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Metabolically active microbial communities are present in a wide range of subsurface environments. Techniques like enumeration of microbial cells, activity measurements with radiotracer assays and the analysis of porewater constituents are currently being used to explore the subsurface biosphere, alongside with molecular biological analyses. However, many of these techniques reach their detection limits due to low microbial activity and abundance. Direct measurements of microbial turnover not just face issues of insufficient sensitivity, they only provide information about a single specific process but in sediments many different process can occur simultaneously. Therefore, the development of a new technique to measure total microbial activity would be a major improvement. A new tritium-based hydrogenase-enzyme assay appeared to be a promising tool to quantify total living biomass, even in low activity subsurface environments. In this PhD project total microbial biomass and microbial activity was quantified in different subsurface sediments using established techniques (cell enumeration and pore water geochemistry) as well as a new tritium-based hydrogenase enzyme assay. By using a large database of our own cell enumeration data from equatorial Pacific and north Pacific sediments and published data it was shown that the global geographic distribution of subseafloor sedimentary microbes varies between sites by 5 to 6 orders of magnitude and correlates with the sedimentation rate and distance from land. Based on these correlations, global subseafloor biomass was estimated to be 4.1 petagram-C and ~0.6 % of Earth's total living biomass, which is significantly lower than previous estimates. Despite the massive reduction in biomass the subseafloor biosphere is still an important player in global biogeochemical cycles. To understand the relationship between microbial activity, abundance and organic matter flux into the sediment an expedition to the equatorial Pacific upwelling area and the north Pacific Gyre was carried out. Oxygen respiration rates in subseafloor sediments from the north Pacific Gyre, which are deposited at sedimentation rates of 1 mm per 1000 years, showed that microbial communities could survive for millions of years without fresh supply of organic carbon. Contrary to the north Pacific Gyre oxygen was completely depleted within the upper few millimeters to centimeters in sediments of the equatorial upwelling region due to a higher supply of organic matter and higher metabolic activity. So occurrence and variability of electron acceptors over depth and sites make the subsurface a complex environment for the quantification of total microbial activity. Recent studies showed that electron acceptor processes, which were previously thought to thermodynamically exclude each other can occur simultaneously. So in many cases a simple measure of the total microbial activity would be a better and more robust solution than assays for several specific processes, for example sulfate reduction rates or methanogenesis. Enzyme or molecular assays provide a more general approach as they target key metabolic compounds. Since hydrogenase enzymes are ubiquitous in microbes, the recently developed tritium-based hydrogenase radiotracer assay is applied to quantify hydrogenase enzyme activity as a parameter of total living cell activity. Hydrogenase enzyme activity was measured in sediments from different locations (Lake Van, Barents Sea, Equatorial Pacific and Gulf of Mexico). In sediment samples that contained nitrate, we found the lowest cell specific enzyme activity around 10^(-5) nmol H_(2) cell^(-1) d^(-1). With decreasing energy yield of the electron acceptor used, cell-specific hydrogenase activity increased and maximum values of up to 1 nmol H_(2) cell^(-1) d^(-1) were found in samples with methane concentrations of >10 ppm. Although hydrogenase activity cannot be converted directly into a turnover rate of a specific process, cell-specific activity factors can be used to identify specific metabolism and to quantify the metabolically active microbial population. In another study on sediments from the Nankai Trough microbial abundance and hydrogenase activity data show that both the habitat and the activity of subseafloor sedimentary microbial communities have been impacted by seismic activities. An increase in hydrogenase activity near the fault zone revealed that the microbial community was supplied with hydrogen as an energy source and that the microbes were specialized to hydrogen metabolism.
In soils and sediments there is a strong coupling between local biogeochemical processes and the distribution of water, electron acceptors, acids and nutrients. Both sides are closely related and affect each other from small scale to larger scales. Soil structures such as aggregates, roots, layers or macropores enhance the patchiness of these distributions. At the same time it is difficult to access the spatial distribution and temporal dynamics of these parameter. Noninvasive imaging techniques with high spatial and temporal resolution overcome these limitations. And new non-invasive techniques are needed to study the dynamic interaction of plant roots with the surrounding soil, but also the complex physical and chemical processes in structured soils. In this study we developed an efficient non-destructive in-situ method to determine biogeochemical parameters relevant to plant roots growing in soil. This is a quantitative fluorescence imaging method suitable for visualizing the spatial and temporal pH changes around roots. We adapted the fluorescence imaging set-up and coupled it with neutron radiography to study simultaneously root growth, oxygen depletion by respiration activity and root water uptake. The combined set up was subsequently applied to a structured soil system to map the patchy structure of oxic and anoxic zones induced by a chemical oxygen consumption reaction for spatially varying water contents. Moreover, results from a similar fluorescence imaging technique for nitrate detection were complemented by a numerical modeling study where we used imaging data, aiming to simulate biodegradation under anaerobic, nitrate reducing conditions.
