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Plants can be primed to survive the exposure to a severe heat stress (HS) by prior exposure to a mild HS. The information about the priming stimulus is maintained by the plant for several days. This maintenance of acquired thermotolerance, or HS memory, is genetically separable from the acquisition of thermotolerance itself and several specific regulatory factors have been identified in recent years.
On the molecular level, HS memory correlates with two types of transcriptional memory, type I and type II, that characterize a partially overlapping subset of HS-inducible genes. Type I transcriptional memory or sustained induction refers to the sustained transcriptional induction above non-stressed expression levels of a gene for a prolonged time period after the end of the stress exposure. Type II transcriptional memory refers to an altered transcriptional response of a gene after repeated exposure to a stress of similar duration and intensity. In particular, enhanced re-induction refers to a transcriptional pattern in which a gene is induced to a significantly higher degree after the second stress exposure than after the first.
This thesis describes the functional characterization of a novel positive transcriptional regulator of type I transcriptional memory, the heat shock transcription factor HSFA3, and compares it to HSFA2, a known positive regulator of type I and type II transcriptional memory. It investigates type I transcriptional memory and its dependence on HSFA2 and HSFA3 for the first time on a genome-wide level, and gives insight on the formation of heteromeric HSF complexes in response to HS. This thesis confirms the tight correlation between transcriptional memory and H3K4 hyper-methylation, reported here in a case study that aimed to reduce H3K4 hyper-methylation of the type II transcriptional memory gene APX2 by CRISPR/dCas9-mediated epigenome editing. Finally, this thesis gives insight into the requirements for a heat shock transcription factor to function as a positive regulator of transcriptional memory, both in terms of its expression profile and protein abundance after HS and the contribution of individual functional domains.
In summary, this thesis contributes to a more detailed understanding of the molecular processes underlying transcriptional memory and therefore HS memory, in Arabidopsis thaliana.
Duplicate detection describes the process of finding multiple representations of the same real-world entity in the absence of a unique identifier, and has many application areas, such as customer relationship management, genealogy and social sciences, or online shopping. Due to the increasing amount of data in recent years, the problem has become even more challenging on the one hand, but has led to a renaissance in duplicate detection research on the other hand.
This thesis examines the effects and opportunities of transitive relationships on the duplicate detection process. Transitivity implies that if record pairs ⟨ri,rj⟩ and ⟨rj,rk⟩ are classified as duplicates, then also record pair ⟨ri,rk⟩ has to be a duplicate. However, this reasoning might contradict with the pairwise classification, which is usually based on the similarity of objects. An essential property of similarity, in contrast to equivalence, is that similarity is not necessarily transitive.
First, we experimentally evaluate the effect of an increasing data volume on the threshold selection to classify whether a record pair is a duplicate or non-duplicate. Our experiments show that independently of the pair selection algorithm and the used similarity measure, selecting a suitable threshold becomes more difficult with an increasing number of records due to an increased probability of adding a false duplicate to an existing cluster. Thus, the best threshold changes with the dataset size, and a good threshold for a small (possibly sampled) dataset is not necessarily a good threshold for a larger (possibly complete) dataset. As data grows over time, earlier selected thresholds are no longer a suitable choice, and the problem becomes worse for datasets with larger clusters.
Second, we present with the Duplicate Count Strategy (DCS) and its enhancement DCS++ two alternatives to the standard Sorted Neighborhood Method (SNM) for the selection of candidate record pairs. DCS adapts SNMs window size based on the number of detected duplicates and DCS++ uses transitive dependencies to save complex comparisons for finding duplicates in larger clusters. We prove that with a proper (domain- and data-independent!) threshold, DCS++ is more efficient than SNM without loss of effectiveness.
Third, we tackle the problem of contradicting pairwise classifications. Usually, the transitive closure is used for pairwise classifications to obtain a transitively closed result set. However, the transitive closure disregards negative classifications. We present three new and several existing clustering algorithms and experimentally evaluate them on various datasets and under various algorithm configurations. The results show that the commonly used transitive closure is inferior to most other clustering algorithms, especially for the precision of results. In scenarios with larger clusters, our proposed EMCC algorithm is, together with Markov Clustering, the best performing clustering approach for duplicate detection, although its runtime is longer than Markov Clustering due to the subexponential time complexity. EMCC especially outperforms Markov Clustering regarding the precision of the results and additionally has the advantage that it can also be used in scenarios where edge weights are not available.
Isometric muscle function
(2022)
The cumulative dissertation consists of four original articles. These considered isometric muscle ac-tions in healthy humans from a basic physiological view (oxygen and blood supply) as well as possibilities of their distinction. It includes a novel approach to measure a specific form of isometric hold-ing function which has not been considered in motor science so far. This function is characterized by an adaptation to varying external forces with particular importance in daily activities and sports.
The first part of the research program analyzed how the biceps brachii muscle is supplied with oxygen and blood by adapting to a moderate constant load until task failure (publication 1). In this regard, regulative mechanisms were investigated in relation to the issue of presumably compressed capillaries due to high intramuscular pressures (publication 2).
