Soil is an important ecosystem that supports a wide variety of organisms such as bacteria, fungi, arthropods and nematodes. This sensitive ecosystem may be influenced by various factors, including agricultural management practices. With the introduction of genetically modified (GM) glyphosate-tolerant (RoundUp ® Ready: RR) crops, herbicides such as glyphosate have been increasingly used. However, little is known about the effect of glyphosate on the biological communities in these herbicide-sprayed soils. With the intimate proximity that microorganisms and nematodes have with the roots of plants, these organisms can be used to assess changes that may occur in the soil surrounding roots of RR crops. The aim of this study was to determine microbial community structure and nematode diversity, with emphasis on that of non-parasitic nematodes, in soil samples from conventional soybean (CS) - and RR- soybean fields compared to that in adjacent natural veld (NV) areas. Samples were collected from twenty three sites at six localities that are situated within the soybean-production areas of South Africa. These sites represented fields where RR and CS soybean grew, as well as surrounding NV. All RR fields have been treated with glyphosate for no less than five years. Microbial community structures of the twenty three sites in the RR, CS and NV ecosystems were determined by phospholipid fatty acid (PLFA) analyses. Nematode diversity was determined by extracting the nematodes from soil samples and conducting a faunal analysis. Soil physical and chemical properties were determined by an independent laboratory, Eco-Analytica (North West University, Potchefstroom) according to standard procedures. Results from this study indicated differences in microbial community structure between the various localities. However, there were no significant (p ≤ 0.05) differences in microbial community structures between RR- and CS ecosystems. Soils of both RR- and CS crops were primarily dominated by bacteria. Nematode identification and faunal analysis also indicated no significant (p ≤ 0.05) differences between the different non-parasitic/beneficial nematodes that were present in soils of these two ecosystems during the time of sampling. Non-parasitic nematode communities were primarily dominated by bacterivores. A faunal analysis indicated that most of the sites contained enriched, but unstructured soil food-webs. However, four of the sites showed enriched and structured food webs due to the presence of non-parasitic nematodes with high coloniser-persister (cp) values. Relationships between non-parasitic nematode – and microbial communities showed that there was a positive relationship between nematode functional groups and their corresponding microbial prey. From the results obtained in this study, it can be concluded that the community structures of both non-parasitic nematodes and microorganisms shared similarities. These community structures showed no long-term detrimental effects of glyphosate application in the soils surrounding roots of RR soybean crops. Relationships existed between non-parasitic nematode and microbial communities in the rhizosphere of soybean crops and natural veld. For example, bacterivore nematodes had a strong positive relationship with gram-negative bacteria. Similar but weaker relationships also existed between carnivores, omnivores, plantparasitic nematodes and gram-negative bacteria. A positive relationship also existed between fungivores and fungal fatty acids. This emphasises the value of these organisms as indicators of soil health and also the impact that agricultural practices can have on soils. / MSc (Environmental Sciences), North-West University, Potchefstroom Campus, 2014

Global change phenomena, such as forest disturbance and land-use change significantly affect elemental balances as well as the structure and function of terrestrial ecosystems. Inappropriate land management often causes nutrient losses and finally soil degradation and loss of soil functioning. Especially in tropical ecoregions, soil degradation by nutrient losses is widely abundant. Soil microorganisms are the proximate agents of many processes performed in soils and are regarded as sensitive bio-indicators. However, the incorporation of microbial responses to the definition of critical soil conditions is not intensively developed. In the present thesis, several data analyses of the relationships between ecosystem disturbance and land-use change (natural forest, pastures of different ages, secondary succession) and a diverse set of soil ecological characteristics in the tropical mountain rainforest region of southern Ecuador were compiled. In particular, it was tested whether soil microbial biomass and community functioning were sensitive to land-use change effects. Furthermore, an information-theoretic approach was applied to find the factors that regulate soil microbial biomass and community function. Finally, in a nutrient enrichment experiment the above- and belowground responses to N and P additions were examined. The tested research questions and results were linked to the theory of ecological stoichiometry in order to connect the research to a sound and unifying scientific basis. Soil and microbial stoichiometry were affected by both land-use change and soil depth. After forest disturbance, significant decreases of soil C:N:P ratios at the pastures were fol-lowed by increases during secondary succession. Microbial C:N ratios varied slightly in response to land-use change, whereas no fixed microbial C:P and N:P ratios were observed. Shifts in microbial community composition were associated with soil and microbial stoichiometry. Strong positive relationships between PLFA-markers 18:2n6,9c (saprotrophic fungi) and 20:4 (animals) and