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Microbial mats are self-sustaining benthic ecosystems composed of highly diverse microbial communities. It has been proposed that microbial mats were widespread in Proterozoic marine environments, prior to the emergence of bioturbating organisms at the Precambrian–Cambrian transition. One characteristic feature of Precambrian biomarker records is that steranes are typically absent or occur in very low concentrations. This has been explained by low eukaryotic source inputs, or degradation of primary produced sterols in benthic microbial mats (“mat-seal effect”). To better understand the preservational pathways of sterols in microbial mats, we analyzed freely extractable and carbonate-bound lipid fractions as well as decalcified extraction residues in different layers of a recent calcifying mat (∼1500 years) from the hypersaline Lake 2 on the island of Kiritimati, central Pacific. A variety of C27–C29 sterols and distinctive C31 4α-methylsterols (4α-methylgorgosterol and 4α-methylgorgostanol, biomarkers for dinoflagellates) were detected in freely extractable and carbonate-bound lipid pools. These sterols most likely originated from organisms living in the water column and the upper mat layers. This autochthonous biomass experienced progressive microbial transformation and degradation in the microbial mat, as reflected by a significant drop in total sterol concentrations, up to 98 %, in the deeper layers, and a concomitant decrease in total organic carbon. Carbonate-bound sterols were generally low in abundance compared to the freely extractable portion, suggesting that incorporation into the mineral matrix does not play a major role in the preservation of eukaryotic sterols in this mat. Likewise, pyrolysis of extraction residues suggested that sequestration of steroid carbon skeletons into insoluble organic matter was low compared to hopanoids. Taken together, our findings argue for a major mat-seal effect affecting the distribution and preservation of steroids in the mat studied. This result markedly differs from recent findings made for another microbial mat growing in the nearby hypersaline Lake 22 on the same island, where sterols showed no systematic decrease with depth. The observed discrepancies in the taphonomic pathways of sterols in microbial mats from Kiritimati may be linked to multiple biotic and abiotic factors including salinity and periods of subaerial exposure, implying that caution has to be exercised in the interpretation of sterol distributions in modern and ancient microbial mat settings. more...

Organic matter in Archean hydrothermal cherts may provide an important archive for molecular traces of the earliest life on Earth. The geobiological interpretation of this archive, however, requires a sound understanding of organic matter preservation and alteration in hydrothermal systems. Here we report on organic matter (including molecular biosignatures) enclosed in hydrothermally influenced cherts of the Pleistocene Lake Magadi (Kenya; High Magadi Beds and Green Beds). The Magadi cherts contain low organic carbon (n-alkanes, hopanes, and polycyclic aromatic hydrocarbons (PAHs) indicate that a fraction of the bitumen has been thermally altered to early or peak oil window maturity. This more mature fraction likely originated from defunctionalization of dissolved organic matter and/or hydrothermal petroleum formation at places of higher thermal flux. Like the bitumens, the kerogens also show variations in thermal maturities, which can partly be explained by admixture of thermally pre-altered macromolecules. However, findings of archaea-derived isoprenoid moieties (C20 and C25 chains) in kerogen pyrolysates indicate rapid sequestration of some archaeal lipids into kerogen while hydrothermal alteration was active. We posit that such early sequestration may enhance the resistance of molecular biosignatures against in situ hydrothermal and post-depositional alteration. Furthermore, the co-occurrence of organic matter with different thermal maturities in the Lake Magadi cherts suggests that similar findings in Archean hydrothermal deposits could partly reflect original environmental conditions and not exclusively post-depositional overprint or contamination. Our results support the view that kerogen in Archean hydrothermal cherts may contain important information on early life. Our study also highlights the suitability of Lake Magadi as an analog system for hydrothermal chert environments on the Archean Earth. more...