Multi-messenger constraints and pressure from dark matter annihilation into electron-positron pairs
(2013)
Despite striking evidence for the existence of dark matter from astrophysical observations, dark matter has still escaped any direct or indirect detection until today. Therefore a proof for its existence and the revelation of its nature belongs to one of the most intriguing challenges of nowadays cosmology and particle physics. The present work tries to investigate the nature of dark matter through indirect signatures from dark matter annihilation into electron-positron pairs in two different ways, pressure from dark matter annihilation and multi-messenger constraints on the dark matter annihilation cross-section. We focus on dark matter annihilation into electron-positron pairs and adopt a model-independent approach, where all the electrons and positrons are injected with the same initial energy E_0 ~ m_dm*c^2. The propagation of these particles is determined by solving the diffusion-loss equation, considering inverse Compton scattering, synchrotron radiation, Coulomb collisions, bremsstrahlung, and ionization. The first part of this work, focusing on pressure from dark matter annihilation, demonstrates that dark matter annihilation into electron-positron pairs may affect the observed rotation curve by a significant amount. The injection rate of this calculation is constrained by INTEGRAL, Fermi, and H.E.S.S. data. The pressure of the relativistic electron-positron gas is computed from the energy spectrum predicted by the diffusion-loss equation. For values of the gas density and magnetic field that are representative of the Milky Way, it is estimated that the pressure gradients are strong enough to balance gravity in the central parts if E_0 < 1 GeV. The exact value depends somewhat on the astrophysical parameters, and it changes dramatically with the slope of the dark matter density profile. For very steep slopes, as those expected from adiabatic contraction, the rotation curves of spiral galaxies would be affected on kiloparsec scales for most values of E_0. By comparing the predicted rotation curves with observations of dwarf and low surface brightness galaxies, we show that the pressure from dark matter annihilation may improve the agreement between theory and observations in some cases, but it also imposes severe constraints on the model parameters (most notably, the inner slope of the halo density profile, as well as the mass and the annihilation cross-section of dark matter particles into electron-positron pairs). In the second part, upper limits on the dark matter annihilation cross-section into electron-positron pairs are obtained by combining observed data at different wavelengths (from Haslam, WMAP, and Fermi all-sky intensity maps) with recent measurements of the electron and positron spectra in the solar neighbourhood by PAMELA, Fermi, and H.E.S.S.. We consider synchrotron emission in the radio and microwave bands, as well as inverse Compton scattering and final-state radiation at gamma-ray energies. For most values of the model parameters, the tightest constraints are imposed by the local positron spectrum and synchrotron emission from the central regions of the Galaxy. According to our results, the annihilation cross-section should not be higher than the canonical value for a thermal relic if the mass of the dark matter candidate is smaller than a few GeV. In addition, we also derive a stringent upper limit on the inner logarithmic slope α of the density profile of the Milky Way dark matter halo (α < 1 if m_dm < 5 GeV, α < 1.3 if m_dm < 100 GeV and α < 1.5 if m_dm < 2 TeV) assuming a dark matter annihilation cross-section into electron-positron pairs (σv) = 3*10^−26 cm^3 s^−1, as predicted for thermal relics from the big bang.