negative associations between 20:4 and microbial N:P point to land-use change affecting the structure of soil food webs. Significant deviations from global soil and microbial C:N:P ratios indicated a major force of land-use change to alter stoichiometric relationships and to structure biological systems. Data analysis reveals a strong impact of land-use change on soil microbial biomass, C-mineralization, gross-NH4-consumption and –production rates. According to the results of the IT-approach, combined models better describe effects of land-use change on soil microorganisms than single explanation models. Microbial resources and soil chemical environment were important pre-dictors for soil microbial biomass and community functioning. Little is known about the environmental drivers of the catalytic properties of EHEs (e.g., pH, nutrients) and their functional link to the structure of soil microbial communities. The activities of the six hydrolytic enzymes were tested. Microbial production of AP responded to the low P status of the sites by a higher investment in the acquisition of P compared to C. Three major drivers of enzyme activities were found to be significant for enzyme production: 1.) Microbial demand for P regulated the production of AP, provided that N and C were available. At the natural forest site the two-fold higher specific activity of AP pointed to a high microbial P-demand, whereas the production of AP was constrained by the availability of N and DOC after pasture abandonment. 2.) Microbial biomass that was controlled by pH and resource availability was the main driver for CBH, BG and NAG activities. 3.) Substrate induction due to increased litter inputs of herbaceous plant species seemed to regulate AG and XYL activities during secondary succession. The enzymes’ affinity to substrate, as a potentially critically enzyme kinetic parameter is understudied. The data analysis suggests that microbial communities adapted to environmental changes, demonstrated high flexibility of extracellular enzyme systems and selected for enzymes with higher catalytic efficiency compared with pure cultures. Under in situ conditions, enzyme-specific environmental drivers of the Km, e.g., the pH for XYL, the C:N ratio for AP, and the C availability for NAG were found. The data demonstrated that the higher substrate affinity of XYL and AP was associated with more abundance of Gram(-) bacteria. The catalytic efficiency of enzymes decomposing cellulose, hemicellulose, and starch positively correlated with the relative abundance of Gram(-) bacteria. The turnover rate of the tested substrates was three to four times faster at the young pasture site compared with the longterm pasture and secondary succession sites. Nutrient inputs by atmospheric deposition are known to affect terrestrial ecosystems. However, little is known about how N and P co-limited ecosystems respond to single nutrient enrichment. In this work the susceptibility of above- and belowground ecosystem compo-nents and of their linkages in an N and P co-limited pasture to N- and P-enrichment was assessed. It was tested if the plants´responses can be explained by the concept of serially linked nutrients introduced by Ågren (2004). In this concept, the control of the growth rate by one nutrient is assumed to depend on the control of a different cellular process by another nutrient. The responses of shoot and root biomass and C:N:P stoichiometry of the grass Setaria sphacelata (Schumach.) to moderate N, P, and N+P application over five years were investigated. In addition, the effects of nutrient enrichment on soil nutrient pools, on arbuscular mycorrhizal fungi (AMF) as well as on microbial biomass, activity, and community structure were tested. In order to evaluate the importance of different factors explaining microbial responses, a likelihood-based information-theoretic approach was applied. The application of N+P increased aboveground grass biomass. Root biomass was stimulated by P-treatment. Grass C:N:P stoichiometry responded by altering the P-uptake or by translocating P from shoot to root. In particular, root C:N and C:P stoichiometry decreased in P- and in N-treatment. Extractable fractions of soil C, N, and P were significantly affected by nutrient enrichment. P application increased the biomass of Gram-positive bacteria and the abundance of AMF, however, results of the IT-approach suggested indirect effects of nutrient enrichment on microbes. The responses of the N and P co-limited pasture to particular nutrient enrichment support the concept of serially linked nutrients. The present study provides evidence for the fundamental importance of P for controlling resource allocation of plants in responses to nutrient enrichment. Resource allocation of the grass rather than direct effects of nutrient additions drives changes in AMF, microbial biomass, community structure, and activity. / Seit dem Übergang vom Holozän zum Anthropozän greift der Mensch immer stärker in globale und regionale Stoffkreisläufe ein. Durch die Zerstörung von Naturwäldern und Landnutzungswandel werden die Strukturen und die Funktionen der Ökosysteme stark verändert. Unangepasste Landnutzung führt zu Nährelementverlusten, die mittel- bis langfristige zur Bodendegradation und zur Reduktion von Bodenfunktionen führen. Solche Veränderungen sind insbesondere in den Tropen zu beobachten. Bodenmikroorganismen spielen in den Stoffkreisläufen eine zentrale Rolle. Zudem sind sie sensitive Bioindikatoren für den Zustand von Ökosystemen. Im Gegensatz dazu, werden die Bodenmikroorganismen noch nicht ausreichend für die Zustandsbewertung von Ökosystemen verwendet. In der vorliegenden Dissertation werden verschiedene Datenanalysen zu den Beziehungen von Landnutzungswandel (Naturwald, Weiden verschiedener