Tobias Goldhammer

Thermococcus kodakarensis modulates its polar membrane lipids and elemental composition according to growth stage and phosphate availability

We observed significant changes in the elemental and intact polar lipid (IPL) composition of the archaeon Thermococcus kodakarensis (KOD1) in response to growth stage and phosphorus supply. Reducing the amount of organic supplements and phosphate in growth media resulted in significant decreases in cell size and cellular quotas of carbon (C), nitrogen (N), and phosphorus (P), which coincided with significant increases in cellular IPL quota and IPLs comprising multiple P atoms and hexose moieties. Relatively more cellular P was stored as IPLs in P-limited cells (2–8%) compared to control cells (<0.8%). We also identified a specific IPL biomarker containing a phosphatidyl-N-acetylhexoseamine headgroup that was relatively enriched during rapid cell division. These observations serve as empirical evidence of IPL adaptations in Archaea that will help to interpret the distribution of these biomarkers in natural systems. The reported cell quotas of C, N, and P represent the first such data for a specific archaeon and suggest that thermophiles are C-rich compared to the cell carbon-to-volume relationship reported for planktonic bacteria.

Extensive nitrogen loss from permeable sediments off North-West Africa

The upwelling area off North-West Africa is characterized by high export production, high nitrate and low oxygen concentration in bottom waters. The underlying sediment consists of sands that cover most of the continental shelf. Due to their permeability sands allow for fast advective pore water transport and can exhibit high rates of nitrogen (N) loss via denitrification as reported for anthropogenically eutrophied regions. However, N loss from sands underlying naturally eutrophied waters is not well studied, and in particular, N loss from the North-West African shelf is poorly constrained. During two research cruises in April/May 2010/2011, sediment was sampled along the North-West African shelf and volumetric denitrification rates were measured in sediment layers down to 8 cm depth using slurry incubations with 15N-labeled nitrate. Areal N loss was calculated by integrating volumetric rates down to the nitrate penetration depth derived from pore water profiles. Areal N loss was neither correlated with water depth nor with bottom water concentrations of nitrate and oxygen but was strongly dependent on sediment grain size and permeability. The derived empirical relation between benthic N loss and grains size suggests that pore water advection is an important regulating parameter for benthic denitrification in sands and further allowed extrapolating rates to an area of 53,000 km2 using detailed sediment maps. Denitrification from this region amounts to 995 kt yr-1 (average 3.6 mmol m-2 d-1) which is 4 times higher than previous estimates based on diffusive pore water transport. Sandy sediments cover 50-60% of the continental shelf and thus may contribute significantly to the global benthic N loss.

Ammonia‐oxidizing Bacteria of the Nitrosospira cluster 1 dominate over ammonia‐oxidizing Archaea in oligotrophic surface sediments near the South Atlantic Gyre

Summary Sediments across the Namibian continental margin feature a strong microbial activity gradient at their surface. This is reflected in ammonium concentrations of < 10 μM in oligotrophic abyssal plain sediments near the South Atlantic Gyre compared with ammonium concentrations of > 700 μM in upwelling areas near the coast. Here we address changes in apparent abundance and structure of ammonia‐oxidizing archaeal and bacterial communities (AOA and AOB) along a transect of seven sediment stations across the Namibian shelf by analysing their respective ammonia monooxygenase genes (amoA). The relative abundance of archaeal and bacterial amoA (g−1 DNA) decreased with increasing ammonium concentrations, and bacterial amoA frequently outnumbered archaeal amoA at the sediment–water interface [0–1 cm below seafloor (cmbsf)]. In contrast, AOA were apparently as abundant as AOB or dominated in several deeper (> 10 cmbsf), anoxic sediment layers. Phylogenetic analyses showed a change within the AOA community along the transect, from two clusters without cultured representatives at the gyre to N itrososphaera and N itrosopumilus clusters in the upwelling region. AOB almost exclusively belonged to the N itrosospira cluster 1. Our results suggest that this predominantly marine AOB lineage without cultured representatives can thrive at low ammonium concentrations and is active in the marine nitrogen cycle.

Near-surface heating of young rift sediment causes mass production and discharge of reactive dissolved organic matter

Ocean margin sediments have been considered as important sources of dissolved organic carbon (DOC) to the deep ocean, yet the contribution from advective settings has just started to be acknowledged. Here we present evidence showing that near-surface heating of sediment in the Guaymas Basin, a young extensional depression, causes mass production and discharge of reactive dissolved organic matter (DOM). In the sediment heated up to ~100 °C, we found unexpectedly low DOC concentrations in the pore waters, reflecting the combined effect of thermal desorption and advective fluid flow. Heating experiments suggested DOC production to be a rapid, abiotic process with the DOC concentration increasing exponentially with temperature. The high proportions of total hydrolyzable amino acids and presence of chemical species affiliated with activated hydrocarbons, carbohydrates and peptides indicate high reactivity of the DOM. Model simulation suggests that at the local scale, near-surface heating of sediment creates short and massive DOC discharge events that elevate the bottom-water DOC concentration. Because of the heterogeneous distribution of high heat flow areas, the expulsion of reactive DOM is spotty at any given time. We conclude that hydrothermal heating of young rift sediments alter deep-ocean budgets of bioavailable DOM, creating organic-rich habitats for benthic life.

