent control over soil basal respiration, whereas microbial biomass explained soil basal respiration independent of spatial scale. Factors explaining the temperature sensitivity of soil respiration also differed by spatial scale; extractable organic C and soil pH were important only at the landscape scale, whereas soil texture as a control was independent of spatial scale. In conclusion, this thesis provides an enhanced understanding of the response of microbial C dynamics to climate change at large scales by combining field measurements with innovative laboratory assays and modelling tools. Component specific degradation rates of SOC using extracellular enzyme measurements as a proxy, group-specific temperature sensitivities of microbial key players, and the demonstrated scale-specificity of factors controlling microbial processes could potentially improve the predictive power of currently available C models at regional scale.
Microplastics interactions with soil organisms
(2022) Schöpfer, Lion; Kandeler, Ellen
Microplastics (MP) are plastic particles from 100 nm to 5 mm with different shapes and chemical compositions. In aquatic ecosystems, MP have proven to affect the biological fitness of aquatic organisms, enter the food web, and act as vectors of pollutants. Agricultural soils are sinks for MP due to inputs via sewage sludges, plastic mulches, and organic fertilizers. However, ecological consequences of MP in agricultural soils are unknown. This doctoral thesis aimed to evaluate the risk of conventional and biodegradable MP for soil organisms in agricultural soils. A microcosm study was combined with a field study and a nematode study to investigate background concentrations, the persistence, and the biodegradation of MP in the soil, and effects of MP on soil microorganisms and nematodes. In the microcosm study, the influence of plastic type, particle size, and soil moisture on the biodegradation of MP in the soil and on effects on soil microorganisms were examined under controlled conditions (25 °C, 230 days). The abundance and composition of the main soil microbial groups was analyzed via phospholipid fatty acids (PLFAs) as biomarkers; activities of C cycling enzymes driving the decomposition of differently complex substances were analyzed as proxies for C turnover. To understand better the role of MP as an interface for specific microbial processes in the soil, e.g. the enzymatic hydrolysis of MP, enzyme activities of individual MP particles extracted from the soil were measured. The site of the field study was a conventionally managed agricultural soil (silt-loam Luvisol) of the Heidfeldhof, University of Hohenheim. No practices associated with significant inputs of MP have been conducted at the site in the past (sewage sludge, organic fertilizers, plastic mulch). In a randomized complete block design, the effects of MP, organic fertilizers (digestate and compost), and their interactions on soil microbiological indicators (microbial biomass, soil enzymes) were studied. Before the setup of the field study, MP background concentrations (particle-based) in the soil were analyzed. The persistence of added MP in the soil was evaluated by comparing MP concentrations in the soil after 1 month and 17 months with initial MP concentrations after addition. In the nematode study, the soil-dwelling nematode Caenorhabditis elegans was exposed to MP feed suspensions on agar plates. The uptake of MP through nematodes and the influence of plastic type and concentration on MP effects on nematode reproduction and body length were examined. In all studies, artificially fragmented MP from a conventional polymer (low-density polyethylene, LDPE) and a biodegradable polymer blend (poly(lactic acid) and poly(butylene adipate-co-terephtalate), PLA/PBAT) were used. The occurrence of both LDPE- and PLA/PBAT-MP is likely in agricultural soils because these are used for plastic mulches and compost bags. Results from this thesis suggest that (1) agricultural soils, including those without management practices related to significant MP entry, contain various MP, indicating diffuse MP inputs via atmospheric deposition, littering, and the abrasion of machinery coatings (a possible newly identified pathway), (2) also biodegradable MP persist and are slowly biodegraded in the soil implying a long term exposure risk for soil organisms to MP, (3) MP have no acute negative effects on microorganisms and C turnover, (4) MP form a specific habitat in the soil, the plastisphere, where MP-specific processes take place, e.g. the enzymatic hydrolysis of PLA/PBAT, (5) MP can enter the soil food web via nematodal uptake and affect nematode reproduction, which could destabilize the soil food web.