Alter, sekundäre Sukzession) und den Eigenschaften der Bodenmikroorganismen in einer tropischen Bergregenwaldregion Süd-Ecuadors zusammengefasst. Ein besonderer Fokus lag darauf zu prüfen, ob die mikrobielle Biomasse und die Funktionen die von der mikrobiellen Gemeinschaft geleistet werden (z.B. Enzymaktivitäten) durch den Landnutzungswandel beeinflusst werden. Ein informations-theoretischer Ansatz wurde verwendet um verschiedene Erklärungsansätze der steuernden Faktoren vergleichend zu testen. Darüber hinaus wurden in einem Weidedüngungsexperiment die Reaktionen der ober- und der unterirdischen Ökosystemkomponenten auf die Anreicherung mit N und P getestet. Um die Ergebnisse auf eine breite wissenschaftliche Basis zu stellen wurde die Untersuchungen in den Kontext der Theorie die Ökologischen Stöchiometrie eingeordnet. Die C:N:P Stöchiometrie im Boden und in den Mikroorganismen veränderte sich durch den Landnutzungswandel und mit der Bodentiefe. Mit der Weideetablierung nahmen die C:N:P Verhältnisse im Boden deutlich ab, stiegen dann nach dem Verlassen der Weiden im Zuge der sekundären Sukzession wieder an. Das mikrobielle C:N Verhältnis variierte nur leicht, dagegen zeigten das C:P und N:P Verhältnis deutliche Veränderungen durch den Landnutzungswandel. Mit diesen Veränderungen in der Boden- und Organismenstöchiometrie waren auch Veränderungen in der Struktur der mikrobiellen Gemeinschaften verbunden. Deutliche positive Beziehungen existierten zwischen den saprotrophen Pilzen und den Protozoen. Die steigenden Mengen von Protozoen waren wiederrum mit sinkendem mikrobiellen N:P verbunden. Diese Muster weisen auf Veränderungen in den Bodennahrungsnetzten durch Landnutzungsänderungen hin. Sehr deutliche Abweichungen von globalen Mustern der C:N:P Stöchiometrie deuten darauf hin, dass der Landnutzungswandel signifikanten Einfluss auf die C:N:P Stöchiometrie ausübt. Der Landnutzungswandel beeinflusste auch die mikrobielle Biomasse, die Basalatmung, sowie die mikrobielle Aufnahme und Produktion von NH4-N im Boden. Dabei zeigten kombinierte Erklärungsansätze die adäquateren Beschreibungen der Muster. In den kombinierten Modellen zur Erklärung der mikrobiellen Biomasse und der mikrobiellen Leistungen überwogen Prädiktoren der mikrobiellen Ressourcen und der bodenchemischen Umwelt. Ein weiterer Schwerpunkt der Untersuchungen lag auf der Erfassung der Effekte des Land-nutzungswandels auf die Aktivität von extrazellulären Bodenenzymen. Bisher ist wenig darüber bekannt, welche Faktoren die katalytischen Eigenschaften steuern und beispielsweise, ob es Zusammenhänge zur mikrobiellen Gemeinschaftsstruktur gibt. Um diese Fragen näher zu beleuchten wurden sechs hydrolytische Enzyme basierend auf MUF-Substraten untersucht. Die mikrobielle Produktion von AP stand dabei in Zusammenhang mit dem niedrigen P-Status der untersuchten Böden. Das wurde besonders durch die hohe AP Produktion im Vergleich zu BG belegt. Im Allgemeinen konnten drei verschiedene Mechanismen festgestellt werden, die die Produktion der untersuchten EHEs vermutlich steuerten. 1.) Der P-Bedarf der Mikroorganismen regulierte die Produktion von AP, vorausgesetzt, dass ausreichend N und C zur Enzymsynthese zur Verfügung standen. 2.) Die Höhe der mikrobiellen Biomasse hat sich als wichtiger Faktor für die Produktion von CBH, BG und NAG gezeigt. Das deutet auf die konstitutive Produktion dieser Enzyme hin. 3.) Die substratinduzierte Produktion von Enzymen ist vermutlich entscheidend für die Aktivität von AG und XYL. Die Berücksichtigung der Enzymkinetiken, insbesondere der Michaelis-Menten-Konstante lieferte weitere Aufschlüsse über relevante Faktoren. Im Allgemeinen so scheint es, haben sich die mikrobiellen Gemeinschaften an die starken Umweltgradienten, die durch den Landnutzungswandel erzeugt worden angepasst. Im Vergleich zu den verfügbaren Daten aus Reinkulturen, wiesen die mikrobiellen Gemeinschaften der untersuchten Böden in der Regel eine deutlich höhere katalytische Effizienz auf. Auch für die Michaelis-Menten-Konstante sind die Faktoren enzymspezifisch. So ist für die Km von XYL der Boden-pH-Wert, für AP das C:N Verhältnis und für NAG die DOC-Menge entscheidend. Darüber hinaus haben sich deutliche Beziehungen zwischen der Menge an Gram(-)-Bakterien und der Substrataffinitäten von XYL und AP ergeben. Je höher die Gram(-)-Abundanz, desto höher war die Substrataffinität der Enzymsysteme. Gegenüber alter und degradierter Weiden, war der Umsatz der untersuchten Substrate im Oberboden der aktiv genutzten Weide drei- bis vierfach erhöht. In einem 5-jährigen Düngeexperiment in der Bergregenwaldregion der Anden Süd-Ecuadors wurden die Reaktionen des auf dieser Fläche N/P co-limitierten Grases (Setaria sphacelata), der Arbuskulären Mykorrhiza (AMF) sowie der Bodenmikroorganismen auf moderate N, P und N+P-Düngung untersucht. Die Zugabe von N+P erhöhte die oberirdische Biomasse (+61%) wohingegen die Wurzelbiomasse durch die Zugabe von P (+45%) anstieg. Die C:N:P Verhältnisse weisen auf veränderte P-Aufnahme oder Translokation von P in die Wurzeln hin. Im Besonderen verengte sich das Wurzel C:N and C:P in der P- und der N-Zugabe. Die aus dem Boden extrahierbaren C, N und P-Fraktionen wurden deutlich beeinflusst. Die Zugabe von P stimulierte die Biomasse Gram-(+)-Bakterien (+22%), die Abundanz der AMF (+46%) und die Brutto-N-Mineralisierung. Die Auswertungen deuten darauf hin, dass die Nährstoffanreicherung indirekt über die Veränderungen der Graswurzeln auf die Bodenorganismen wirkte. Die Ergebnisse bestätigen, dass N und P in den Reaktionen von co-limitierten Pflanzen eng miteinander verbunden sind. Vor allem aber steuert P grundlegend die Allokation von Ressourcen und wirkt damit auf andere Ökosystem-komponenten, z.B. auf die Struktur und Aktivität der Bodenmikroorganismen.

