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The effects of bottom water oxygen concentration on sediment oxygen uptake, oxygen penetration depth, nitrate and ammonium fluxes, anammox, denitrification, dissimilatory nitrate reduction to ammonium, nitrification, and mineralization were investigated off the Changjiang estuary and its adjacent East China Sea, by combining a seasonal comparison with three artificially induced bottom water oxygen conditions (oxic, ambient, and severe hypoxia). A 50% decrease in in-situ bottom water oxygen concentrations between May and August, led to decreases in the average sediment oxygen uptake and oxygen penetration depth by 23% and 29%, respectively. Anammox rates decreased by a factor of 2.5, and the relative contribution of anammox to the total benthic N-loss decreased from 20% to 7.4%. However, denitrification rates increased, leading to an overall benthic N-loss rate of 0.85 mmol N m(-2)d(-1). At the same time, an increasing contribution of dissimilatory nitrate reduction to ammonium to total nitrate reduction led to higher recycling of inorganic nitrogen during hypoxia in August. Under artificially induced conditions of severe hypoxia, there was a sharp decrease in both sediment oxygen uptake and benthic N-loss rates by 88% and 38%, respectively. Nitrate and ammonium fluxes showed complex behavior at different sites which could be related to the repression of sedimentary nitrification below a bottom water oxygen threshold of 9.7 mu M and increasing dissimilatory nitrate reduction to ammonium. Taken together, our results indicate that changes in benthic nutrient cycling under seasonal hypoxia enhance the retention of both organic and inorganic nitrogen, thereby exacerbating oxygen deficiency.

Diatoms are among the few eukaryotes known to store nitrate (NO3−) and to use it for dissimilatory nitrate/nitrite reduction to ammonium (DNRA) to generate enegry in the absence of light and O2. We used stable isotope incubations and in situ microsensor measurements over complete light cycles to study the diel activity transitions of the NO3−-storing benthic diatom Craticula cuspidata in the submerged Middle Island Sinkhole, Lake Huron (USA). We found that this diatom links NO3− respiration to diel migration into deep (~4 cm) sulfidic sediments below the microbial mat. This pattern was accompanied by pronounced diel changes in the depth of sulphide consumption. During the day sulphide was consumed by anoxygenic photosynthesis and aerobic sulphide oxidation in the uppermost few mm. Surprisingly, the consumption zone moved downward in the evening and was deepest in the sediment at night. Thus, the sulphide consumption zone strikingly overlapped with the depth of DNRA-performing diatom residence. Using an enrichment of Craticula cuspidata, we found that the nitrate respiration via DNRA was ~10-fold higher in the presence of sulphide. Overall, our data therefore indicate that C. cuspidata and/or their microbiome link NO3− reduction to sulphide oxidation.

The surface waters of the open ocean are mostly oversaturated with methane and thus act as a net source to the atmosphere. In situ production from methylated compounds or inorganic carbon has been proposed to act as a potential source of methane in the oxygenated surface waters. However, the distribution and importance of aerobic methane production in different marine regions remains poorly constrained. We investigated the processes and microorganisms involved in aerobic methane production in the surface waters of the western Tropical North Atlantic off Barbados. Using stable isotope incubation experiments, we showed that within 24 hours methane was readily produced from methylphosphonate (MPn) but not from dissolved inorganic carbon. MPn-derived methane production reached up to ca. 8 nmol l-1 d-1, with the highest rates measured in surface waters above the deep chlorophyll maximum (DCM). Additions of inorganic phosphate resulted in the suppression of methane production in the surface waters but not below the DCM. Additional controlling factors of MPn-derived methane production, both physicochemical (depth, nutrients) as well as biological (primary production, nitrogen fixation), were also investigated. Our metagenomic and metatranscriptomic analyses revealed that various microbial groups, including Trichodesmium and Alphaproteobacteria, had the capacity to utilise MPn in situ, making them potential contributors to methane production in this region. Overall, our results highlight the importance of MPn-derived methane production in the phosphate-limited western Tropical North Atlantic and identify the controlling factors that may regulate aerobic methane production and thereby exert control over marine methane emissions.