Furthermore, it was examined if oxygenation and time to task failure (TTF) differs compared to an-other isometric muscle function (publication 3). This function is mainly of diagnostic interest by measuring the maximal voluntary isometric contraction (MVIC) as a gold standard. For that, a person pulls on or pushes against an insurmountable resistance. However, the underlying pulling or pushing form of isometric muscle action (PIMA) differs compared to the holding one (HIMA).
HIMAs have mainly been examined by using constant loads. In order to quantify the adaptability to varying external forces, a new approach was necessary and considered in the second part of the research program. A device was constructed based on a previously developed pneumatic measurement system. The device should have been able to measure the Adaptive Force (AF) of elbow ex-tensor muscles. The AF determines the adaptability to increasing external forces under isometric (AFiso) and eccentric (AFecc) conditions. At first, it was questioned if these parameters can be relia-bly assessed by use of the new device (publication 4). Subsequently, the main research question was investigated: Is the maximal AFiso a specific and independent variable of muscle function in comparison to the MVIC? Furthermore, both research parts contained a sub-question of how results can be influenced.
Parameters of local oxygen saturation (SvO2) and capillary blood filling (rHb) were non-invasively recorded by a spectrophotometer during maximal and submaximal HIMAs and PIMAs.
These were the main findings: Under load, SvO2 and rHb always adjusted into a steady state after an initial decrease. Nevertheless, their behavior could roughly be categorized into two types. In type I, both parameters behaved nearly parallel to each other. In contrast, their progression over time was partly inverse in type II. The inverse behavior probably depends on the level of deoxygenation since rHb increased reliably at a suggested threshold of about 59% SvO2. This triggered mechanism and the found homeostatic steady states seem to be in conflict with the concept of mechanically compressed capillaries and consequently with a restricted blood flow. Anatomical configuration of blood vessels might provide one hypothetical explanation of how blood flow might be maintained. HIMA and PIMA did not differ regarding oxygenation and allocation to the described types. The TTF tended to be longer during PIMA.
As a sub-question, oxygenation and TTF were compared between (HIMA) and intermittent voluntary muscle twitches during a weight holding task. TTF but not oxygenation differed significantly
(Twitch > HIMA). A changed neuromuscular control might serve as a speculative explanation of how the results can be explained. This is supported by the finding that the TTF did not correlate significantly with the extent of deoxygenation irrespective of the performed task (HIMA, PIMA or Twitch).
Other neuromuscular aspects of muscle function were considered in second part of the re-search program. The new device mentioned above detected different force capacities within four trials at two days each. Among AF measurements, the functional counterpart of a concentric muscle action merging into an isometric one was analyzed in comparison to the MVIC.
Based on the results, it can be assumed that a prior concentric muscle action does not influence the MVIC. However, the results were inconsistent and possibly influenced by systematic errors. In con-trast, maximal variables of the AF (AFisomax and AFeccmax) could be measured in a reliable way which is indicated by a high test-retest reliability. Despite substantial correlations between force variables, the AFisomax differed significantly from MVIC and AFmax, which was identical with AFeccmax in almost all cases. Moreover, AFisomax revealed the highest variability between trials.
These results indicate that maximal force capacities should be assessed separately. The adaptive holding capacity of a muscle can be lower compared to a commonly determined MVIC. This is of relevance since muscles frequently need to respond adequately to external forces. If their response does not correspond to the external impact, the muscle is forced to lengthen. In this scenario, joints are not completely stabilized and an injury may occur. This outlined issue should be addressed in future research in the field of sport and health sciences.
At last, the dissertation presents another possibility to quantify the AFisomax by use of a handheld device applied in combination with a manual muscle test. This assessment delivers a more practical way for clinical purposes.
Stimuli-promoted in situ formation of hydrogels with thiol/thioester containing peptide precursors
(2022)
Hydrogels are potential synthetic ECM-like substitutes since they provide functional and structural similarities compared to soft tissues. They can be prepared by crosslinking of macromolecules or by polymerizing suitable precursors. The crosslinks are not necessarily covalent bonds, but could also be formed by physical interactions such as π-π interactions, hydrophobic interactions, or H-bonding. On demand in situ forming hydrogels have garnered increased interest especially for biomedical applications over preformed gels due to the relative ease of in vivo delivery and filling of cavities. The thiol-Michael addition reaction provides a straightforward and robust strategy for in situ gel formation with its fast reaction kinetics and ability to proceed under physiological conditions. The incorporation of a trigger function into a crosslinking system becomes even more interesting since gelling can be controlled with stimulus of choice. The use of small molar mass crosslinker precursors with active groups orthogonal to thiol-Michael reaction type electrophile provides the opportunity to implement an on-demand in situ crosslinking without compromising the fast reaction kinetics.
It was postulated that short peptide sequences due to the broad range structural-function relations available with the different constituent amino acids, can be exploited for the realisation of stimuli-promoted in situ covalent crosslinking and gelation applications. The advantages of this system over conventional polymer-polymer hydrogel systems are the ability tune and predict material property at the molecular level.