Global change phenomena, such as forest disturbance and land-use change significantly affect elemental balances as well as the structure and function of terrestrial ecosystems. Inappropriate land management often causes nutrient losses and finally soil degradation and loss of soil functioning. Especially in tropical ecoregions, soil degradation by nutrient losses is widely abundant. Soil microorganisms are the proximate agents of many processes performed in soils and are regarded as sensitive bio-indicators. However, the incorporation of microbial responses to the definition of critical soil conditions is not intensively developed. In the present thesis, several data analyses of the relationships between ecosystem disturbance and land-use change (natural forest, pastures of different ages, secondary succession) and a diverse set of soil ecological characteristics in the tropical mountain rainforest region of southern Ecuador were compiled. In particular, it was tested whether soil microbial biomass and community functioning were sensitive to land-use change effects. Furthermore, an information-theoretic approach was applied to find the factors that regulate soil microbial biomass and community function. Finally, in a nutrient enrichment experiment the above- and belowground responses to N and P additions were examined. The tested research questions and results were linked to the theory of ecological stoichiometry in order to connect the research to a sound and unifying scientific basis. Soil and microbial stoichiometry were affected by both land-use change and soil depth. After forest disturbance, significant decreases of soil C:N:P ratios at the pastures were fol-lowed by increases during secondary succession. Microbial C:N ratios varied slightly in response to land-use change, whereas no fixed microbial C:P and N:P ratios were observed. Shifts in microbial community composition were associated with soil and microbial stoichiometry. Strong positive relationships between PLFA-markers 18:2n6,9c (saprotrophic fungi) and 20:4 (animals) and negative associations between 20:4 and microbial N:P point to land-use change affecting the structure of soil food webs. Significant deviations from global soil and microbial C:N:P ratios indicated a major force of land-use change to alter stoichiometric relationships and to structure biological systems. Data analysis reveals a strong impact of land-use change on soil microbial biomass, C-mineralization, gross-NH4-consumption and –production rates. According to the results of the IT-approach, combined models better describe effects of land-use change on soil microorganisms than single explanation models. Microbial resources and soil chemical environment were important pre-dictors for soil microbial biomass and community functioning. Little is known about the environmental drivers of the catalytic properties of EHEs (e.g., pH, nutrients) and their functional link to the structure of soil microbial communities. The activities of the six hydrolytic enzymes were tested. Microbial production of AP responded to the low P status of the sites by a higher investment in the acquisition of P compared to C. Three major drivers of enzyme activities were found to be significant for enzyme production: 1.) Microbial demand for P regulated the production of AP, provided that N and C were available. At the natural forest site the two-fold higher specific activity of AP pointed to a high microbial P-demand, whereas the production of AP was constrained by the availability of N and DOC after pasture abandonment. 2.) Microbial biomass that was controlled by pH and resource availability was the main driver for CBH, BG and NAG activities. 3.) Substrate induction due to increased litter inputs of herbaceous plant species seemed to regulate AG and XYL activities during secondary succession. The enzymes’ affinity to substrate, as a potentially critically enzyme kinetic parameter is understudied. The data analysis suggests that microbial communities adapted to environmental changes, demonstrated high flexibility of extracellular enzyme systems and selected for enzymes with higher catalytic efficiency compared with pure cultures. Under in situ conditions, enzyme-specific environmental drivers of the Km, e.g., the pH for XYL, the C:N ratio for AP, and the C availability for NAG were found. The data demonstrated that the higher substrate affinity of XYL and AP was associated with more abundance of Gram(-) bacteria. The catalytic efficiency of enzymes decomposing cellulose, hemicellulose, and