Oligotrophic ocean gyre ecosystems may be expanding due to rising global temperatures [1–5]. Models predicting carbon flow through these changing ecosystems require accurate descriptions of phytoplankton communities and their metabolic activities [6]. We therefore measured distributions and activities of cyanobacteria and small photosynthetic eukaryotes throughout the euphotic zone on a zonal transect through the South Pacific Ocean, focusing on the ultraoligotrophic waters of the South Pacific Gyre (SPG). Bulk rates of CO2 fixation were low (0.1 µmol C l−1 d−1) but pervasive throughout both the surface mixed-layer (upper 150 m), as well as the deep chlorophyll a maximum of the core SPG. Chloroplast 16S rRNA metabarcoding, and single-cell 13CO2 uptake experiments demonstrated niche differentiation among the small eukaryotes and picocyanobacteria. Prochlorococcus abundances, activity, and growth were more closely associated with the rims of the gyre. Small, fast-growing, photosynthetic eukaryotes, likely related to the Pelagophyceae, characterized the deep chlorophyll a maximum. In contrast, a slower growing population of photosynthetic eukaryotes, likely comprised of Dictyochophyceae and Chrysophyceae, dominated the mixed layer that contributed 65–88% of the areal CO2 fixation within the core SPG. Small photosynthetic eukaryotes may thus play an underappreciated role in CO2 fixation in the surface mixed-layer waters of ultraoligotrophic ecosystems., The ISME Journal, 16 (2), ISSN:1751-7362, ISSN:1751-7370

Significance Seagrass meadows colonize shallow coastlines around the world and represent sites of intense carbon cycling. Due to their capacity to produce methane, seagrass ecosystems constitute net sources of methane to the atmosphere. Here, we identify key processes and microorganisms responsible for methane formation in seagrass-covered sediments in the Mediterranean Sea. Our work shows that methane is solely formed from methylated compounds that are produced and released by the plant itself. Due to the persistence of these compounds in buried plant material, microbial methane production continues long after the death of the living plant. These results provide a comprehensive understanding of methane production in seagrass habitats, thereby contributing to our knowledge on these important blue carbon ecosystems.

Global-scale N-oxide contamination of groundwater within aquifers occurs due to the widespread use of N-bearing fertilizers and chemicals, threatening both human and environmental health. However, the conversion of these pollutants in active nitrogen (N) cycling processes in the subsurface biosphere still remains unclear. This study investigates the global occurrence of anaerobic ammonium oxidation (anammox) in aquifers, where anammox was found to be turned on and off between saturated and unsaturated soil horizons, and contributed 36.8–79.5% to N loss in saturated soil horizons, the remainder being due to denitrification which has traditionally been considered the main pathway for removal of N-pollutants from aquifers. Although anammox activity was undetectable in the unsaturated soil horizons, it could potentially be activated by contact with ascending groundwater. High-throughput pyrosequencing analysis identified Candidatus Brocadia anammoxidans as being the most abundant anammox bacterium in the saturated soils investigated. However, the anammox bacterial abundance was determined by the relative richness of Candidatus Jettenia asiatica. Isotopic pairing experiments revealed that coupling anammox with ammonium oxidation and respiratory ammonification enabled the formation of a revised N cycle in aquifer systems, in which respiratory ammonification acted as an important coordinator. Anammox can therefore contribute substantially to aquifer N cycling and its role in remediation of aquifers contaminated with N-oxides may be of global importance.

The subtropical South Pacific Gyre (SPG) encompasses the largest oligotrophic region of the global ocean. In these remote waters dissolved organic matter (DOM) accumulates in the surface waters, though constituting a potential source of nutrients and energy to sustain microbial life. On a zonal transect across the SPG, we quantified bulk dissolved organic carbon (DOC) and assessed the DOM composition via ultrahigh resolution mass spectrometry (UHR-MS) of solid-phase extracted DOC (SPE-DOC) to elucidate the molecular-level reasons behind the apparent recalcitrance of the DOM prevailing in the SPG. We included a comparison between two individual formula assignment approaches to UHR-MS data in absorption and magnitude mode which yielded consistent results. DOC concentrations exceeding 100 μmol C L−1 in the warm and saline waters of the central gyre were higher than in the surface waters of the adjacent western South Pacific. Along the transect, concentrations of SPE-DOC were less variable than bulk DOC. Nevertheless, molecular-level investigation revealed that the composition of the DOM accumulated in the central SPG generally conformed to characteristics of surface ocean DOM, but all assessed properties were more pronounced. We found high abundances of potentially labile unsaturated aliphatic molecular formulas and a low calculated degradation index for the DOM of likely marine microbial origin. Markedly decreased molar N/C ratios in the central gyre indicated preferential microbial utilization of nitrogen-containing DOM. A distinct imprint of extensive photochemical reworking was manifested in the low aromaticity of the DOM in the photic layer. Over the whole water column, ageing of DOM was evident through the small, but significant contribution of SPE-DOC to apparent oxygen utilization as well as on molecular level. Our findings demonstrate that SPE-DOC captures carbon fractions relevant on timescales of seasons to timescales covering ocean circulation and biogeochemical processes in stable gyre systems are imprinted in the DOM molecular composition.