Functional metabolism of storage carbohydrates is vital to plants and animals. The water-soluble glycogen in animal cells and the amylopectin which is the major component of water-insoluble starch granules residing in plant plastids are chemically similar as they consist of α-1,6 branched α-1,4 glucan chains. Synthesis and degradation of transitory starch and of glycogen are accomplished by a set of enzymatic activities that to some extend are also similar in plants and animals. Chain elongation, branching, and debranching are achieved by synthases, branching enzymes, and debranching enzymes, respectively. Similarly, both types of polyglucans contain low amounts of phosphate esters whose abundance varies depending on species and organs. Starch is selectively phosphorylated by at least two dikinases (GWD and PWD) at the glucosyl carbons C6 and C3 and dephosphorylated by the phosphatase SEX4 and SEX4-like enzymes. In Arabidopsis insufficiency in starch phosphorylation or dephosphorylation results in largely impaired starch turnover, starch accumulation, and often in retardation of growth. In humans the progressive neurodegenerative epilepsy, Lafora disease, is the result of a defective enzyme (laforin) that is functional equivalent to the starch phosphatase SEX4 and capable of glycogen dephosphorylation. Patients lacking laforin progressively accumulate unphysiologically structured insoluble glycogen-derived particles (Lafora bodies) in many tissues including brain. Previous results concerning the carbon position of glycogen phosphate are contradictory. Currently it is believed that glycogen is esterified exclusively at the carbon positions C2 and C3 and that the monophosphate esters, being incorporated via a side reaction of glycogen synthase (GS), lack any specific function but are rather an enzymatic error that needs to be corrected. In this study a versatile and highly sensitive enzymatic cycling assay was established that enables quantification of very small G6P amounts in the presence of high concentrations of non-target compounds as present in hydrolysates of polysaccharides, such as starch, glycogen, or cytosolic heteroglycans in plants. Following validation of the G6P determination by analyzing previously characterized starches G6P was quantified in hydrolysates of various glycogen samples and in plant heteroglycans. Interestingly, glucosyl C6 phosphate is present in all glycogen preparations examined, the abundance varying between glycogens of different sources. Additionally, it was shown that carbon C6 is severely hyperphosphorylated in glycogen of Lafora disease mouse model and that laforin is capable of removing C6 phosphate from glycogen. After enrichment of phosphoglucans from amylolytically degraded glycogen, several techniques of two-dimensional NMR were applied that independently proved the existence of 6-phosphoglucosyl residues in glycogen and confirmed the recently described phosphorylation sites C2 and C3. C6 phosphate is neither Lafora disease- nor species-, or organ-specific as it was demonstrated in liver glycogen from laforin-deficient mice and in that of wild type rabbit skeletal muscle. The distribution of 6-phosphoglucosyl residues was analyzed in glycogen molecules and has been found to be uneven. Gradual degradation experiments revealed that C6 phosphate is more abundant in central parts of the glycogen molecules and in molecules possessing longer glucan chains. Glycogen of Lafora disease mice consistently contains a higher proportion of longer chains while most short chains were reduced as compared to wild type. Together with results recently published (Nitschke et al., 2013) the findings of this work completely unhinge the hypothesis of GS-mediated phosphate incorporation as the respective reaction mechanism excludes phosphorylation of this glucosyl carbon, and as it is difficult to explain an uneven distribution of C6 phosphate by a stochastic event. Indeed the results rather point to a specific function of 6-phosphoglucosyl residues in the metabolism of polysaccharides as they are present in starch, glycogen, and, as described in this study, in heteroglycans of Arabidopsis. In the latter the function of phosphate remains unclear but this study provides evidence that in starch and glycogen it is related to branching. Moreover a role of C6 phosphate in the early stages of glycogen synthesis is suggested. By rejecting the current view on glycogen phosphate to be a stochastic biochemical error the results permit a wider view on putative roles of glycogen phosphate and on alternative biochemical ways of glycogen phosphorylation which for many reasons are likely to be mediated by distinct phosphorylating enzymes as it is realized in starch metabolism of plants. Better understanding of the enzymology underlying glycogen phosphorylation implies new possibilities of Lafora disease treatment.
In children the way of life, nutrition and recreation changed in recent years and as a consequence body composition shifted as well. It is established that overweight belongs to a global problem. In addition, German children exhibit a less robust skeleton than ten years ago. These developments may elevate the risk of cardiovascular diseases and skeletal modifications. Heredity and environmental factors as nutrition, socioeconomic status, physical activity and inactivity influence fat accumulation and the skeletal system. Based on these negative developments associations between type of body shape, skeletal measures and physical activity; relations between external skeletal robustness, physical activity and inactivity, BMI and body fat and also the progress of body composition especially external skeletal robustness in comparison in Russian and German children were investigated. In a cross-sectional study 691 German boys and girls aged 6 to 10 years were examined. Anthropometric measurements were taken and questionnaires about physical activity and inactivity were answered by parents. Additionally, pedometers were worn to determinate the physical activity in children. To compare the body composition in Russian and German children data from the years 2000 and 2010 were used. The study has shown that pyknomorphic individuals exhibit the highest external skeletal robustness and leptomorphic ones the lowest. Leptomorphic children may have a higher risk for bone diseases in adulthood. Pyknomorphic boys are more physically active by tendency. This is assessed as positive because pyknomorphic types display the highest BMI and body fat. Results showed that physical activity may reduce BMI and body fat. In contrast physical inactivity may lead to an increase of BMI and body fat and may rise with increasing age. Physical activity encourages additionally a robust skeleton. Furthermore external skeletal robustness is associated with BMI in order that BMI as a measure of overweight should be consider critically. The international 10-year comparison has shown an increase of BMI in Russian children and German boys. Currently, Russian children exhibit a higher external skeletal robustness than the Germans. However, in Russian boys skeleton is less robust than ten years ago. This trend should be observed in the future as well in other countries. All in all, several measures should be used to describe health situation in children and adults. Furthermore, in children it is essential to support physical activity in order to reduce the risk of obesity and to maintain a robust skeleton. In this way diseases are able to prevent in adulthood.