starch positively correlated with the relative abundance of Gram(-) bacteria. The turnover rate of the tested substrates was three to four times faster at the young pasture site compared with the longterm pasture and secondary succession sites. Nutrient inputs by atmospheric deposition are known to affect terrestrial ecosystems. However, little is known about how N and P co-limited ecosystems respond to single nutrient enrichment. In this work the susceptibility of above- and belowground ecosystem compo-nents and of their linkages in an N and P co-limited pasture to N- and P-enrichment was assessed. It was tested if the plants´responses can be explained by the concept of serially linked nutrients introduced by Ågren (2004). In this concept, the control of the growth rate by one nutrient is assumed to depend on the control of a different cellular process by another nutrient. The responses of shoot and root biomass and C:N:P stoichiometry of the grass Setaria sphacelata (Schumach.) to moderate N, P, and N+P application over five years were investigated. In addition, the effects of nutrient enrichment on soil nutrient pools, on arbuscular mycorrhizal fungi (AMF) as well as on microbial biomass, activity, and community structure were tested. In order to evaluate the importance of different factors explaining microbial responses, a likelihood-based information-theoretic approach was applied. The application of N+P increased aboveground grass biomass. Root biomass was stimulated by P-treatment. Grass C:N:P stoichiometry responded by altering the P-uptake or by translocating P from shoot to root. In particular, root C:N and C:P stoichiometry decreased in P- and in N-treatment. Extractable fractions of soil C, N, and P were significantly affected by nutrient enrichment. P application increased the biomass of Gram-positive bacteria and the abundance of AMF, however, results of the IT-approach suggested indirect effects of nutrient enrichment on microbes. The responses of the N and P co-limited pasture to particular nutrient enrichment support the concept of serially linked nutrients. The present study provides evidence for the fundamental importance of P for controlling resource allocation of plants in responses to nutrient enrichment. Resource allocation of the grass rather than direct effects of nutrient additions drives changes in AMF, microbial biomass, community structure, and activity. / Seit dem Übergang vom Holozän zum Anthropozän greift der Mensch immer stärker in globale und regionale Stoffkreisläufe ein. Durch die Zerstörung von Naturwäldern und Landnutzungswandel werden die Strukturen und die Funktionen der Ökosysteme stark verändert. Unangepasste Landnutzung führt zu Nährelementverlusten, die mittel- bis langfristige zur Bodendegradation und zur Reduktion von Bodenfunktionen führen. Solche Veränderungen sind insbesondere in den Tropen zu beobachten. Bodenmikroorganismen spielen in den Stoffkreisläufen eine zentrale Rolle. Zudem sind sie sensitive Bioindikatoren für den Zustand von Ökosystemen. Im Gegensatz dazu, werden die Bodenmikroorganismen noch nicht ausreichend für die Zustandsbewertung von Ökosystemen verwendet. In der vorliegenden Dissertation werden verschiedene Datenanalysen zu den Beziehungen von Landnutzungswandel (Naturwald, Weiden verschiedener Alter, sekundäre Sukzession) und den Eigenschaften der Bodenmikroorganismen in einer tropischen Bergregenwaldregion Süd-Ecuadors zusammengefasst. Ein besonderer Fokus lag darauf zu prüfen, ob die mikrobielle Biomasse und die Funktionen die von der mikrobiellen Gemeinschaft geleistet werden (z.B. Enzymaktivitäten) durch den Landnutzungswandel beeinflusst werden. Ein informations-theoretischer Ansatz wurde verwendet um verschiedene Erklärungsansätze der steuernden Faktoren vergleichend zu testen. Darüber hinaus wurden in einem Weidedüngungsexperiment die Reaktionen der ober- und der unterirdischen Ökosystemkomponenten auf die Anreicherung mit N und P getestet. Um die Ergebnisse auf eine breite wissenschaftliche Basis zu stellen wurde die Untersuchungen in den Kontext der Theorie die Ökologischen Stöchiometrie eingeordnet. Die C:N:P Stöchiometrie im Boden und in den Mikroorganismen veränderte sich durch den Landnutzungswandel und mit der Bodentiefe. Mit der Weideetablierung nahmen die C:N:P Verhältnisse im Boden deutlich ab, stiegen dann nach dem Verlassen der Weiden im Zuge der sekundären Sukzession wieder an. Das mikrobielle C:N Verhältnis variierte nur leicht, dagegen zeigten das C:P und N:P Verhältnis deutliche Veränderungen durch