Filaments and fronts play a crucial role for a net offshore and downward nutrient transport in Eastern Boundary Upwelling Systems (EBUSs) and thereby reduce regional primary production. Most studies on this topic are based on either observations or model simulations, but only seldom are both approaches are combined quantitatively to assess the importance of filaments for primary production and nutrient transport. Here we combine targeted interdisciplinary shipboard observations of a cold filament off Peru with submesoscale-permitting (1/45∘) coupled physical (Coastal and Regional Ocean Community model, CROCO) and biogeochemical (Pelagic Interaction Scheme for Carbon and Ecosystem Studies, PISCES) model simulations to (i) evaluate the model simulations in detail, including the timescales of biogeochemical modification of the newly upwelled water, and (ii) quantify the net effect of submesoscale fronts and filaments on primary production in the Peruvian upwelling system. The observed filament contains relatively cold, fresh, and nutrient-rich waters originating in the coastal upwelling. Enhanced nitrate concentrations and offshore velocities of up to 0.5 m s−1 within the filament suggest an offshore transport of nutrients. Surface chlorophyll in the filament is a factor of 4 lower than at the upwelling front, while surface primary production is a factor of 2 higher. The simulation exhibits filaments that are similar in horizontal and vertical scale compared to the observed filament. Nitrate concentrations and primary production within filaments in the model are comparable to observations as well, justifying further analysis of nitrate uptake and subduction using the model. Virtual Lagrangian floats were released in the subsurface waters along the shelf and biogeochemical variables tracked along the trajectories of floats upwelled near the coast. In the submesoscale-permitting (1/45∘) simulation, 43 % of upwelled floats and 40 % of upwelled nitrate are subducted within 20 d after upwelling, which corresponds to an increase in nitrate subduction compared to a mesoscale-resolving (1/9∘) simulation by 14 %. Taking model biases into account, we give a best estimate for subduction of upwelled nitrate off Peru between 30 %– 40 %. Our results suggest that submesoscale processes further reduce primary production by amplifying the downward and offshore export of nutrients found in previous mesoscale studies, which are thus likely to underestimate the reduction in primary production due to eddy fluxes. Moreover, this downward and offshore transport could also enhance the export of fresh organic matter below the euphotic zone and thereby potentially stimulate microbial activity in regions of the upper offshore oxygen minimum zone.

Diatoms are among the few eukaryotes known to store nitrate (NO3- ) and to use it as an electron acceptor for respiration in the absence of light and O2 . Using microscopy and 15 N stable isotope incubations, we studied the relationship between dissimilatory nitrate/nitrite reduction to ammonium (DNRA) and diel vertical migration of diatoms in phototrophic microbial mats and the underlying sediment of a sinkhole in Lake Huron (USA). We found that the diatoms rapidly accumulated NO3- at the mat-water interface in the afternoon and 40% of the population migrated deep into the sediment, where they were exposed to dark and anoxic conditions for ~75% of the day. The vertical distribution of DNRA rates and diatom abundance maxima coincided, suggesting that DNRA was the main energy generating metabolism of the diatom population. We conclude that the illuminated redox-dynamic ecosystem selects for migratory diatoms that can store nitrate for respiration in the absence of light. A major implication of this study is that the dominance of DNRA over denitrification is not explained by kinetics or thermodynamics. Rather, the dynamic conditions select for migratory diatoms that perform DNRA and can outcompete sessile denitrifiers.