Several methanogenic pathways in oxic surface waters were recently discovered, but their relevance in the natural environment is still unknown. Our study examines distinct methane (CH4) enrichments that repeatedly occur below the thermocline during the summer months in the central Baltic Sea. In agreement with previous studies in this region, we discovered differences in the methane distributions between the western and eastern Gotland Basin, pointing to in situ methane production below the thermocline in the latter (concentration of CH4 14.1±6.1 nM, δ13C CH4 −62.9 ‰). Through the use of a high-resolution hydrographic model of the Baltic Sea, we showed that methane below the thermocline can be transported by upwelling events towards the sea surface, thus contributing to the methane flux at the sea–air interface. To quantify zooplankton-associated methane production rates, we developed a sea-going methane stripping-oxidation line to determine methane release rates from copepods grazing on 14C-labelled phytoplankton. We found that (1) methane production increased with the number of copepods, (2) higher methane production rates were measured in incubations with Temora longicornis (125±49 fmol methane copepod−1 d−1) than in incubations with Acartia spp. (84±19 fmol CH4 copepod−1 d−1) dominated zooplankton communities, and (3) methane was only produced on a Rhodomonas sp. diet, and not on a cyanobacteria diet. Furthermore, copepod-specific methane production rates increased with incubation time. The latter finding suggests that methanogenic substrates for water-dwelling microbes are released by cell disruption during feeding, defecation, or diffusion from fecal pellets. In the field, particularly high methane concentrations coincided with stations showing a high abundance of DMSP/DMSO-rich Dinophyceae. Lipid biomarkers extracted from phytoplankton- and copepod-rich samples revealed that Dinophyceae are a major food source of the T. longicornis dominated zooplankton community, supporting the proposed link between copepod grazing, DMSP/DMSO release, and the build-up of subthermocline methane enrichments in the central Baltic Sea. more...

Magmatic sill intrusions into organic-rich sediments cause the release of thermogenic CH4 and CO2. Pore fluids from the Guaymas Basin (Gulf of California), a sedimentary basin with recent magmatic activity, were investigated to constrain the link between sill intrusions and fluid seepage as well as the timing of sill-induced hydrothermal activity. Sampling sites were close to a hydrothermal vent field at the northern rift axis and at cold seeps located up to 30 km away from the rift. Pore fluids close to the active hydrothermal vent field showed a slight imprint by hydrothermal fluids and indicated a shallow circulation system transporting seawater to the hydrothermal catchment area. Geochemical data of pore fluids at cold seeps showed a mainly ambient diagenetic fluid composition without any imprint related to high temperature processes at greater depth. Seep communities at the seafloor were mainly sustained by microbial methane, which rose along pathways formed earlier by hydrothermal activity, driving the anaerobic oxidation of methane (AOM) and the formation of authigenic carbonates.Overall, our data from the cold seep sites suggest that at present, sill-induced hydrothermalism is not active away from the ridge axis, and the vigorous venting of hydrothermal fluids is restricted to the ridge axis. Using the sediment thickness above extinct conduits and carbonate dating, we calculated that deep fluid and thermogenic gas flow ceased 28 to 7 kyr ago. These findings imply a short lifetime of hydrothermal systems, limiting the time of unhindered carbon release as suggested in previous modeling studies. Consequently, activation and deactivation mechanisms of these systems need to be better constrained for the use in climate modeling approaches. more...

Archaean hydrothermal chert veins commonly contain abundant organic carbon of uncertain origin (abiotic vs. biotic). In this study, we analysed kerogen contained in a hydrothermal chert vein from the ca. 3.5 Ga Dresser Formation (Pilbara Craton, Western Australia). Catalytic hydropyrolysis (HyPy) of this kerogen yielded n-alkanes up to n-C22, with a sharp decrease in abundance beyond n-C18. This distribution ( ≤ n-C18) is very similar to that observed in HyPy products of recent bacterial biomass, which was used as reference material, whereas it differs markedly from the unimodal distribution of abiotic compounds experimentally formed via Fischer–Tropsch-type synthesis. We therefore propose that the organic matter in the Archaean chert veins has a primarily microbial origin. The microbially derived organic matter accumulated in anoxic aquatic (surface and/or subsurface) environments and was then assimilated, redistributed and sequestered by the hydrothermal fluids (hydrothermal pump hypothesis). more...

The water column of the Landsort Deep, central Baltic Sea, is stratified into an oxic, suboxic, and anoxic zone. This stratification controls the distributions of individual microbial communities and biogeochemical processes. In summer 2011, particulate organic matter was filtered from these zones using an in situ pump. Lipid biomarkers were extracted from the filters to establish water-column profiles of individual hydrocarbons, alcohols, phospholipid fatty acids, and bacteriohopanepolyols (BHPs). As a reference, a cyanobacterial bloom sampled in summer 2012 in the central Baltic Sea Gotland Deep was analyzed for BHPs. The biomarker data from the surface layer of the oxic zone showed major inputs from cyanobacteria, dinoflagellates, and ciliates, while the underlying cold winter water layer was characterized by a low diversity and abundance of organisms, with copepods as a major group. The suboxic zone supported bacterivorous ciliates, type I aerobic methanotrophic bacteria, sulfate-reducing bacteria, and, most likely, methanogenic archaea. In the anoxic zone, sulfate reducers and archaea were the dominating microorganisms as indicated by the presence of distinctive branched fatty acids: archaeol and pentamethylicosane (PMI) derivatives, respectively. Our study of in situ biomarkers in the Landsort Deep thus provided an integrated insight into the distribution of relevant compounds and describes useful tracers to reconstruct stratified water columns in the geological record. more...

