ALBERT

Description / Table of Contents: This thesis investigates how the permafrost microbiota responds to global warming. In detail, the constraints behind methane production in thawing permafrost were linked to methanogenic activity, abundance and composition. Furthermore, this thesis offers new insights into microbial adaptions to the changing environmental conditions during global warming. This was assesed by investigating the potential ecological relevant functions encoded by plasmid DNA within the permafrost microbiota. Permafrost of both interglacial and glacial origin spanning the Holocene to the late Pleistocene, including Eemian, were studied during long-term thaw incubations. Furthermore, several permafrost cores of different stratigraphy, soil type and vegetation cover were used to target the main constraints behind methane production during short-term thaw simulations. Short- and long-term incubations simulating thaw with and without the addition of substrate were combined with activity measurements, amplicon and metagenomic sequencing of permanently frozen and seasonally thawed active layer. Combined, it allowed to address the following questions. i) What constraints methane production when permafrost thaws and how is this linked to methanogenic activity, abundance and composition? ii) How does the methanogenic community composition change during long-term thawing conditions? iii) Which potential ecological relevant functions are encoded by plasmid DNA in active layer soils? The major outcomes of this thesis are as follows. i) Methane production from permafrost after long-term thaw simulation was found to be constrained mainly by the abundance of methanogens and the archaeal community composition. Deposits formed during periods of warmer temperatures and increased precipitation, (here represented by deposits from the Late Pleistocene of both interstadial and interglacial periods) were found to respond strongest to thawing conditions and to contain an archaeal community dominated by methanogenic archaea (40% and 100% of all detected archaea). Methanogenic population size and carbon density were identified as main predictors for potential methane production in thawing permafrost in short-term incubations when substrate was sufficiently available. ii) Besides determining the methanogenic activity after long-term thaw, the paleoenvironmental conditions were also found to influence the response of the methanogenic community composition. Substantial shifts within methanogenic community structure and a drop in diversity were observed in deposits formed during warmer periods, but not in deposits from stadials, when colder and drier conditions occurred. Overall, a shift towards a dominance of hydrogenotrophic methanogens was observed in all samples, except for the oldest interglacial deposits from the Eemian, which displayed a potential dominance of acetoclastic methanogens. The Eemian, which is discussed to serve as an analogue to current climate conditions, contained highly active methanogenic communities. However, all potential limitation of methane production after permafrost thaw, it means methanogenic community structure, methanogenic population size, and substrate pool might be overcome after permafrost had thawed on the long-term. iii) Enrichments with soil from the seasonally thawed active layer revealed that its plasmid DNA (‘metaplasmidome’) carries stress-response genes. In particular it encoded antibiotic resistance genes, heavy metal resistance genes, cold shock proteins and genes encoding UV-protection. Those are functions that are directly involved in the adaptation of microbial communities to stresses in polar environments. It was further found that metaplasmidomes from the Siberian active layer originate mainly from Gammaproteobacteria. By applying enrichment cultures followed by plasmid DNA extraction it was possible to obtain a higher average contigs length and significantly higher recovery of plasmid sequences than from extracting plasmid sequences from metagenomes. The approach of analyzing ‘metaplasmidomes’ established in this thesis is therefore suitable for studying the ecological role of plasmids in polar environments in general. This thesis emphasizes that including microbial community dynamics have the potential to improve permafrost-carbon projections. Microbially mediated methane release from permafrost environments may significantly impact future climate change. This thesis identified drivers of methanogenic composition, abundance and activity in thawing permafrost landscapes. Finally, this thesis underlines the importance to study how the current warming Arctic affects microbial communities in order to gain more insight into microbial response and adaptation strategies.

Description / Table of Contents: Assumed comparable environmental conditions of early Mars and early Earth in 3.7 Ga ago – at a time when first fossil records of life on Earth could be found – suggest the possibility of life emerging on both planets in parallel. As conditions changed, the hypothetical life on Mars either became extinct or was able to adapt and might still exist in biological niches. The controversial discussed detection of methane on Mars led to the assumption, that it must have a recent origin – either abiotic through active volcanism or chemical processes, or through biogenic production. Spatial and seasonal variations in the detected methane concentrations and correlations between the presence of water vapor and geological features such as subsurface hydrogen, which are occurring together with locally increased detected concentrations of methane, gave fuel to the hypothesis of a possible biological source of the methane on Mars. Therefore the phylogenetically old methanogenic archaea, which have evolved under early Earth conditions, are often used as model-organisms in astrobiological studies to investigate the potential of life to exist in possible extraterrestrial habitats on our neighboring planet. In this thesis methanogenic archaea originating from two extreme environments on Earth were investigated to test their ability to be active under simulated Mars analog conditions. These extreme environments – the Siberian permafrost-affected soil and the chemoautotrophically based terrestrial ecosystem of Movile cave, Romania – are regarded as analogs for possible Martian (subsurface) habitats. Two novel species of methanogenic archaea isolated from these environments were described within the frame of this thesis. It could be shown that concentrations up to 1 wt% of Mars regolith analogs added to the growth media had a positive influence on the methane production rates of the tested methanogenic archaea, whereas higher concentrations resulted in decreasing rates. Nevertheless it was possible for the organisms to metabolize when incubated on water-saturated soil matrixes made of Mars regolith analogs without any additional nutrients. Long-term desiccation resistance of more than 400 days was proven with reincubation and indirect counting of viable cells through a combined treatment with propidium monoazide (to inactivate DNA of destroyed cells) and quantitative PCR. Phyllosilicate rich regolith analogs seem to be the best soil mixtures for the tested methanogenic archaea to be active under Mars analog conditions. Furthermore, in a simulation chamber experiment the activity of the permafrost methanogen strain Methanosarcina soligelidi SMA-21 under Mars subsurface analog conditions could be proven. Through real-time wavelength modulation spectroscopy measurements the increase in the methane concentration at temperatures down to -5 °C could be detected. The results presented in this thesis contribute to the understanding of the activity potential of methanogenic archaea under Mars analog conditions and therefore provide insights to the possible habitability of present-day Mars (near) subsurface environments. Thus, it contributes also to the data interpretation of future life detection missions on that planet. For example the ExoMars mission of the European Space Agency (ESA) and Roscosmos which is planned to be launched in 2018 and is aiming to drill in the Martian subsurface

Description / Table of Contents: The arctic-boreal treeline is a transition zone from taiga to tundra covering a vast area in Siberia. It often features large environmental gradients and reacts sensitively to changes in the environment. For example, the expansion of shrubs and a northward movement of the treeline are observable in Siberia as a response to the warming climate. The changes in vegetation across the treeline are known to influence the water chemistry in the lakes. This causes further alteration to the composition and diversity of sensitive aquatic organisms such as diatoms and macrophytes. Despite the rising awareness of the complex climate-feedback mechanisms of terrestrial plants, the understanding of their assembly rules and about responses of aquatic biomes in the surrounding treeline lakes is still limited. The goal of this thesis is to examine the previous and present biodiversity of terrestrial and freshwater biomes from the Siberian treeline ecotone, as well as their reactions to environmental changes. In particular, this thesis attempts to ...