Stratification, nitrogen fixation, and cyanobacterial bloom stage regulate the planktonic food web structure.

Changes in the complexity of planktonic food webs may be expected in future aquatic systems due to increases in sea surface temperature and an enhanced stratification of the water column. Under these conditions, the growth of unpalatable, filamentous, N2 -fixing cyanobacterial blooms, and their effect on planktonic food webs will become increasingly important. The planktonic food web structure in aquatic ecosystems at times of filamentous cyanobacterial blooms is currently unresolved, with discordant lines of evidence suggesting that herbivores dominate the mesozooplankton or that mesozooplankton organisms are mainly carnivorous. Here, we use a set of proxies derived from amino acid nitrogen stable isotopes from two mesozooplankton size fractions to identify changes in the nitrogen source and the planktonic food web structure across different microplankton communities. A transition from herbivory to carnivory in mesozooplankton between more eutrophic, near-coastal sites and more oligotrophic, offshore sites was accompanied by an increasing diversity of microplankton communities with aging filamentous cyanobacterial blooms. Our analyses of 124 biotic and abiotic variables using multivariate statistics confirmed salinity as a major driver for the biomass distribution of non-N2 -fixing microplankton species such as dinoflagellates. However, we provide strong evidence that stratification, N2 fixation, and the stage of the cyanobacterial blooms regulated much of the microplankton diversity and the mean trophic position and size of the metabolic nitrogen pool in mesozooplankton. Our empirical, macroscale data set consistently unifies contrasting results of the dominant feeding mode in mesozooplankton during blooms of unpalatable, filamentous, N2 -fixing cyanobacteria by identifying the at times important role of heterotrophic microbial food webs. Thus, carnivory, rather than herbivory, dominates in mesozooplankton during aging and decaying cyanobacterial blooms with hitherto uncharacterized consequences for the biogeochemical functions of mesozooplankton.

The Impact of the Major Baltic Inflow of December 2014 on the Mercury Species Distribution in the Baltic Sea.

The Baltic Sea is a marginal sea characterized by stagnation periods of several years. Oxygen consumption in its deep waters leads to the buildup of sulfide from sulfate reduction. Some of the microorganisms responsible for these processes also transform reactive ionic mercury to neurotoxic methylmercury. Episodic inflows of oxygenated saline water from the North Sea temporally re-establish oxic life in deep waters of the Baltic Sea. Thus, this sea is an especially important region to better understand mercury species distributions in connection with variable redox conditions. Mercury species were measured on three Baltic Sea campaigns, during the preinflow, ongoing inflow, and subsiding inflow of water, respectively, to the central basin. The inflowing water caused the removal of total mercury by 600 nmol m-2 and of methylmercury by 214 nmol m-2 in the Gotland Deep, probably via attachment of the mercury compounds to sinking particles. It appears likely that the consequences of the oxygenation of Baltic Sea deep waters, which are the coprecipitation of mercury species and the resettlement of the oxic deep waters, could lead to the enhanced transfer of accumulated mercury and methylmercury to the planktonic food chain and finally to fish.

Detoxification of sulphidic African shelf waters by blooming chemolithotrophs.

Coastal waters support approximately 90 per cent of global fisheries and are therefore an important food reserve for our planet. Eutrophication of these waters, due to human activity, leads to severe oxygen depletion and the episodic occurrence of hydrogen sulphide-toxic to multi-cellular life-with disastrous consequences for coastal ecosytems. Here we show that an area of approximately 7,000 km(2) of African shelf, covered by sulphidic water, was detoxified by blooming bacteria that oxidized the biologically harmful sulphide to environmentally harmless colloidal sulphur and sulphate. Combined chemical analyses, stoichiometric modelling, isotopic incubations, comparative 16S ribosomal RNA, functional gene sequence analyses and fluorescence in situ hybridization indicate that the detoxification proceeded by chemolithotrophic oxidation of sulphide with nitrate and was mainly catalysed by two discrete populations of gamma- and epsilon-proteobacteria. Chemolithotrophic bacteria, accounting for approximately 20 per cent of the bacterioplankton in sulphidic waters, created a buffer zone between the toxic sulphidic subsurface waters and the oxic surface waters, where fish and other nekton live. This is the first time that large-scale detoxification of sulphidic waters by chemolithotrophs has been observed in an open-ocean system. The data suggest that sulphide can be completely consumed by bacteria in the subsurface waters and, thus, can be overlooked by remote sensing or monitoring of shallow coastal waters. Consequently, sulphidic bottom waters on continental shelves may be more common than previously believed, and could therefore have an important but as yet neglected effect on benthic communities.

A full year of turbulence measurements from a drift campaign in the Arctic Ocean 2019-2020.

Ocean turbulent mixing is a key process in the global climate system, regulating ocean circulation and the uptake and redistribution of heat, carbon, nutrients, oxygen and other tracers. In polar oceans, turbulent heat transport additionally affects the sea ice mass balance. Due to the inaccessibility of polar regions, direct observations of turbulent mixing are sparse in the Arctic Ocean. During the year-long drift expedition "Multidisciplinary drifting Observatory for the Study of Arctic Climate" (MOSAiC) from September 2019 to September 2020, we obtained an unprecedented data set of vertical profiles of turbulent dissipation rate and water column properties, including oxygen concentration and fluorescence. Nearly 1,700 profiles, covering the upper ocean down to approximately 400 m, were collected in sets of 3 or more consecutive profiles every day, and complemented with several intensive sampling periods. This data set allows for the systematic assessment of upper ocean mixing in the Arctic, and the quantification of turbulent heat and nutrient fluxes, and can help to better constrain turbulence parameterizations in ocean circulation models.