The main aim of this work was to develop a simplified and biologically-friendly stimuli-promoted in situ crosslinking and hydrogelation system using peptide mimetics as latent crosslinkers. The approach aims at using a single thiodepsipeptide sequence to achieve separate pH- and enzyme-promoted gelation systems with little modification to the thiodepsipeptide sequence. The realization of this aim required the completion of three milestones.
In the first place, after deciding on the thiol-Michael reaction as an effective in situ crosslinking strategy, a thiodepsipeptide, Ac-Pro-Leu-Gly-SLeu-Leu-Gly-NEtSH (TDP) with expected propensity towards pH-dependent thiol-thioester exchange (TTE) activation, was proposed as a suitable crosslinker precursor for pH-promoted gelation system. Prior to the synthesis of the proposed peptide-mimetic, knowledge of the thiol-Michael reactivity of the would-be activated thiol moiety SH-Leu, which is internally embedded in the thiodepsipeptide was required. In line with pKa requirements for a successful TTE, the reactivity of a more acidic thiol, SH-Phe was also investigated to aid the selection of the best thiol to be incorporated in the thioester bearing peptide based crosslinker precursor. Using ‘pseudo’ 2D-NMR investigations, it was found that only reactions involving SH-Leu yielded the expected thiol-Michael product, an observation that was attributed to the steric hindrance of the bulkier nature of SH-Phe. The fast reaction rates and complete acrylate/maleimide conversion obtained with SH-Leu at pH 7.2 and higher aided the direct elimination of SH-Phe as a potential thiol for the synthesis of the peptide mimetic.
Based on the initial studies, for the pH-promoted gelation system, the proposed Ac-Pro-Leu-Gly-SLeu-Leu-Gly-NEtSH was kept unmodified. The subtle difference in pKa values between SH-Leu (thioester thiol) and the terminal cysteamine thiol from theoretical conditions should be enough to effect a ‘pseudo’ intramolecular TTE. In polar protic solvents and under basic aqueous conditions, TDP successfully undergoes a ‘pseudo’ intramolecular TTE reaction to yield an α,ω-dithiol tripeptide, HSLeu-Leu-Gly-NEtSH. The pH dependence of thiolate ion generation by the cysteamine thiol aided the incorporation of the needed stimulus (pH) for the overall success of TTE (activation step) – thiol-Michael addition (crosslinking) strategy.
Secondly, with potential biomedical applications in focus, the susceptibility of TDP, like other thioesters, to intermolecular TTE reaction was probed with a group of thiols of varying thiol pKa values, since biological milieu characteristically contain peptide/protein thiols. L-cysteine, which is a biologically relevant thiol, and a small molecular weight thiol, methylthioglycolate both with relatively similar thiol pKa, values, led to an increase concentration of the dithiol crosslinker when reacted with TDP. In the presence of acidic thiols (p-NTP and 4MBA), a decrease in the dithiol concentration was observed, an observation that can be attributed to the inability of the TTE tetrahedral intermediate to dissociate into exchange products and is in line with pKa requirements for successful TTE reaction. These results additionally makes TDP more attractive and the potentially the first crosslinker precursor for applications in biologically relevant media.
Finally, the ability of TDP to promote pH-sensitive in situ gel formation was probed with maleimide functionalized 4-arm polyethylene glycol polymers in tris-buffered media of varying pHs. When a 1:1 thiol: maleimide molar ratio was used, TDP-PEG4MAL hydrogels formed within 3, 12 and 24 hours at pH values of 8.5, 8.0 and 7.5 respectively. However, gelation times of 3, 5 and 30 mins were observed for the same pH trend when the thiol: maleimide molar was increased to 2:1.
A direct correlation of thiol content with G’ of the gels at each pH could also be drawn by comparing gels with thiol: maleimide ratios of 1:1 to those with 2:1 thiol: maleimide mole ratios. This is supported by the fact that the storage modulus (G') is linearly dependent on the crosslinking density of the polymer. The values of initial G′ for all gels ranged between (200 – 5000 Pa), which falls in the range of elasticities of certain tissue microenvironments for example brain tissue 200 – 1000 Pa and adipose tissue (2500 – 3500 Pa).
Knowledge so far gained from the study on the ability to design and tune the exchange reaction of thioester containing peptide mimetic will give those working in the field further insight into the development of new sequences tailored towards specific applications.
TTE substrate design using peptide mimetic as presented in this work has revealed interesting new insights considering the state-of-the-art. Using the results obtained as reference, the strategy provides a possibility to extend the concept to the controlled delivery of active molecules needed for other robust and high yielding crosslinking reactions for biomedical applications. Application for this sequentially coupled functional system could be seen e.g. in the treatment of inflamed tissues associated with urinary tract like bladder infections for which pH levels above 7 were reported. By the inclusion of cell adhesion peptide motifs, the hydrogel network formed at this pH could act as a new support layer for the healing of damage epithelium as shown in interfacial gel formation experiments using TDP and PEG4MAL droplets.