The western Indian continental shelf houses the world's largest naturally formed coastal low-oxygen zone that develops seasonally during the summer monsoon. We investigated multiple reductive nitrogen transformation pathways and quantified their rates in this system through anaerobic incubations with additions of N-15-labeled substrates during the anoxic period for three consecutive years (2008-2010). Addition of N-15 labeled ammonium ((NH4+)-N-15) resulted in low to moderate anaerobic ammonia oxidation (Anammox) rates in about half of our incubations from the oxygen depleted waters. In contrast, incubations with labeled nitrite ((NO2-)-N-15) led to large production of N-30(2) over N-29(2) in all incubation experiments, indicating denitrification to be the dominant N-loss pathway. Rates of dissimilatory nitrate/nitrite reduction to ammonium (DNRA) were found to be highly variable and were lower by an order of magnitude than the denitrification rates. Extrapolation of average rates over the sampling periods and volume of anoxic waters showed large nitrogen removal (3.70-11.1 Tg year(-1)) which is about three times as high as the previously reported estimate (1.3-3.8 Tg year(-1)). Despite the small area it occupies, this shallow seasonal anoxic zone may account for as much as 20-60% the of the total annual fixed nitrogen loss in the perennial oxygen minimum zone of the Arabian Sea.

The pool of dissolved organic matter (DOM) in the deep ocean represents one of the largest carbon sinks on the planet. In recent years, studies have shown that most of this pool is recalcitrant, because individual compounds are present at low concentrations and because certain compounds seem resistant to microbial degradation. The formation of the diverse and recalcitrant deep ocean DOM pool has been attributed to repeated and successive processing of DOM by microorganisms over time scales of weeks to years. Little is known however, about the transformation and cycling that labile DOM undergoes in the first hours upon its release from phytoplankton. Here we provide direct experimental evidence showing that within hours of labile DOM release, its breakdown and recombination with ambient DOM leads to the formation of a diverse array of new molecules in oligotrophic North Atlantic surface waters. Furthermore, our results reveal a preferential breakdown of N and P containing molecules versus those containing only carbon. Hence, we show the preferential breakdown and molecular diversification are the crucial first steps in the eventual formation of carbon rich DOM that is resistant to microbial remineralization.

Populations of genetically identical cells can display marked variation in phenotypic traits; such variation is termed phenotypic heterogeneity. Here, we investigate the effect of substrate and electron donor limitation on phenotypic heterogeneity in N2 and CO2 fixation in the green sulphur bacterium Chlorobium phaeobacteroides. We grew populations in chemostats and batch cultures and used stable isotope labelling combined with nanometer-scale secondary ion mass spectrometry (NanoSIMS) to quantify phenotypic heterogeneity. Experiments in H2 S (i.e. electron donor) limited chemostats show that varying levels of NH4+ limitation induce heterogeneity in N2 fixation. Comparison of phenotypic heterogeneity between chemostats and batch (unlimited for H2 S) populations indicates that electron donor limitation drives heterogeneity in N2 and CO2 fixation. Our results demonstrate that phenotypic heterogeneity in a certain metabolic activity can be driven by different modes of limitation and that heterogeneity can emerge in different metabolic processes upon the same mode of limitation. In conclusion, our data suggest that limitation is a general driver of phenotypic heterogeneity in microbial populations.

The N2 -fixing (diazotrophic) community in marine ecosystems is dominated by non-cyanobacterial microorganisms. Yet, very little is known about their identity, function and ecological relevance due to a lack of cultured representatives. Here we report a novel heterotrophic diazotroph isolated from the oxygen minimum zone (OMZ) off Peru. The new species belongs to the genus Sagittula (Rhodobacteraceae, Alphaproteobacteria) and its capability to fix N2 was confirmed in laboratory experiments. Genome sequencing revealed that it is a strict heterotroph with a high versatility in substrate utilization and energy acquisition mechanisms. Pathways for sulfide oxidation and nitrite reduction to nitrous oxide are encoded in the genome and might explain the presence throughout the Peruvian OMZ. The genome further indicates that this novel organism could be in direct interaction with other microbes or particles. NanoSIMS analyses were used to compare the metabolic potential of S. castanea with single-cell activity in situ; however, N2 fixation by this diazotroph could not be detected at the isolation site. While the biogeochemical impact of S. castanea is yet to be resolved, its abundance and widespread distribution suggests that its potential to contribute to the marine N input could be significant at a larger geographical scale.