den Landnutzungswandel. Mit diesen Veränderungen in der Boden- und Organismenstöchiometrie waren auch Veränderungen in der Struktur der mikrobiellen Gemeinschaften verbunden. Deutliche positive Beziehungen existierten zwischen den saprotrophen Pilzen und den Protozoen. Die steigenden Mengen von Protozoen waren wiederrum mit sinkendem mikrobiellen N:P verbunden. Diese Muster weisen auf Veränderungen in den Bodennahrungsnetzten durch Landnutzungsänderungen hin. Sehr deutliche Abweichungen von globalen Mustern der C:N:P Stöchiometrie deuten darauf hin, dass der Landnutzungswandel signifikanten Einfluss auf die C:N:P Stöchiometrie ausübt. Der Landnutzungswandel beeinflusste auch die mikrobielle Biomasse, die Basalatmung, sowie die mikrobielle Aufnahme und Produktion von NH4-N im Boden. Dabei zeigten kombinierte Erklärungsansätze die adäquateren Beschreibungen der Muster. In den kombinierten Modellen zur Erklärung der mikrobiellen Biomasse und der mikrobiellen Leistungen überwogen Prädiktoren der mikrobiellen Ressourcen und der bodenchemischen Umwelt. Ein weiterer Schwerpunkt der Untersuchungen lag auf der Erfassung der Effekte des Land-nutzungswandels auf die Aktivität von extrazellulären Bodenenzymen. Bisher ist wenig darüber bekannt, welche Faktoren die katalytischen Eigenschaften steuern und beispielsweise, ob es Zusammenhänge zur mikrobiellen Gemeinschaftsstruktur gibt. Um diese Fragen näher zu beleuchten wurden sechs hydrolytische Enzyme basierend auf MUF-Substraten untersucht. Die mikrobielle Produktion von AP stand dabei in Zusammenhang mit dem niedrigen P-Status der untersuchten Böden. Das wurde besonders durch die hohe AP Produktion im Vergleich zu BG belegt. Im Allgemeinen konnten drei verschiedene Mechanismen festgestellt werden, die die Produktion der untersuchten EHEs vermutlich steuerten. 1.) Der P-Bedarf der Mikroorganismen regulierte die Produktion von AP, vorausgesetzt, dass ausreichend N und C zur Enzymsynthese zur Verfügung standen. 2.) Die Höhe der mikrobiellen Biomasse hat sich als wichtiger Faktor für die Produktion von CBH, BG und NAG gezeigt. Das deutet auf die konstitutive Produktion dieser Enzyme hin. 3.) Die substratinduzierte Produktion von Enzymen ist vermutlich entscheidend für die Aktivität von AG und XYL. Die Berücksichtigung der Enzymkinetiken, insbesondere der Michaelis-Menten-Konstante lieferte weitere Aufschlüsse über relevante Faktoren. Im Allgemeinen so scheint es, haben sich die mikrobiellen Gemeinschaften an die starken Umweltgradienten, die durch den Landnutzungswandel erzeugt worden angepasst. Im Vergleich zu den verfügbaren Daten aus Reinkulturen, wiesen die mikrobiellen Gemeinschaften der untersuchten Böden in der Regel eine deutlich höhere katalytische Effizienz auf. Auch für die Michaelis-Menten-Konstante sind die Faktoren enzymspezifisch. So ist für die Km von XYL der Boden-pH-Wert, für AP das C:N Verhältnis und für NAG die DOC-Menge entscheidend. Darüber hinaus haben sich deutliche Beziehungen zwischen der Menge an Gram(-)-Bakterien und der Substrataffinitäten von XYL und AP ergeben. Je höher die Gram(-)-Abundanz, desto höher war die Substrataffinität der Enzymsysteme. Gegenüber alter und degradierter Weiden, war der Umsatz der untersuchten Substrate im Oberboden der aktiv genutzten Weide drei- bis vierfach erhöht. In einem 5-jährigen Düngeexperiment in der Bergregenwaldregion der Anden Süd-Ecuadors wurden die Reaktionen des auf dieser Fläche N/P co-limitierten Grases (Setaria sphacelata), der Arbuskulären Mykorrhiza (AMF) sowie der Bodenmikroorganismen auf moderate N, P und N+P-Düngung untersucht. Die Zugabe von N+P erhöhte die oberirdische Biomasse (+61%) wohingegen die Wurzelbiomasse durch die Zugabe von P (+45%) anstieg. Die C:N:P Verhältnisse weisen auf veränderte P-Aufnahme oder Translokation von P in die Wurzeln hin. Im Besonderen verengte sich das Wurzel C:N and C:P in der P- und der N-Zugabe. Die aus dem Boden extrahierbaren C, N und P-Fraktionen wurden deutlich beeinflusst. Die Zugabe von P stimulierte die Biomasse Gram-(+)-Bakterien (+22%), die Abundanz der AMF (+46%) und die Brutto-N-Mineralisierung. Die Auswertungen deuten darauf hin, dass die Nährstoffanreicherung indirekt über die Veränderungen der Graswurzeln auf die Bodenorganismen wirkte. Die Ergebnisse bestätigen, dass N und P in den Reaktionen von co-limitierten Pflanzen eng miteinander verbunden sind. Vor allem aber steuert P grundlegend die Allokation von Ressourcen und wirkt damit auf andere Ökosystem-komponenten, z.B. auf die Struktur und Aktivität der Bodenmikroorganismen.