The versatility of the thiodepsipeptide sequence, Ac-Pro-Leu-Gly-SLeu-Leu-Gly-(TDPo) was extended for the design and synthesis of a MMP-sensitive 4-arm PEG-TDPo conjugate. The purported cleavage of TDPo at the Gly-SLeu bond yields active thiol units for subsequent reaction of orthogonal Michael acceptor moieties. One of the advantages of stimuli-promoted in situ crosslinking systems using short peptides should be the ease of design of required peptide molecules due to the predictability of peptide functions their sequence structure. Consequently the functionalisation of a 4-arm PEG core with the collagenase active TDPo sequence yielded an MMP-sensitive 4-arm thiodepsipeptide-PEG conjugate (PEG4TDPo) substrate.
Cleavage studies using thiol flourometric assay in the presence of MMPs -2 and -9 confirmed the susceptibility of PEG4TDPo towards these enzymes. The resulting time-dependent increase in fluorescence intensity in the presence of thiol assay signifies the successful cleavage of TDPo at the Gly-SLeu bond as expected. It was observed that the cleavage studies with thiol flourometric assay introduces a sigmoid non-Michaelis-Menten type kinetic profile, hence making it difficult to accurately determine the enzyme cycling parameters, kcat and KM .
Gelation studies with PEG4MAL at 10 % wt. concentrations revealed faster gelation with MMP-2 than MMP-9 with 28 and 40 min gelation times respectively. Possible contributions by hydrolytic cleavage of PEG4TDPo has resulted in the gelation of PEG4MAL blank samples but only after 60 minutes of reaction. From theoretical considerations, the simultaneous gelation reaction would be expected to more negatively impact the enzymatic than hydrolytic cleavage. The exact contributions from hydrolytic cleavage of PEG4TDPo would however require additional studies.
In summary this new and simplified in situ crosslinking system using peptide-based crosslinker precursors with tuneable properties exhibited in situ crosslinking gelation kinetics on similar levels with already active dithiols reported. The advantageous on-demand functionality associated with its pH-sensitivity and physiological compatibility makes it a strong candidate worth further research as biomedical applications in general and on-demand material synthesis is concerned.
Results from MMP-promoted gelation system unveils a simple but unexplored approach for in situ synthesis of covalently crosslinked soft materials, that could lead to the development of an alternative pathway in addressing cancer metastasis by making use of MMP overexpression as a trigger. This goal has so far not being reach with MMP inhibitors despite the extensive work this regard.
Selbstwirksamkeitserwartungen von Lehramtsstudierenden im Kontext von schulpraktischen Erfahrungen
(2022)
Selbstwirksamkeitserwartungen spielen eine wichtige Rolle für das professionelle Verhalten von Lehrkräften im Unterricht (Tschannen-Moran et al., 1998) sowie für die Leistungen und das Verhalten der Schülerinnen und Schüler (Mojavezi & Tamiz, 2012). Selbstwirksamkeitserwartungen von Lehrkräften sind definiert als die Überzeugung von Lehrkräften, dass sie in der Lage sind, bestimmte Ziele in einer spezifischen Situation zu erreichen (Dellinger et al., 2008; Tschannen-Moran & Hoy, 2001). Aufgrund der bedeutenden Rolle der Lehrkräfte im Bildungssystem und in der Gesellschaft ist es wichtig, das Wohlbefinden, die Produktivität und die Wirksamkeit von Lehrkräften zu fördern (Kasalak & Dagyar, 2020). Empirische Befunde unterstreichen die positiven Effekte von Selbstwirksamkeitserwartungen bei Lehrkräften auf ihr Wohlbefinden (Perera & John, 2020) und auf das Lernen sowie die Leistungen der Schülerinnen und Schüler (Zee & Koomen, 2016). Dabei mangelt es jedoch an empirischer Forschung, die die Bedeutung von Selbstwirksamkeitserwartungen bei Lehramtsstudierende in der Lehrkräftebildung untersucht (Yurekli et al., 2020), insbesondere während schulpraktischen Ausbildungsphasen. Ausgehend von der Bedeutung eigener Unterrichtserfahrungen, die als mastery experience, d.h. als stärkste Quelle von Selbstwirksamkeit für Lehramtsstudierende, beschrieben wurden (Pfitzner-Eden, 2016b), werden in dieser Dissertation Praxiserfahrungen als Quelle von Selbstwirksamkeit von Lehramtsstudierenden und die Veränderung der Selbstwirksamkeit von Lehramtsstudierenden während der Lehrkräfteausbildung untersucht. Studie 1 konzentriert sich daher auf die Veränderung der Selbstwirksamkeit von Lehramtsstudierenden während kurzer praktischer Unterrichtserfahrungen im Vergleich zur Online-Lehre ohne Unterrichtserfahrung. Aufgrund inkonsistenter Befunde zu den wechselseitigen Beziehungen zwischen den Selbstwirksamkeitserwartungen von Lehrkräften und ihrem Unterrichtsverhalten (Holzberger et al., 2013; Lazarides et al., 2022) wurde in Studie 2 der Zusammenhang zwischen der Selbstwirksamkeit von Lehramtsstudierenden und ihrem Unterrichtsverhalten während des Lehramtsstudiums untersucht. Da Feedback als verbale Überzeugung (verbal persuasion) dienen kann und somit eine wichtige Quelle für Selbstwirksamkeitserwartungen ist, die das Gefühl der Kompetenz stärkt (Pfitzner-Eden, 2016b), fokussiert Studie 2 den Zusammenhang zwischen der Veränderung der Selbstwirksamkeit von Lehramtsstudierenden und der wahrgenommenen Qualität des Peer-Feedbacks im Kontext kurzer schulpraktischer Erfahrungen während des Lehramtsstudiums. Darüber hinaus ist es für die Untersuchung der Veränderung von Selbstwirksamkeit bei Lehramtsstudierenden wichtig, individuelle Persönlichkeitsaspekte und spezifische Bedingungen der Lernumgebung in der Lehrkräftebildung zu untersuchen (Bach, 2022). Ausgehend von der Annahme, dass die Unterstützung von Reflexionsprozessen in der Lehrkräftebildung (Menon & Azam, 2021) und der Einsatz innovativer Lernsettings wie VR-Videos (Nissim & Weissblueth, 2017) die Entwicklung von Selbstwirksamkeitserwartungen von Lehramtsstudierenden fördern, werden in Studie 3 und Studie 4 Reflexionsprozesse bei Lehramtsstudierenden in Bezug auf ihre eigenen Unterrichtserfahrungen bzw. stellvertretenden Unterrichtserfahrungen anderer untersucht. Vor dem Hintergrund inkonsistenter Befunde und fehlender empirischer Forschung zu den Zusammenhängen zwischen Selbstwirksamkeit von Lehramtsstudierenden und verschiedenen Faktoren, die das Lernumfeld oder persönliche Merkmale betreffen, sind weitere empirische Studien erforderlich, die verschiedene Quellen und Zusammenhänge der Selbstwirksamkeitserwartungen von Lehramtsstudierenden während des Lehramtsstudiums untersuchen. In diesem Zusammenhang wird in der vorliegenden Dissertation der Frage nachgegangen, welche individuellen Merkmale und Lernumgebungen die Selbstwirksamkeit von Lehramtsstudierenden – insbesondere während kurzer schulpraktischer Phasen im Lehramtsstudium fördern können. Darüber hinaus schließt die Dissertation mit der Diskussion der Ergebnisse aus den vier Teilstudien ab, indem Stärken und Schwächen jeder Studie gesamtheitlich in den Blick genommen werden. Abschließend werden Limitationen und Implikationen für die weitere Forschung und die Praxis diskutiert.
Polyglot programming allows developers to use multiple programming languages within the same software project. While it is common to use more than one language in certain programming domains, developers also apply polyglot programming for other purposes such as to re-use software written in other languages. Although established approaches to polyglot programming come with significant limitations, for example, in terms of performance and tool support, developers still use them to be able to combine languages.
Polyglot virtual machines (VMs) such as GraalVM provide a new level of polyglot programming, allowing languages to directly interact with each other. This reduces the amount of glue code needed to combine languages, results in better performance, and enables tools such as debuggers to work across languages. However, only a little research has focused on novel tools that are designed to support developers in building software with polyglot VMs. One reason is that tool-building is often an expensive activity, another one is that polyglot VMs are still a moving target as their use cases and requirements are not yet well understood.
In this thesis, we present an approach that builds on existing self-sustaining programming systems such as Squeak/Smalltalk to enable exploratory programming, a practice for exploring and gathering software requirements, and re-use their extensive tool-building capabilities in the context of polyglot VMs. Based on TruffleSqueak, our implementation for the GraalVM, we further present five case studies that demonstrate how our approach helps tool developers to design and build tools for polyglot programming. We further show that TruffleSqueak can also be used by application developers to build and evolve polyglot applications at run-time and by language and runtime developers to understand the dynamic behavior of GraalVM languages and internals. Since our platform allows all these developers to apply polyglot programming, it can further help to better understand the advantages, use cases, requirements, and challenges of polyglot VMs. Moreover, we demonstrate that our approach can also be applied to other polyglot VMs and that insights gained through it are transferable to other programming systems.
We conclude that our research on tools for polyglot programming is an important step toward making polyglot VMs more approachable for developers in practice. With good tool support, we believe polyglot VMs can make it much more common for developers to take advantage of multiple languages and their ecosystems when building software.
High-mountain regions provide valuable ecosystem services, including food, water, and energy production, to more than 900 million people worldwide. Projections hold, that this population number will rapidly increase in the next decades, accompanied by a continued urbanisation of cities located in mountain valleys. One of the manifestations of this ongoing socio-economic change of mountain societies is a rise in settlement areas and transportation infrastructure while an increased power need fuels the construction of hydropower plants along rivers in the high-mountain regions of the world. However, physical processes governing the cryosphere of these regions are highly sensitive to changes in climate and a global warming will likely alter the conditions in the headwaters of high-mountain rivers. One of the potential implications of this change is an increase in frequency and magnitude of outburst floods – highly dynamic flows capable of carrying large amounts of water and sediments. Sudden outbursts from lakes formed behind natural dams are complex geomorphological processes and are often part of a hazard cascade. In contrast to other types of natural hazards in high-alpine areas, for example landslides or avalanches, outburst floods are highly infrequent. Therefore, observations and data describing for example the mode of outburst or the hydraulic properties of the downstream propagating flow are very limited, which is a major challenge in contemporary (glacial) lake outburst flood research. Although glacial lake outburst floods (GLOFs) and landslide-dammed lake outburst floods (LLOFs) are rare, a number of documented events caused high fatality counts and damage. The highest documented losses due to outburst floods since the start of the 20th century were induced by only a few high-discharge events. Thus, outburst floods can be a significant hazard to downvalley communities and infrastructure in high-mountain regions worldwide.