In the context of ecological risk assessment of chemicals, individual-based population models hold great potential to increase the ecological realism of current regulatory risk assessment procedures. However, developing and parameterizing such models is time-consuming and often ad hoc. Using standardized, tested submodels of individual organisms would make individual-based modelling more efficient and coherent. In this thesis, I explored whether Dynamic Energy Budget (DEB) theory is suitable for being used as a standard submodel in individual-based models, both for ecological risk assessment and theoretical population ecology. First, I developed a generic implementation of DEB theory in an individual-based modeling (IBM) context: DEB-IBM. Using the DEB-IBM framework I tested the ability of the DEB theory to predict population-level dynamics from the properties of individuals. We used Daphnia magna as a model species, where data at the individual level was available to parameterize the model, and population-level predictions were compared against independent data from controlled population experiments. We found that DEB theory successfully predicted population growth rates and peak densities of experimental Daphnia populations in multiple experimental settings, but failed to capture the decline phase, when the available food per Daphnia was low. Further assumptions on food-dependent mortality of juveniles were needed to capture the population dynamics after the initial population peak. The resulting model then predicted, without further calibration, characteristic switches between small- and large-amplitude cycles, which have been observed for Daphnia. We conclude that cross-level tests help detecting gaps in current individual-level theories and ultimately will lead to theory development and the establishment of a generic basis for individual-based models and ecology. In addition to theoretical explorations, we tested the potential of DEB theory combined with IBMs to extrapolate effects of chemical stress from the individual to population level. For this we used information at the individual level on the effect of 3,4-dichloroanailine on Daphnia. The individual data suggested direct effects on reproduction but no significant effects on growth. Assuming such direct effects on reproduction, the model was able to accurately predict the population response to increasing concentrations of 3,4-dichloroaniline. We conclude that DEB theory combined with IBMs holds great potential for standardized ecological risk assessment based on ecological models.
Körperliche Attraktivität und gutes Aussehen spielen in der heutigen Gesellschaft eine entscheidende Rolle, was bereits frühzeitig auch Kinder und Jugendliche in ihren Einstellungen und der Wahrnehmung ihres Körpers prägt. Sorgen um den eigenen Körper gelten als normatives Problem unter Jugendlichen und bergen nicht selten das Risiko für gesundheitsgefährdendes Verhalten und psychische Erkrankungen. In der Suche nach den Ursachen gerieten in den letzten Jahren insbesondere soziokulturelle Faktoren, insbesondere der Einfluss von medial vermittelten Schönheitsidealen, in den Fokus der Forschung. Es ist jedoch fraglich, warum nicht alle Jugendlichen in gleicher Weise auf den allgegenwärtigen Mediendruck reagieren. Naheliegend ist, dass die Jugendlichen besonders gefährdet sind, deren unmittelbares soziales Umfeld das geltende Schönheitsideal direkt oder indirekt vermittelt und verstärkt. Das Verständnis der Rolle sozialen Drucks ist jedoch bislang noch durch zahlreiche inhaltliche und methodische Aspekte beschränkt (z.B. Einschränkungen in der Operationalisierung, ungenügende