Carbon assimilating fungi from surface ocean to subseafloor revealed by coupled phylogenetic and stable isotope analysis.

Fungi are ubiquitous in the ocean and hypothesized to be important members of marine ecosystems, but their roles in the marine carbon cycle are poorly understood. Here, we use 13C DNA stable isotope probing coupled with phylogenetic analyses to investigate carbon assimilation within diverse communities of planktonic and benthic fungi in the Benguela Upwelling System (Namibia). Across the redox stratified water column and in the underlying sediments, assimilation of 13C-labeled carbon from diatom extracellular polymeric substances (13C-dEPS) by fungi correlated with the expression of fungal genes encoding carbohydrate-active enzymes. Phylogenetic analysis of genes from 13C-labeled metagenomes revealed saprotrophic lineages related to the facultative yeast Malassezia were the main fungal foragers of pelagic dEPS. In contrast, fungi living in the underlying sulfidic sediments assimilated more 13C-labeled carbon from chemosynthetic bacteria compared to dEPS. This coincided with a unique seafloor fungal community and dissolved organic matter composition compared to the water column, and a 100-fold increased fungal abundance within the subseafloor sulfide-nitrate transition zone. The subseafloor fungi feeding on 13C-labeled chemolithoautotrophs under anoxic conditions were affiliated with Chytridiomycota and Mucoromycota that encode cellulolytic and proteolytic enzymes, revealing polysaccharide and protein-degrading fungi that can anaerobically decompose chemosynthetic necromass. These subseafloor fungi, therefore, appear to be specialized in organic matter that is produced in the sediments. Our findings reveal that the phylogenetic diversity of fungi across redox stratified marine ecosystems translates into functionally relevant mechanisms helping to structure carbon flow from primary producers in marine microbiomes from the surface ocean to the subseafloor.

Author Correction: Metabolic activity analyses demonstrate that Lokiarchaeon exhibits homoacetogenesis in sulfidic marine sediments.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Metabolic activity analyses demonstrate that Lokiarchaeon exhibits homoacetogenesis in sulfidic marine sediments.

The genomes of the Asgard superphylum of Archaea hold clues pertaining to the nature of the host cell that acquired the mitochondrion at the origin of eukaryotes1-4. Representatives of the Asgard candidate phylum Candidatus Lokiarchaeota (Lokiarchaeon) have the capacity for acetogenesis and fermentation5-7, but how their metabolic activity responds to environmental conditions is poorly understood. Here, we show that in anoxic Namibian shelf sediments, Lokiarchaeon gene expression levels are higher than those of bacterial phyla and increase with depth below the seafloor. Lokiarchaeon gene expression was significantly different across a hypoxic-sulfidic redox gradient, whereby genes involved in growth, fermentation and H2-dependent carbon fixation had the highest expression under the most reducing (sulfidic) conditions. Quantitative stable isotope probing revealed that anaerobic utilization of CO2 and diatomaceous extracellular polymeric substances by Lokiarchaeon was higher than the bacterial average, consistent with higher expression of Lokiarchaeon genes, including those involved in transport and fermentation of sugars and amino acids. The quantitative stable isotope probing and gene expression data demonstrate homoacetogenic activity of Candidatus Lokiarchaeota, whereby fermentative H2 production from organic substrates is coupled with the Wood-Ljungdahl carbon fixation pathway8. The high energetic efficiency provided by homoacetogenesis8 helps to explain the elevated metabolic activity of Lokiarchaeon in this anoxic, energy-limited setting.

The tropical-subtropical coupling in the Southeast Atlantic from the perspective of the northern Benguela upwelling system.

In the Benguela upwelling system, the environmental conditions are determined to a large extent by central water masses advected from remote areas onto the shelf. The origin, spreading pathways and fate of those water masses are investigated with a regional ocean model that is analysed using Eulerian passive tracers and on the basis of Lagrangian trajectories. Two major water masses influencing the Benguela upwelling system are identified: tropical South Atlantic Central Water (SACW) and subtropical Eastern South Atlantic Central Water (ESACW). The spreading of tropical waters into the subtropical Benguela upwelling system is mediated by equatorial currents and their continuation in the Southeast Atlantic. This tropical-subtropical connection has been attributed to signal propagation in the equatorial and coastal waveguides. However, there exists an additional spreading path for tropical central water in the open ocean. This mass transport fluctuates on a seasonal scale around an averaged meridional transport in Sverdrup balance. The inter-annual variability of the advection of tropical waters is related to Benguela Niños, as evidenced by the 2010/2011 event. The northern Benguela upwelling system is a transition zone between SACW and ESACW since they encounter each other at about 20°S. Both water masses have seasonal variable shares in the upwelled water there. To summarise the main pathways of central water mass transport, an enhanced scheme for the subsurface circulation in the Southeast Atlantic is presented.

Expanding oxygen-minimum zones in the tropical oceans.

Oxygen-poor waters occupy large volumes of the intermediate-depth eastern tropical oceans. Oxygen-poor conditions have far-reaching impacts on ecosystems because important mobile macroorganisms avoid or cannot survive in hypoxic zones. Climate models predict declines in oceanic dissolved oxygen produced by global warming. We constructed 50-year time series of dissolved-oxygen concentration for select tropical oceanic regions by augmenting a historical database with recent measurements. These time series reveal vertical expansion of the intermediate-depth low-oxygen zones in the eastern tropical Atlantic and the equatorial Pacific during the past 50 years. The oxygen decrease in the 300- to 700-m layer is 0.09 to 0.34 micromoles per kilogram per year. Reduced oxygen levels may have dramatic consequences for ecosystems and coastal economies.