This thesis focuses on the Greater Himalayan region, a vast mountain belt stretching across 0.89 million km2. Although potentially hundreds of outburst floods have occurred there since the beginning of the 20th century, data on these events is still scarce. Projections of cryospheric change, including glacier-mass wastage and permafrost degradation, will likely result in an overall increase of the water volume stored in meltwater lakes as well as the destabilisation of mountain slopes in the Greater Himalayan region. Thus, the potential for outburst floods to affect the increasingly more densely populated valleys of this mountain belt is also likely to increase in the future. A prime example of one of these valleys is the Pokhara valley in Nepal, which is drained by the Seti Khola, a river crossing one of the steepest topographic gradients in the Himalayas. This valley is also home to Nepal’s second largest, rapidly growing city, Pokhara, which currently has a population of more than half a million people – some of which live in informal settlements within the floodplain of the Seti Khola. Although there is ample evidence for past outburst floods along this river in recent and historic times, these events have hardly been quantified.
The main motivation of my thesis is to address the data scarcity on past and potential future outburst floods in the Greater Himalayan region, both at a regional and at a local scale. For the former, I compiled an inventory of >3,000 moraine-dammed lakes, of which about 1% had a documented sudden failure in the past four decades. I used this data to test whether a number of predictors that have been widely applied in previous GLOF assessments are statistically relevant when estimating past GLOF susceptibility. For this, I set up four Bayesian multi-level logistic regression models, in which I explored the credibility of the predictors lake area, lake-area dynamics, lake elevation, parent-glacier-mass balance, and monsoonality. By using a hierarchical approach consisting of two levels, this probabilistic framework also allowed for spatial variability on GLOF susceptibility across the vast study area, which until now had not been considered in studies of this scale. The model results suggest that in the Nyainqentanglha and Eastern Himalayas – regions with strong negative glacier-mass balances – lakes have been more prone to release GLOFs than in regions with less negative or even stable glacier-mass balances. Similarly, larger lakes in larger catchments had, on average, a higher probability to have had a GLOF in the past four decades. Yet, monsoonality, lake elevation, and lake-area dynamics were more ambiguous. This challenges the credibility of a lake’s rapid growth in surface area as an indicator of a pending outburst; a metric that has been applied to regional GLOF assessments worldwide.
At a local scale, my thesis aims to overcome data scarcity concerning the flow characteristics of the catastrophic May 2012 flood along the Seti Khola, which caused 72 fatalities, as well as potentially much larger predecessors, which deposited >1 km³ of sediment in the Pokhara valley between the 12th and 14th century CE. To reconstruct peak discharges, flow depths, and flow velocities of the 2012 flood, I mapped the extents of flood sediments from RapidEye satellite imagery and used these as a proxy for inundation limits. To constrain the latter for the Mediaeval events, I utilised outcrops of slackwater deposits in the fills of tributary valleys. Using steady-state hydrodynamic modelling for a wide range of plausible scenarios, from meteorological (1,000 m³ s-1) to cataclysmic outburst floods (600,000 m³ s-1), I assessed the likely initial discharges of the recent and the Mediaeval floods based on the lowest mismatch between sedimentary evidence and simulated flood limits. One-dimensional HEC-RAS simulations suggest, that the 2012 flood most likely had a peak discharge of 3,700 m³ s-1 in the upper Seti Khola and attenuated to 500 m³ s-1 when arriving in Pokhara’s suburbs some 15 km downstream.
Simulations of flow in two-dimensions with orders of magnitude higher peak discharges in ANUGA show extensive backwater effects in the main tributary valleys. These backwater effects match the locations of slackwater deposits and, hence, attest for the flood character of Mediaeval sediment pulses. This thesis provides first quantitative proof for the hypothesis, that the latter were linked to earthquake-triggered outbursts of large former lakes in the headwaters of the Seti Khola – producing floods with peak discharges of >50,000 m³ s-1.
Building on this improved understanding of past floods along the Seti Khola, my thesis continues with an analysis of the impacts of potential future outburst floods on land cover, including built-up areas and infrastructure mapped from high-resolution satellite and OpenStreetMap data. HEC-RAS simulations of ten flood scenarios, with peak discharges ranging from 1,000 to 10,000 m³ s-1, show that the relative inundation hazard is highest in Pokhara’s north-western suburbs. There, the potential effects of hydraulic ponding upstream of narrow gorges might locally sustain higher flow depths. Yet, along this reach, informal settlements and gravel mining activities are close to the active channel. By tracing the construction dynamics in two of these potentially affected informal settlements on multi-temporal RapidEye, PlanetScope, and Google Earth imagery, I found that exposure increased locally between three- to twentyfold in just over a decade (2008 to 2021).