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 eukaryotes1,2,3,4. Representatives of the Asgard candidate phylum Candidatus Lokiarchaeota (Lokiarchaeon) have the capacity for acetogenesis and fermentation5,6,7, 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 proposed Asgard superphylum of Archaea comprises the closest archaeal relatives of eukaryotes, whose genomes hold clues pertaining to the nature host cell that acquired the mitochondrion at the origin of eukaryotes1-4. Genomes of the Asgard candidate Phylum ‘CandidatusLokiarchaeota’ [Lokiarchaeon] suggest an acetogenic H2–dependent lifestyle5and mixotrophic capabilities6. However, data on the activity of Lokiarchaeon are currently lacking, and the ecology of the host cell that acquired the mitochondrion is debated4,7. Here, we show that in anoxic marine sediments underlying highly productive waters on the Namibian continental shelf Lokiarchaeon gene expression increases with depth below the seafloor, and was significantly different across a redox gradient spanning hypoxic to sulfidic conditions. Notably, Lokiarchaeon increased expression of genes involved in growth, carbohydrate metabolism, and the H2-dependent Wood-Ljungdahl (WLP) carbon fixation pathway under the most reducing (sulfidic) conditions. Quantitative stable isotope probing experiments revealed multiple populations of Lokiarchaeota utilizing both CO2and diatomaceous extracellular polymeric substances (dEPS) as carbon sources over a 10-day incubation under anoxic conditions. This apparent anaerobic mixotrophic metabolism was consistent with the expression of Lokiarchaeon genes involved in transport and fermentation of sugars and amino acids. Remarkably, several Asgard populations were more enriched with13C-dEPS compared to the community average, indicating a preference for dEPS as a growth substrate. The qSIP and gene expression data indicate a metabolism of “CandidatusLokiarchaeota” similar to that of sugar fermenting homoacetogenic bacteria8, namely that Lokiarchaeon can couple fermentative H2production from organic substrates with electron bifurcation and the autotrophic and H2-dependent WLP. Homoacetogenesis allows to access a wide range of substrates and relatively high ATP gain during acetogenic sugar fermentation8thereby providing an ecological advantage for Lokiarchaeon in anoxic, energy limited settings.

The 15N isotope pairing technique is widely used to quantify anammox, denitrification, and dissimilatory nitrate reduction to ammonium (DNRA) in sediments. However, the effects of DNRA on anammox and denitrification in slurry incubations and on genuine N2 production in intact core incubations have not been fully explored. We developed a mathematical model describing these effects of DNRA, and tested this model in field and computational studies. Our model calculations revealed that for slurry incubations the presence of DNRA tends to under -and overestimate the actual anammox and denitrification rates, respectively, when calculated according to Thamdrup and Dalsgaard (2002). We found this underestimate of anammox to be proportional to the 15 NH4+ mole fraction (FA), and the overestimate of denitrification to be related to both FA and the relative contribution of anammox to the total N2 production (ra). We propose three alternative procedures to better quantify anammox and denitrification rates in slurry incubations and two procedures for intact core incubations. Our model calculations also revealed that for intact core incubations, the presence of DNRA leads to an overestimate of the genuine N2 production rate when calculated according to Risgaard-Petersen et al. (2003). This overestimate depends on the ra and the mole fractions of 15 NH4+ and 15 NO3− in intact core incubations. The results of our field experiments and numerical modeling indicated that the overestimate of the genuine N2 production was less than 1% at two sites in the East China Sea.

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.