The evolution of Saharan dust input on Lanzarote (Canary Islands) — influenced by human activity in the Northwest Sahara during the early Holocene?

An overall Holocene increase of Saharan dust input to the Canary Islands and to the North Canary Basin is accompanied by a strong coarsening of Saharan dust in loess-like sediments deposited on Lanzarote from ~7—8 ka. No similar coarsening events are indicated in investigations of the sedimentological record for the last 180 ka, a period showing several dramatic climate changes. Therefore a mobilisation of Holocene dust by anthropogenic activity in the northwest Sahara east of the Canary Islands is assumed. Although scarce archaeological data from the coastal area of that region does not point to strong anthropogenic activity during the early Holocene, a high density of unexplored archaeological remains is reported from the coastal hinterlands in the Western Sahara. Thus, the hypothesis of early anthropogenic activity cannot be excluded.

Intense biological phosphate uptake onto particles in subeuphotic continental margin waters

Elucidating the processes that affect particulate phosphorus (P) export from the euphotic zone and burial in sediments is important for models of global phosphorus, nitrogen and carbon cycling. We investigated dissolved inorganic Pi incorporation into particles (>0.2 µm) in the sub-euphotic zone and benthic boundary layer (BBL) of high productivity Mauritanian and Namibian shelf waters, using 33PO43- tracer experiments combined with a sequential chemical extraction analysis. Pi uptake (5.4 to19.9 nmol P L-1d-1) by particulate matter was biologically mediated (~50% into the organic fraction), and similar to estimated rates of heterotrophic growth. Thus, a substantial fraction of Pi must be recycled through a particle-associated microbial pool. Rapid adsorption of 33P in the anoxic waters of Namibia indicated the additional existence of a large pool of surface exchangeable P. Particle associated Pi recycling and adsorption may influence the export flux and ultimate fate of particle bound P in continental shelf waters.

Temporal stability and origin of chemoclines in the deep hypersaline anoxic Urania basin

Submarine brine lakes feature sharp and persistent concentration gradients between seawater and brine, though these should be smoothed out by free diffusion in open ocean settings. The anoxic Urania basin of the eastern Mediterranean contains an ultrasulfidic, hypersaline brine of Messinian origin above a thick layer of suspended sediments. With a dual modeling approach we reconstruct its contemporary stratification by geochemical solute transport fundamentals and show that thermal convection is required to maintain mixing in the brine and mud layer. The origin of the Urania basin stratification was dated to 1650 years B.P., which may be linked to a major earthquake in the region. The persistence of the chemoclines may be key to the development of diverse and specialized microbial communities. Ongoing thermal convection in the fluid mud layer may have important yet unresolved consequences for sedimentological and geochemical processes, also in similar environments.

Isoprenoid Quinones Resolve the Stratification of Redox Processes in a Biogeochemical Continuum from the Photic Zone to Deep Anoxic Sediments of the Black Sea

ABSTRACT The stratified water column of the Black Sea serves as a model ecosystem for studying the interactions of microorganisms with major biogeochemical cycles. Here, we provide detailed analysis of isoprenoid quinones to study microbial redox processes in the ocean. In a continuum from the photic zone through the chemocline into deep anoxic sediments of the southern Black Sea, diagnostic quinones and inorganic geochemical parameters indicate niche segregation between redox processes and corresponding shifts in microbial community composition. Quinones specific for oxygenic photosynthesis and aerobic respiration dominate oxic waters, while quinones associated with thaumarchaeal ammonia oxidation and bacterial methanotrophy, respectively, dominate a narrow interval in suboxic waters. Quinone distributions indicate highest metabolic diversity within the anoxic zone, with anoxygenic photosynthesis being a major process in its photic layer. In the dark anoxic layer, quinone profiles indicate the occurrence of bacterial sulfur and nitrogen cycling, archaeal methanogenesis, and archaeal methanotrophy. Multiple novel ubiquinone isomers, possibly originating from unidentified intra-aerobic anaerobes, occur in this zone. The respiration modes found in the anoxic zone continue into shallow subsurface sediments, but quinone abundances rapidly decrease within the upper 50 cm below the sea floor, reflecting the transition to lower energy availability. In the deep subseafloor sediments, quinone distributions and geochemical profiles indicate archaeal methanogenesis/methanotrophy and potentially bacterial fermentative metabolisms. We observed that sedimentary quinone distributions track lithology, which supports prior hypotheses that deep biosphere community composition and metabolisms are determined by environmental conditions during sediment deposition. IMPORTANCE Microorganisms play crucial roles in global biogeochemical cycles, yet we have only a fragmentary understanding of the diversity of microorganisms and their metabolisms, as the majority remains uncultured. Thus, culture-independent approaches are critical for determining microbial diversity and active metabolic processes. In order to resolve the stratification of microbial communities in the Black Sea, we comprehensively analyzed redox process-specific isoprenoid quinone biomarkers in a unique continuous record from the photic zone through the chemocline into anoxic subsurface sediments. We describe an unprecedented quinone diversity that allowed us to detect distinct biogeochemical processes, including oxygenic photosynthesis, archaeal ammonia oxidation, aerobic methanotrophy, and anoxygenic photosynthesis in defined geochemical zones.