The Baltic Sea, one of the world's largest brackish-marine basins, established after deglaciation of Scandinavia about 17 000 to 15 000 yr ago. In the changeable history of the Baltic Sea, the initial freshwater system was connected to the North Sea about 8000 yr ago and the modern brackish-marine setting (Littorina Sea) was established. Today, a relatively stable stratification has developed in the water column of the deep basins due to salinity differences. Stratification is only occasionally interrupted by mixing events, and it controls nutrient availability and growth of specifically adapted microorganisms and algae. We studied bacteriohopanepolyols (BHPs), lipids of specific bacterial groups, in a sediment core from the central Baltic Sea (Gotland Deep) and found considerable differences between the distinct stages of the Baltic Sea's history. Some individual BHP structures indicate contributions from as yet unknown redoxcline-specific bacteria (bacteriohopanetetrol isomer), methanotrophic bacteria (35-aminobacteriohopanetetrol), cyanobacteria (bacteriohopanetetrol cyclitol ether isomer) and from soil bacteria (adenosylhopane) through allochthonous input after the Littorina transgression, whereas the origin of other BHPs in the core has still to be identified. Notably high BHP abundances were observed in the deposits of the brackish-marine Littorina phase, particularly in laminated sediment layers. Because these sediments record periods of stable water column stratification, bacteria specifically adapted to these conditions may account for the high portions of BHPs. An additional and/or accompanying source may be nitrogen-fixing (cyano)bacteria, which is indicated by a positive correlation of BHP abundances with Corg and δ15N. more...

Water column samples taken in summer 2008 from the stratified Gotland Deep (central Baltic Sea) showed a strong gradient in dissolved methane concentrations from high values in the saline deep water (max. 504 nM) to low concentrations in the less dense, brackish surface water (about 4 nM). The steep methane-gradient (between 115 and 135 m water depth) within the redox-zone, which separates the anoxic deep part from the oxygenated surface water (oxygen concentration 0–0.8 mL L−1), implies a methane consumption rate of 0.28 nM d−1. The process of microbial methane oxidation within this zone was evident by a shift of the stable carbon isotope ratio of methane between the bottom water (δ13C CH4 = −82.4‰ and the redox-zone (δ13C CH4 = −38.7‰. Water column samples between 80 and 119 m were studied to identify the microorganisms responsible for the methane turnover in that depth interval. Notably, methane monooxygenase gene expression analyses for water depths covering the whole redox-zone demonstrated that accordant methanotrophic activity was probably due to only one phylotype of the aerobic type I methanotrophic bacteria. An imprint of these organisms on the particular organic matter was revealed by distinctive lipid biomarkers showing bacteriohopanepolyols and lipid fatty acids characteristic for aerobic type I methanotrophs (e.g., 35-aminobacteriohopane-30,31,32,33,34-pentol), corroborating their role in aerobic methane oxidation in the redox-zone of the central Baltic Sea. more...

Methane-related carbonates from Hydrate Ridge typically show several macroscopically distinguishable mineral phases, namely whitish aragonite, lucent aragonite, and gray micrite. The relationship of these phases to particular microorganisms or biogeochemical processes is as yet unclear. We used a miniaturized biomarker technique on mg samples, combined with factor analysis and subsequent electron microprobe analysis, to study lipid biomarkers and chemical compositions of the individual phases. This allows us to identify particular mechanisms involved in the formation of the different carbonate precipitates. Our combined analysis of biomarkers and petrographic traits shows that most of the lipids related to the anaerobic oxidation of methane (>90% by weight) are concentrated within only a minor compartment (~20% by volume) of the Hydrate Ridge carbonates, the whitish aragonite. The patterns indicate that the whitish aragonite represents fossilized biofilms of methanotrophic consortia containing mainly archaea of the ANME-2 group and sulfate reducing bacteria, whereas the precipitation of the lucent aragonite may have lacked the immediate proximity of microorganisms during formation. By contrast, the gray micrite formed by incorporation of allochthonous organic and inorganic matter during carbonate precipitation induced by the anaerobic oxidation of methane involving ANME-1 archaea. more...