Despite decades of research on the physiology and biochemistry of nitrate/nitrite-respiring microorganisms, little is known regarding their metabolic response to temperature, especially under in situ conditions. The temperature regulation of microbial communities that mediate anammox and denitrification was investigated in near shore permeable sediments at polar, temperate, and subtropical sites with annual mean temperatures ranging from −5 to 23 °C. Total N2 production rates were determined using the isotope pairing technique in intact core incubations under diffusive and simulated advection conditions and ranged from 2 to 359 μmol N m−2 d−1. For the majority of sites studied, N2 removal was 2 to 7 times more rapid under advective flow conditions. Anammox comprised 6 to 14% of total N2 production at temperate and polar sites and was not detected at the subtropical site. Potential rates of denitrification and anammox were determined in anaerobic slurries in a temperature gradient block incubator across a temperature range of −1 to 42 °C. The highest optimum temperature (Topt) for denitrification was 36 °C and was observed in subtropical sediments, while the lowest Topt of 21 °C was observed at the polar site. Seasonal variation in the Topt was observed at the temperate site with values of 26 and 34 °C in winter and summer, respectively. The Topt values for anammox were 9 and 26 °C at the polar and temperate sites, respectively. The results demonstrate adaptation of denitrifying communities to in situ temperatures in permeable marine sediments across a wide range of temperatures, whereas marine anammox bacteria may be predominately psychrophilic to psychrotolerant. To our knowledge, we provide the first rates of denitrification and anammox from permeable sediments of a polar permanently cold ecosystem. The adaptation of microbial communities to in situ temperatures suggests that the relationship between temperature and rates of N removal is highly dependent on community structure.

In the Arctic Ocean, increased sea surface temperature and sea ice retreat have triggered shifts in phytoplankton communities. In Fram Strait, coccolithophorids have been occasionally observed to replace diatoms as the dominating taxon of spring blooms. Deep-sea benthic communities depend strongly on such blooms, but with a change in quality and quantity of primarily produced organic matter (OM) input, this may likely have implications for deep-sea life. We compared the in situ responses of Arctic deep-sea benthos to input of phytodetritus from a diatom (Thalassiosira sp.) and a coccolithophorid (Emiliania huxleyi) species. We traced the fate of 13C- and 15N-labelled phytodetritus into respiration, assimilation by bacteria and infauna in a 4-day and 14-day experiment. Bacteria were key assimilators in the Thalassiosira OM degradation, whereas Foraminifera and other infauna were at least as important as bacteria in the Emiliania OM assimilation. After 14 days, 5 times less carbon and 3.8 times less nitrogen of the Emiliania detritus was recycled compared to Thalassiosira detritus. This implies that the utilization of Emiliania OM may be less efficient than for Thalassiosira OM. Our results indicate that a shift from diatom-dominated input to a coccolithophorid-dominated pulse could entail a delay in OM cycling, which may affect benthopelagic coupling.

We investigated microbial pathways of nitrogen transformation in highly permeable sediments from the German Bight (South-East North Sea) by incubating sediment cores percolated with 15N-labeled substrates under near in situ conditions. In incubations with added math formula, production of math formula occurred while the sediment was oxic, indicating ammonia oxidation. Similarly, math formula production during math formula incubations indicated nitrite oxidation. Taken together these findings provide direct evidence of high nitrification rates within German Bight sands. The production of 15N-N2 on addition of math formula revealed high denitrification rates within the sediment under oxic and anoxic conditions. Denitrification rates were strongly and positively correlated with oxygen consumption rates, suggesting that denitrification is controlled by organic matter availability. Nitrification and denitrification rates were of the same magnitude and the rapid production of 15N-N2 in incubations with added math formula confirmed close coupling of the two processes. Areal rates of N-transformation were estimated taking advective transport of substrates into account and integrating volumetric rates over modeled oxygen and nitrate penetration depths, these ranged between 22 μmol N m−2 h−1 and 94 μmol N m−2 h−1. Furthermore, results from the 15N-labeling experiments show that these subtidal permeable sediments are, in sharp contrast to common belief, a substantial source of N2O. Our combined results show that nitrification fuels denitrification by providing an additional source of nitrate, and as such masks true N-losses from these highly eutrophic sediments. Given the widespread occurrence of anthropogenically influenced permeable sediments, coupled benthic nitrification–denitrification might have an important but so far neglected role in N-loss from shelf sediments.