Berücksichtigung geschlechtsspezifischer Mechanismen, fehlende längsschnittliche Belege). Daher widmet sich die vorliegende Arbeit der Bedeutung aussehensbezogenen sozialen Drucks in der Entstehung von Körperunzufriedenheit im Jugendalter in drei aufeinander aufbauenden Untersuchungsschritten. Ausgehend von der Entwicklung eines umfassenden und zuverlässigen Erhebungsinstruments zielt die Arbeit darauf ab, unterschiedliche Aspekte sozialen Drucks gegenüberzustellen und hinsichtlich ihrer Verbreitung und Risikowirkung zu vergleichen. Die Umsetzung des Forschungsvorhabens erfolgte in unterschiedlichen Schülerstichproben der Klassen 7 bis 9 unterschiedlicher Gymnasien und Gesamtschulen (Hauptstichprobe N = 1112, im Mittel = 13.4 ± 0.8 Jahre). Dabei wurden sowohl quer- als auch längsschnittliche Analysen durchgeführt. Zusätzlich wurden zur Erprobung des Fragebogenverfahrens klinische Stichproben mit Ess- und Gewichtsstörungen herangezogen. Zur detaillierten Erfassung unterschiedlicher Formen aussehensbezogenen sozialen Drucks erfolgte im ersten Schritt die Entwicklung des Fragebogen zum aussehensbezogen sozialen Druck (FASD), welcher acht unterschiedliche Formen aussehensbezogene sozialen Drucks ausgehend von Eltern und Gleichaltrigen reliabel und valide erfasst. Dabei erwies sich das Verfahren gleichermaßen für Jungen und Mädchen, wie für Jugendliche mit unterschiedlichem Gewichtsstatus geeignet. Die psychometrische Güte des Verfahrens konnte sowohl für populationsbasierte als auch für klinische Stichproben mit Ess- und Gewichtsstörung belegt werden, wodurch eine breite Einsatzmöglichkeit in Forschung und Praxis denkbar ist. Im zweiten Schritt erfolgte die Untersuchung der Verbreitung aussehensbezogenen sozialen Drucks unter besonderer Berücksichtigung von Geschlechts-, Alters- und Gewichtsgruppenunterschieden. Dabei erwiesen sich Mädchen als stärker von aussehensbezogenem Druck durch Gleichaltrige betroffen als Jungen. Darüberhinaus legen die Ergebnisse nahe, dass Übergewicht ungeachtet des Geschlechts mit verstärkten aussehensbezogenen Abwertungen und Ausgrenzungserfahrungen verbunden ist. Zudem deuten die Alterseffekte der Studie darauf hin, dass der Übergang von früher zu mittlerer Adoleszenz aber auch Schulwechsel besonderes kritische Zeitpunkte für die Etablierung aussehensbezogener Einflüsse darstellen. Abschließend widmete sich die Arbeit der längsschnittlichen Risikowirkung unterschiedlicher Aspekte aussehensbezogenen sozialen Drucks in der Entstehung von Körperunzufriedenheit. Aussehensbezogene Einflüsse von Freunden verstärkten längsschnittlich Körpersorgen sowohl bei Mädchen als auch bei Jungen. Zudem ergab sich das Erleben von Ausgrenzung durch Gleichaltrige als entscheidender Risikofaktor für gewichtsbezogene Körpersorgen unter Jungen. Als bedeutsamster elterlicher Einfluss erwiesen sich Aufforderungen auf die Figur zu achten. Diese Aufforderungen verstärkten gleichermaßen für Mädchen und Jungen gewichtsbezogene Körpersorgen. Die vorliegende Arbeit widmete sich dem Ziel, die Rolle aussehensbezogener sozialer Einflüsse weiter aufzuklären. Das dazu vorgelegte umfassende Instrument ermöglichte eine differenzierte Betrachtung der Verbreitung und Wirkung unterschiedlicher Formen sozialen Drucks. Hierdurch weisen die Ergebnisse nicht nur auf wichtige geschlechtsspezifische Mechanismen hin, sondern leisten ebenso einen Beitrag zum vertieften Verständnis der Risikowirkung sozialen Drucks. Diese Erkenntnisse liefern somit einerseits konkrete Ansatzpunkte für Prävention und Intervention und ermöglichen andererseits auch eine weitere Konkretisierung bereits etablierter soziokultureller Wirkmodelle.