In conclusion, this thesis provides new quantitative insights into the past controls on the susceptibility of glacial lakes to sudden outburst at a regional scale and the flow dynamics of propagating flood waves released by past events at a local scale, which can aid future hazard assessments on transient scales in the Greater Himalayan region. My subsequent exploration of the impacts of potential future outburst floods to exposed infrastructure and (informal) settlements might provide valuable inputs to anticipatory assessments of multiple risks in the Pokhara valley.
The Greenland Ice Sheet is the second-largest mass of ice on Earth. Being almost 2000 km long, more than 700 km wide, and more than 3 km thick at the summit, it holds enough ice to raise global sea levels by 7m if melted completely. Despite its massive size, it is particularly vulnerable to anthropogenic climate change: temperatures over the Greenland Ice Sheet have increased by more than 2.7◦C in the past 30 years, twice as much as the global mean temperature. Consequently, the ice sheet has been significantly losing mass since the 1980s and the rate of loss has increased sixfold since then. Moreover, it is one of the potential tipping elements of the Earth System, which might undergo irreversible change once a warming threshold is exceeded. This thesis aims at extending the understanding of the resilience of the Greenland Ice Sheet against global warming by analyzing processes and feedbacks relevant to its centennial to multi-millennial stability using ice sheet modeling.
One of these feedbacks, the melt-elevation-feedback is driven by the temperature rise with decreasing altitudes: As the ice sheet melts, its thickness and surface elevation decrease, exposing the ice surface to warmer air and thus increasing the melt rates even further. The glacial isostatic adjustment (GIA) can partly mitigate this melt-elevation feedback as the bedrock lifts in response to an ice load decrease, forming the negative GIA feedback. In my thesis, I show that the interaction between these two competing feedbacks can lead to qualitatively different dynamical responses of the Greenland Ice Sheet to warming – from permanent loss to incomplete recovery, depending on the feedback parameters. My research shows that the interaction of those feedbacks can initiate self-sustained oscillations of the ice volume while the climate forcing remains constant.
Furthermore, the increased surface melt changes the optical properties of the snow or ice surface, e.g. by lowering their albedo, which in turn enhances melt rates – a process known as the melt-albedo feedback. Process-based ice sheet models often neglect this melt-albedo feedback. To close this gap, I implemented a simplified version of the diurnal Energy Balance Model, a computationally efficient approach that can capture the first-order effects of the melt-albedo feedback, into the Parallel Ice Sheet Model (PISM). Using the coupled model, I show in warming experiments that the melt-albedo feedback almost doubles the ice loss until the year 2300 under the low greenhouse gas emission scenario RCP2.6, compared to simulations where the melt-albedo feedback is neglected,
and adds up to 58% additional ice loss under the high emission scenario RCP8.5. Moreover, I find that the melt-albedo feedback dominates the ice loss until 2300, compared to the melt-elevation feedback.
Another process that could influence the resilience of the Greenland Ice Sheet is the warming induced softening of the ice and the resulting increase in flow. In my thesis, I show with PISM how the uncertainty in Glen’s flow law impacts the simulated response to warming. In a flow line setup at fixed climatic mass balance, the uncertainty in flow parameters leads to a range of ice loss comparable to the range caused by different warming levels.
While I focus on fundamental processes, feedbacks, and their interactions in the first three projects of my thesis, I also explore the impact of specific climate scenarios on the sea level rise contribution of the Greenland Ice Sheet. To increase the carbon budget flexibility, some warming scenarios – while still staying within the limits of the Paris Agreement – include a temporal overshoot of global warming. I show that an overshoot by 0.4◦C increases the short-term and long-term ice loss from Greenland by several centimeters. The long-term increase is driven by the warming at high latitudes, which persists even when global warming is reversed. This leads to a substantial long-term commitment of the sea level rise contribution from the Greenland Ice Sheet.
Overall, in my thesis I show that the melt-albedo feedback is most relevant for the ice loss of the Greenland Ice Sheet on centennial timescales. In contrast, the melt-elevation feedback and its interplay with the GIA feedback become increasingly relevant on millennial timescales. All of these influence the resilience of the Greenland Ice Sheet against global warming, in the near future and on the long term.