It is increasingly being recognized that the soil microbes can mineralize recalcitrant soil organic matter (SOM) by using the fresh carbon (C) as a source of energy, a process called priming effect (PE). It has been shown mostly in lab incubations that PE can have important consequences for sequestration of organic C in soils. However, the importance of PE in C and N dynamics of ecosystems remains little known. The soil-plant interactions and rhizospheric processes can modulate the rates of PE and its consequences on C and N dynamics in an ecosystem. The objective of this thesis was to determine the role of PE in the C and N dynamics of permanent grasslands and the modulation of this role in response to management (plant clipping, fertilization) and global warming. Moreover, it was aimed to identify the microbial groups involved in PE and to unravel the way, e.g. absorption of N, root exudations and litter deposition, by which plant can induce PE. The thesis was based on a new approach allowing continuous dual labelling of multiple grassland plants with 13C- and 14C-CO2. The dual labelling permitted the separation of soil-derived CO2 from plant-derived CO2, the calculation of PE and the determination of mean age of soil-derived CO2-C. Moreover, phospholipids fatty-acids analysis (PLFA) permitted to correlate the variation of PE with changes in microbial community composition. Our work showed that the increased SOM mineralization under grasses was consistently two to three times more than that in bare soils (i.e. PE) over long term (511 days). This reveals that the PE plays key role in ecosystem CO2-C flux and indicates that a very large pool of SOM is under the control of PE. Moreover, we report that 15,000 years old organic C from an undisturbed deep soil can be mineralized after the supply of fresh C by living plants to soil microbes. This result supports the idea that the SOM in deep soils is stable due to the energy-limitation of microbes and the ‘inert' pool of organic C defined in current models is not so ‘inert' finally. The supply of N in soil-plant system through the use of fertilizer or legume decreased the PE suggesting that the C storage in soils is limited by nutrient supply. Similarly, plant clipping reduced the plant N uptake thereby PE. Collectively these results suggest synchronization between plant N uptake and SOM mineralization supporting the idea that soils under permanent plant cover function as a bank of nutrients for the plant, maximizing plant productivity and nutrient retention. An innovative method clearly showed that the root exudation is the major way by which grassland plants induce PE. Moreover, saprophytic fungi are suggested as the key actors in the mineralization of recalcitrant SOM & PE. Lastly, we developed a new theory on temperature response of SOM mineralization by taking into account the energy-limitation of microbes and the temperature-dependent inactivation of enzymes. This theory predicts a negative relationship between temperature and mineralization of recalcitrant SOM, which was supported by experimental results. This finding challenges the classical paradigm of positive relationship between temperature and recalcitrant SOM mineralization. Overall, these investigations on plant-soil systems reinforce the idea that PE and underlying mechanisms play a key role in ecosystem C and N dynamics and even suggest that this role was underestimated in lab experiments. / It is increasingly being recognized that the soil microbes can mineralize recalcitrant soil organic matter (SOM) by using the fresh carbon (C) as a source of energy, a process called priming effect (PE). It has been shown mostly in lab incubations that PE can have important consequences for sequestration of organic C in soils. However, the importance of PE in C and N dynamics of ecosystems remains little known. The soil-plant interactions and rhizospheric processes can modulate the rates of PE and its consequences on C and N dynamics in an ecosystem. The objective of this thesis was to determine the role of PE in the C and N dynamics of permanent grasslands and the modulation of this role in response to management (plant clipping, fertilization) and global warming. Moreover, it was aimed to identify the microbial groups involved in PE and to unravel the way, e.g. absorption of N, root exudations and litter deposition, by which plant can induce PE. The thesis was based on a new approach allowing continuous dual labelling of multiple grassland plants with 13C- and 14C-CO2. The dual labelling permitted the separation of soil-derived CO2 from plant-derived CO2, the calculation of PE and the determination of mean age of soil-derived CO2-C. Moreover, phospholipids fatty-acids analysis (PLFA) permitted to correlate the variation of PE with changes in microbial community composition. Our work showed that the increased SOM mineralization under grasses was consistently two to three times more than that in bare soils (i.e. PE) over long term (511 days). This reveals that the PE plays key role in ecosystem CO2-C flux and indicates that a very large pool of SOM is under the control of PE. Moreover, we report that 15,000 years old organic C from an undisturbed deep soil can be mineralized after the supply of fresh C by living plants to soil microbes. This result supports the idea that the SOM in deep soils is stable due to the energy-limitation of microbes and the ‘inert' pool of organic C defined in current models is not so ‘inert' finally. The supply of N in soil-plant system through the use of fertilizer or legume decreased the PE suggesting that the C storage in soils is limited by nutrient supply. Similarly, plant clipping reduced the plant N uptake thereby PE. Collectively these results suggest synchronization between plant N uptake and SOM mineralization supporting the idea that soils under permanent plant cover function as a bank of nutrients for the plant, maximizing plant productivity and nutrient retention. An innovative method clearly showed that the root exudation is the major way by which grassland plants induce PE. Moreover, saprophytic fungi are suggested as the key actors in the mineralization of recalcitrant SOM & PE. Lastly, we developed a new theory on temperature response of SOM mineralization by taking into account the energy-limitation of microbes and the temperature-dependent inactivation of enzymes. This theory predicts a negative relationship between temperature and mineralization of recalcitrant SOM, which was supported by experimental results. This finding challenges the classical paradigm of positive relationship between temperature and recalcitrant SOM mineralization. Overall, these investigations on plant-soil systems reinforce the idea that PE and underlying mechanisms play a key role in ecosystem C and N dynamics and even suggest that this role was underestimated in lab experiments.