Large areas in the humid tropics are currently undergoing land-use change. The decrease of tropical rainforest, which is felled for land clearing and timber production, is countered by increasing areas of tree plantations and secondary forests. These changes are known to affect the regional water cycle as a result of plant-specific water demand and by influencing key soil properties which determine hydrological flow paths. One of these key properties sensitive to land-use change is the saturated hydraulic conductivity (Ks) as it governs vertical percolation of water within the soil profile. Low values of Ks in a certain soil depth can form an impeding layer and lead to perched water tables and the development of predominantly lateral flow paths such as overland flow. These processes can induce nutrient redistribution, erosion and soil degradation and thus affect ecosystem services and human livelihoods. Due to its sensitivity to land-use change, Ks is commonly used to assess the associated changes in hydrological flow paths. The objective of this dissertation was to assess the effect of land-use change on hydrological flow paths by analysing Ks as indicator variable. Sources of Ks variability, their implications for Ks monitoring and the relationship between Ks and near-surface hydrological flow paths in the context of land-use change were studied. The research area was located in central Panama, a country widely experiencing the abovementioned changes in land use. Ks is dependent on both static, soil-inherent properties such as particle size and clay mineralogy and dynamic, land use-dependent properties such as organic carbon content. By conducting a pair of studies with one of these influences held constant in each, the importance of static and dynamic properties for Ks was assessed. Applying a space-for-time approach to sample Ks under secondary forests of different age classes on comparable soils, a recovery of Ks from the former pasture use was shown to require more than eight years. The process was limited to the 0−6 cm sampling depth and showed large variability among replicates. A wavelet analysis of a Ks transect crossing different soil map units under comparable land cover, old-growth tropical rainforest, showed large small-scale variability, which was attributed to biotic influences, as well as a possible but non-conclusive influence of soil types. The two results highlight the importance of dynamic, land use-dependent influences on Ks. Monitoring studies can help to quantify land use-induced change of Ks, but there is a variety of sampling designs which differ in efficiency of estimating mean Ks. A comparative study of four designs and their suitability for Ks monitoring is used to give recommendations about designing a Ks monitoring scheme. Quantifying changes in spatial means of Ks for small catchments with a rotational stratified sampling design did not prove to be more efficient than Simple Random Sampling. The lack of large-scale spatial structure prevented benefits of stratification, and large small-scale variability resulting from local biotic processes and artificial effects of destructive sampling caused a lack of temporal consistency in the re-sampling of locations, which is part of the rotational design. The relationship between Ks and near-surface hydrological flow paths is of critical importance when assessing the consequences of land-use change in the humid tropics. The last part of this dissertation aimed at disclosing spatial relationships between Ks and overland flow as influenced by different land cover types. The effects of Ks on overland-flow generation were spatially variable, different between planar plots and incised flowlines and strongly influenced by land-cover characteristics. A simple comparison of Ks values and rainfall intensities was insufficient to describe the observed pattern of overland flow. Likewise, event flow in the stream was apparently not directly related to overland flow response patterns within the catchments. The study emphasises the importance of combining pedological, hydrological, meteorological and botanical measurements to comprehensively understand the land use-driven change in hydrological flow paths. In summary, Ks proved to be a suitable parameter for assessing the influence of land-use change on soils and hydrological processes. The results illustrated the importance of land cover and spatial variability of Ks for decisions on sampling designs and for interpreting overland-flow generation. As relationships between Ks and overland flow were shown to be complex and dependent on land cover, an interdisciplinary approach is required to comprehensively understand the effects of land-use change on soils and near-surface hydrological flow paths in the humid tropics.