Die Arbeit gibt einen Einblick in die Verständigungspraxen bei Stadtführungen mit (ehemaligen) Obdachlosen, die in ihrem Selbstverständnis auf die Herstellung von Verständnis, Toleranz und Anerkennung für von Obdachlosigkeit betroffene Personen zielen. Zunächst wird in den Diskurs des Slumtourismus eingeführt und, angesichts der Vielfalt der damit verbundenen Erscheinungsformen, Slumming als organisierte Begegnung mit sozialer Ungleichheit definiert. Die zentralen Diskurslinien und die darin eingewobenen moralischen Positionen werden nachvollzogen und im Rahmen der eigenommenen wissenssoziologischen Perspektive als Ausdruck einer per se polykontexturalen Praxis re-interpretiert. Slumming erscheint dann als eine organisierte Begegnung von Lebensformen, die sich in einer Weise fremd sind, als dass ein unmittelbares Verstehen unwahrscheinlich erscheint und genau aus diesem Grund auf der Basis von gängigen Interpretationen des Common Sense ausgehandelt werden muss. Vor diesem Hintergrund untersucht die vorliegende Arbeit, wie sich Teilnehmer und Stadtführer über die Erfahrung der Obdachlosigkeit praktisch verständigen und welcher Art das hierüber erzeugte Verständnis für die im öffentlichen Diskurs mit vielfältigen stigmatisierenden Zuschreibungen versehenen Obdachlosen ist. Dabei interessiert besonders, in Bezug auf welche Aspekte der Erfahrung von Obdachlosigkeit ein gemeinsames Verständnis möglich wird und an welchen Stellen dieses an Grenzen gerät. Dazu wurden die Gesprächsverläufe auf neun Stadtführungen mit (ehemaligen) obdachlosen Stadtführern unterschiedlicher Anbieter im deutschsprachigen Raum verschriftlicht und mit dem Verfahren der Dokumentarischen Methode ausgewertet. Die vergleichende Betrachtung der Verständigungspraxen eröffnet nicht zuletzt eine differenzierte Perspektive auf die in den Prozessen der Verständigung immer schon eingewobenen Anerkennungspraktiken. Mit Blick auf die moralische Debatte um organisierte Begegnungen mit sozialer Ungleichheit wird dadurch eine ethische Perspektive angeregt, in deren Zentrum Fragen zur Vermittlungsarbeit stehen.
The increasing introduction of non-native plant species may pose a threat to local biodiversity. However, the basis of successful plant invasion is not conclusively understood, especially since these plant species can adapt to the new range within a short period of time despite impoverished genetic diversity of the starting populations. In this context, DNA methylation is considered promising to explain successful adaptation mechanisms in the new habitat. DNA methylation is a heritable variation in gene expression without changing the underlying genetic information. Thus, DNA methylation is considered a so-called epigenetic mechanism, but has been studied in mainly clonally reproducing plant species or genetic model plants. An understanding of this epigenetic mechanism in the context of non-native, predominantly sexually reproducing plant species might help to expand knowledge in biodiversity research on the interaction between plants and their habitats and, based on this, may enable more precise measures in conservation biology.
For my studies, I combined chemical DNA demethylation of field-collected seed material from predominantly sexually reproducing species and rearing offsping under common climatic conditions to examine DNA methylation in an ecological-evolutionary context. The contrast of chemically treated (demethylated) plants, whose variation in DNA methylation was artificially reduced, and untreated control plants of the same species allowed me to study the impact of this mechanism on adaptive trait differentiation and local adaptation. With this experimental background, I conducted three studies examining the effect of DNA methylation in non-native species along a climatic gradient and also between climatically divergent regions.
The first study focused on adaptive trait differentiation in two invasive perennial goldenrod species, Solidago canadensis sensu latu and S. gigantea AITON, along a climate gradient of more than 1000 km in length in Central Europe. I found population differences in flowering timing, plant height, and biomass in the temporally longer-established S. canadensis, but only in the number of regrowing shoots for S. gigantea. While S. canadensis did not show any population structure, I was able to identify three genetic groups along this climatic gradient in S. gigantea. Surprisingly, demethylated plants of both species showed no change in the majority of traits studied. In the subsequent second study, I focused on the longer-established goldenrod species S. canadensis and used molecular analyses to infer spatial epigenetic and genetic population differences in the same specimens from the previous study. I found weak genetic but no epigenetic spatial variation between populations. Additionally, I was able to identify one genetic marker and one epigenetic marker putatively susceptible to selection. However, the results of this study reconfirmed that the epigenetic mechanism of DNA methylation appears to be hardly involved in adaptive processes within the new range in S. canadensis.
Finally, I conducted a third study in which I reciprocally transplanted short-lived plant species between two climatically divergent regions in Germany to investigate local adaptation at the plant family level. For this purpose, I used four plant families (Amaranthaceae, Asteraceae, Plantaginaceae, Solanaceae) and here I additionally compared between non-native and native plant species. Seeds were transplanted to regions with a distance of more than 600 kilometers and had either a temperate-oceanic or a temperate-continental climate. In this study, some species were found to be maladapted to their own local conditions, both in non-native and native plant species alike. In demethylated individuals of the plant species studied, DNA methylation had inconsistent but species-specific effects on survival and biomass production. The results of this study highlight that DNA methylation did not make a substantial contribution to local adaptation in the non-native as well as native species studied.
In summary, my work showed that DNA methylation plays a negligible role in both adaptive trait variation along climatic gradients and local adaptation in non-native plant species that either exhibit a high degree of genetic variation or rely mainly on sexual reproduction with low clonal propagation. I was able to show that the adaptive success of these non-native plant species can hardly be explained by DNA methylation, but could be a possible consequence of multiple introductions, dispersal corridors and meta-population dynamics. Similarly, my results illustrate that the use of plant species that do not predominantly reproduce clonally and are not model plants is essential to characterize the effect size of epigenetic mechanisms in an ecological-evolutionary context.