Agricultural intensification reliant on monocrops could change soil health in a way that does not support maximum crop productivity. Twenty-nine-year-old no-till field plots at the University of Kentucky Spindletop research farm showed a significant reduction in corn yields from continuous corn plots compared to those from plots in various types of rotation. The objective of this study was to determine what role soil microbes might play in yield reduction and how management and time effects microbial community structure. Samples were collected from the following treatments: continuous corn (CC), continuous soybean (SS), a 2-year corn/soybean rotation (CCSS), Corn in rotation with soybean with winter wheat cover (C/W/S), and sod controls (SOD). Soil health-related parameters were determined along with microbial community structure using phospholipid fatty acid analysis (PLFA). Results show that there is a strong seasonal dynamic in microbial communities with May, July and September showing the greatest differentiation between treatments. Nonparametric multidimensional analysis (NMDS) shows that microbial communities under SS, CC treatments were significantly different from the CS and CWS treatments across all four years of the study. My findings will prove useful for assessing the contribution of biological indicators to agroecosystem function and will aid in making recommendations of when and how to manage these parameters to improve soil health and maximize yield.

It is increasingly being recognized that the soil microbes can mineralize recalcitrant soil organic matter (SOM) by using the fresh carbon (C) as a source of energy, a process called priming effect (PE). It has been shown mostly in lab incubations that PE can have important consequences for sequestration of organic C in soils. However, the importance of PE in C and N dynamics of ecosystems remains little known. The soil-plant interactions and rhizospheric processes can modulate the rates of PE and its consequences on C and N dynamics in an ecosystem. The objective of this thesis was to determine the role of PE in the C and N dynamics of permanent grasslands and the modulation of this role in response to management (plant clipping, fertilization) and global warming. Moreover, it was aimed to identify the microbial groups involved in PE and to unravel the way, e.g. absorption of N, root exudations and litter deposition, by which plant can induce PE. The thesis was based on a new approach allowing continuous dual labelling of multiple grassland plants with 13C- and 14C-CO2. The dual labelling permitted the separation of soil-derived CO2 from plant-derived CO2, the calculation of PE and the determination of mean age of soil-derived CO2-C. Moreover, phospholipids fatty-acids analysis (PLFA) permitted to correlate the variation of PE with changes in microbial community composition. Our work showed that the increased SOM mineralization under grasses was consistently two to three times more than that in bare soils (i.e. PE) over long term (511 days). This reveals that the PE plays key role in ecosystem CO2-C flux and indicates that a very large pool of SOM is under the control of PE. Moreover, we report that 15,000 years old organic C from an undisturbed deep soil can be mineralized after the supply of fresh C by living plants to soil microbes. This result supports the idea that the SOM in deep soils is stable due to the energy-limitation of microbes and the 'inert' pool of organic C defined in current models is not so 'inert' finally. The supply of N in soil-plant system through the use of fertilizer or legume decreased the PE suggesting that the C storage in soils is limited by nutrient supply. Similarly, plant clipping reduced the plant N uptake thereby PE. Collectively these results suggest synchronization between plant N uptake and SOM mineralization supporting the idea that soils under permanent plant cover function as a bank of nutrients for the plant, maximizing plant productivity and nutrient retention. An innovative method clearly showed that the root exudation is the major way by which grassland plants induce PE. Moreover, saprophytic fungi are suggested as the key actors in the mineralization of recalcitrant SOM & PE. Lastly, we developed a new theory on temperature response of SOM mineralization by taking into account the energy-limitation of microbes and the temperature-dependent inactivation of enzymes. This theory predicts a negative relationship between temperature and mineralization of recalcitrant SOM, which was supported by experimental results. This finding challenges the classical paradigm of positive relationship between temperature and recalcitrant SOM mineralization. Overall, these investigations on plant-soil systems reinforce the idea that PE and underlying mechanisms play a key role in ecosystem C and N dynamics and even suggest that this role was underestimated in lab experiments.

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This diploma thesis has been carried out as a part of the project Utilization of long term (passive) sampling methods combined with in situ microcosms for assessment of (bio)degradation potential (PASSES). In the frame of the project groundwater remediation took place in the premises of Farmak a.s. in Olomouc using a pilot photooxidation unit and efficiency of the remediation was monitored through passive and active sampling methods. Pilot photooxidation unit is a technology based on the H2O2/UV-C photochemical oxidation of organic pollutants. In this work optimization tests of the pilot photooxidation unit were performed. The residence time of the groundwater in the photoreactors, required for its sufficient decontamination from pharmaceuticals and aromatic hydrocarbons, was 2.5 hours. 91% degradation of the pharmaceuticals and 80% degradation of aromatic hydrocarbons were reached during this interval. Although the removal efficiency of the pharmaceuticals by the photooxidation unit was high, the pilot photooxidation unit was not able to effectively remove the pharmaceuticals at the studied locality. By comparing the results of the pharmaceuticals from active and passive groundwater sampling during the remediation attempt, passive Polar Organic Chemical Integrative Sampler (POCIS) was found to be...

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