Das Selenoprotein Glutathionperoxidase 2 (GPx2) ist ein epithelzellspezifisches, Hydroperoxide-reduzierendes Enzym, welches im Darmepithel, vor allem in den proliferierenden Zellen des Kryptengrundes, exprimiert wird. Die Aufrechterhaltung der GPx2-Expression im Kryptengrund auch bei subadäquatem Selenstatus könnte darauf hinweisen, dass sie hier besonders wichtige Funktionen wahrnimmt. Tatsächlich weisen GPx2 knockout (KO)-Mäuse eine erhöhte Apoptoserate im Kryptengrund auf. Ein Ziel dieser Arbeit war es deshalb, die physiologische Funktion der GPx2 näher zu untersuchen. In Kryptengrundepithelzellen aus dem Colon selenarmer GPx2 KO-Mäuse wurde eine erhöhte Caspase 3/7-Aktivität im Vergleich zum Wildtyp (WT) festgestellt. Zudem wiesen diese Zellen eine erhöhte Suszeptibilität für oxidativen Stress auf. Die GPx2 gewährleistet also den Schutz der proliferierenden Zellen des Kryptengrundes auch bei subadäquater Selenversorgung. Des Weiteren wurde im Colon selenarmer (-Se) und -adäquater (+Se) GPx2 KO-Mäuse im Vergleich zum WT eine erhöhte Tumornekrosefaktor α-Expression und eine erhöhte Infiltration von Makrophagen festgestellt. Durch Fütterung einer selensupplementierten Diät (++Se) konnte dies verhindert werden. In GPx2 KO-Mäusen liegt demnach bereits basal eine niedriggradige Entzündung vor. Dies unterstreicht, dass GPx2 vor allem eine wichtige antiinflammatorische Funktion im Darmepithel besitzt. Dem Mikronährstoff Selen werden protektive Funktionen in der Colonkanzerogenese zugeschrieben. In einem Mausmodell der Colitis-assoziierten Colonkanzerogenese wirkte GPx2 antiinflammatorisch und hemmte so die Tumorentstehung. Auf der anderen Seite wurden jedoch auch prokanzerogene Eigenschaften der GPx2 aufgedeckt. Deshalb sollte in dieser Arbeit untersucht werden, welchen Effekt ein GPx2 knockout in einem Modell der sporadischen, durch Azoxymethan (AOM) induzierten, Colonkanzerogenese hat. Im WT kam es in präneoplastischen Läsionen häufig zu einer erhöhten GPx2-Expression im Vergleich zur normalen Darmmucosa. Eine derartige Steigerung der GPx2-Expression wurde auch in der humanen Colonkanzerogenese beschrieben. Das Fehlen der GPx2 resultierte in einer verminderten Entstehung von Tumoren (-Se und ++Se) und präneoplastischen Läsionen (-Se und +Se). Somit förderte GPx2 die Tumorentstehung im AOM-Modell. Acht Stunden nach AOM-Gabe war im GPx2 KO-Colon im Vergleich zum WT eine erhöhte Apoptoserate in der Kryptenmitte (-Se, +Se), nicht jedoch im Kryptengrund oder in der ++Se-Gruppe zu beobachten. Möglicherweise wirkte GPx2 prokanzerogen, indem sie die effiziente Elimination geschädigter Zellen in der Tumorinitiationsphase verhinderte. Eine ähnliche Wirkung wäre auch durch die erhöhte GPx2-Expression in der Promotionsphase denkbar. So könnte GPx2 proliferierende präneoplastische Zellen vor oxidativem Stress, Apoptosen, oder auch der Antitumorimmunität schützen. Dies könnte durch ein Zusammenwirken mit anderen Selenoproteinen wie GPx1 und Thioredoxinreduktasen, für die ebenfalls auch prokanzerogene Funktionen beschrieben wurden, verstärkt werden. Eine wichtige Rolle könnte hier die Modulation des Redoxstatus in Tumorzellen spielen. Die Variation des Selengehalts der Diät hatte im WT einen eher U-förmigen Effekt. So traten in der –Se und ++Se-Gruppe tendenziell mehr und größere Tumore auf, als in der +Se Gruppe. Zusammenfassend schützt GPx2 also die proliferierenden Zellen des Kryptengrundes. Sie könnte jedoch auch proliferierende transformierte Zellen schützen und so die sporadische, AOM-induzierte Colonkanzerogenese fördern. In einem Modell der Colitis-assoziierten Colonkanzerogenese hatte GPx2 auf Grund ihrer antiinflammatorischen Wirkung einen gegenteiligen Effekt und hemmte die Tumorentstehung. Die Rolle der GPx2 in der Colonkanzerogenese ist also abhängig vom zugrunde liegenden Mechanismus und wird maßgeblich von der Beteiligung einer Entzündung bestimmt.
Unter geeigneten Wachstumsbedingungen weisen Algenkulturen oft eine größere Produktivität der Zellen auf, als sie bei höheren Pflanzen zu beobachten ist. Chlamydomonas reinhardtii-Zellen sind vergleichsweise klein. So beträgt das Zellvolumen während des vegetativen Zellzyklus etwa 50–3500 µm³. Im Vergleich zu höheren Pflanzen ist in einer Algensuspension die Konzentration der Biomasse allerdings gering. So enthält beispielsweise 1 ml einer üblichen Konzentration zwischen 10E6 und 10E7 Algenzellen. Quantifizierungen von Metaboliten oder Makromolekülen, die zur Modellierung von zellulären Prozessen genutzt werden, werden meist im Zellensemble vorgenommen. Tatsächlich unterliegt jedoch jede Algenzelle einer individuellen Entwicklung, die die Identifizierung charakteristischer allgemeingültiger Systemparameter erschwert. Ziel dieser Arbeit war es, biochemisch relevante Messgrößen in-vivo und in-vitro mit Hilfe optischer Verfahren zu identifizieren und zu quantifizieren. Im ersten Teil der Arbeit wurde ein Puls-Amplituden-Modulation(PAM)-Fluorimetriemessplatz zur Messung der durch äußere Einflüsse bedingten veränderlichen Chlorophyllfluoreszenz an einzelnen Zellen vorgestellt. Die Verwendung eines kommerziellen Mikroskops, die Implementierung empfindlicher Nachweiselektronik und einer geeignete Immobilisierungsmethode ermöglichten es, ein Signal-zu-Rauschverhältnis zu erreichen, mit dem Fluoreszenzsignale einzelner lebender Chlamydomonas-Zellen gemessen werden konnten. Insbesondere wurden das Zellvolumen und der als Maß für die Effizienz des Photosyntheseapparats bzw. die Zellfitness geltende Chlorophyllfluoreszenzparameter Fv/Fm ermittelt und ein hohes Maß an Heterogenität dieser zellulären Parameter in verschiedenen Entwicklungsstadien der synchronisierten Chlamydomonas-Zellen festgestellt. Im zweiten Teil der Arbeit wurden die bildgebende Laser-Scanning-Mikroskopie und anschließende Bilddatenanalyse zur quantitativen Erfassung der wachstumsabhängigen zellulären Parameter angewandt. Ein kommerzielles konfokales Mikroskop wurde um die Möglichkeit der nichtlinearen Mikroskopie erweitert. Diese hat den Vorteil einer lokalisierten Anregung, damit verbunden einer höheren Ortsauflösung und insgesamt geringeren Probenbelastung. Weiterhin besteht neben der Signalgewinnung durch Fluoreszenzanregung die Möglichkeit der Erzeugung der Zweiten Harmonischen (SHG) an biophotonischen Strukturen, wie der zellulären Stärke. Anhand der Verteilungsfunktionen war es möglich mit Hilfe von modelltheoretischen Ansätzen zelluläre Parameter zu ermitteln, die messtechnisch nicht unmittelbar zugänglich sind. Die morphologischen Informationen der Bilddaten ermöglichten die Bestimmung der Zellvolumina und die Volumina subzellularer Strukturen, wie Nuclei, extranucleäre DNA oder Stärkegranula. Weiterhin konnte die Anzahl subzellulärer Strukturen innerhalb einer Zelle bzw. eines Zellverbunds ermittelt werden. Die Analyse der in den Bilddaten enthaltenen Signalintensitäten war Grundlage einer relativen Konzentrationsbestimmung von zellulären Komponenten, wie DNA bzw. Stärke. Mit dem hier vorgestellten Verfahren der nichtlinearen Mikroskopie und nachfolgender Bilddatenanalyse konnte erstmalig die Verteilung des zellulären Stärkegehalts in einer Chlamydomonas-Population während des Wachstums bzw. nach induziertem Stärkeabbau verfolgt werden. Im weiteren Verlauf wurde diese Methode auch auf Gefrierschnitte höherer Pflanzen, wie Arabidopsis thaliana, angewendet. Im Ergebnis wurde gezeigt, dass viele zelluläre Parameter, wie das Volumen, der zelluläre DNA- und Stärkegehalt bzw. die Anzahl der Stärkegranula durch eine Lognormalverteilung, mit wachstumsabhängiger Parametrisierung, beschrieben werden. Zelluläre Parameter, wie Stoffkonzentration und zelluläres Volumen, zeigen keine signifikanten Korrelationen zueinander, woraus geschlussfolgert werden muss, dass es ein hohes Maß an Heterogenität der zellulären Parameter innerhalb der synchronisierten Chlamydomonas-Populationen gibt. Diese Aussage gilt sowohl für die als homogenste Form geltenden Synchronkulturen von Chlamydomonas reinhardtii als auch für die gemessenen zellulären Parameter im intakten Zellverbund höherer Pflanzen. Dieses Ergebnis ist insbesondere für modelltheoretische Betrachtungen von Relevanz, die sich auf empirische Daten bzw. zelluläre Parameter stützen welche im Zellensemble gemessen wurden und somit nicht notwendigerweise den zellulären Status einer einzelnen Zelle repräsentieren.