The formation of a subsurface anticyclonic eddy in the Peru-Chile Undercurrent (PCUC) in January and February 2013 is investigated using a multi-platform four-dimensional observational approach. Research vessel, multiple glider and mooring-based measurements were conducted in the Peruvian upwelling regime near 12°30'S. The dataset consists of > 10000 glider profiles and repeated vessel-based hydrography and velocity transects. It allows a detailed description of the eddy formation and its impact on the near-coastal salinity, oxygen and nutrient distributions. In early January, a strong PCUC with maximum poleward velocities of ∼ 0.25 m/s at 100 to 200 m depth was observed. Starting on January 20 a subsurface anticyclonic eddy developed in the PCUC downstream of a topographic bend, suggesting flow separation as the eddy formation mechanism. The eddy core waters exhibited oxygen concentrations

Summary The gutless marine worm O lavius algarvensis lives in symbiosis with chemosynthetic bacteria that provide nutrition by fixing carbon dioxide (CO 2) into biomass using reduced sulfur compounds as energy sources. A recent metaproteomic analysis of the O . algarvensis symbiosis indicated that carbon monoxide (CO) and hydrogen (H 2) might also be used as energy sources. We provide direct evidence that the O . algarvensis symbiosis consumes CO and H 2. Single cell imaging using nanoscale secondary ion mass spectrometry revealed that one of the symbionts, the γ3‐symbiont, uses the energy from CO oxidation to fix CO 2. Pore water analysis revealed considerable in‐situ concentrations of CO and H 2 in the O . algarvensis environment, Mediterranean seagrass sediments. Pore water H 2 concentrations (89–2147 nM) were up to two orders of magnitude higher than in seawater, and up to 36‐fold higher than previously known from shallow‐water marine sediments. Pore water CO concentrations (17–51 nM) were twice as high as in the overlying seawater (no literature data from other shallow‐water sediments are available for comparison). Ex‐situ incubation experiments showed that dead seagrass rhizomes produced large amounts of CO. CO production from decaying plant material could thus be a significant energy source for microbial primary production in seagrass sediments.

Chain-forming diatoms are key CO2-fixing organisms in the ocean. Under turbulent conditions they form fast-sinking aggregates that are exported from the upper sunlit ocean to the ocean interior. A decade-old paradigm states that primary production in chain-forming diatoms is stimulated by turbulence. Yet, direct measurements of cell-specific primary production in individual field populations of chain-forming diatoms are poorly documented. Here we measured cell-specific carbon, nitrate and ammonium assimilation in two field populations of chain-forming diatoms (Skeletonema and Chaetoceros) at low-nutrient concentrations under still conditions and turbulent shear using secondary ion mass spectrometry combined with stable isotopic tracers and compared our data with those predicted by mass transfer theory. Turbulent shear significantly increases cell-specific C assimilation compared to still conditions in the cells/chains that also form fast-sinking, aggregates rich in carbon and ammonium. Thus, turbulence simultaneously stimulates small-scale biological CO2 assimilation and large-scale biogeochemical C and N cycles in the ocean., Chain-forming diatoms are key organisms in the biotic transfer of CO2 from the atmosphere to the ocean interior. Here, the authors show that turbulence stimulates and links small-scale and large scale processes from CO2 assimilation at a diatom cell level to nitrogen cycling in fast-sinking diatom aggregates.

Coastal oceans receive large amounts of anthropogenic fixed nitrogen (N), most of which is denitrified in the sediment before reaching the open ocean. Sandy sediments, which are common in coastal regions, seem to play an important role in catalysing this N‐loss. Permeable sediments are characterized by advective porewater transport, which supplies high fluxes of organic matter into the sediment, but also leads to fluctuations in oxygen and nitrate concentrations. Little is known about how the denitrifying communities in these sediments are adapted to such fluctuations. Our combined results indicate that denitrification in eutrophied sandy sediments from the world's largest tidal flat system, the Wadden Sea, is carried out by different groups of microorganisms. This segregation leads to the formation of N2O which is advectively transported to the overlying waters and thereby emitted to the atmosphere. At the same time, the production of N2O within the sediment supports a subset of Flavobacteriia which appear to be specialized on N2O reduction. If the mechanisms shown here are active in other coastal zones, then denitrification in eutrophied sandy sediments may substantially contribute to current marine N2O emissions. ISSN:1462-2912 ISSN:1462-2920

Dissolved iron (DFe) concentrations in oxygen minimum zones (OMZs) of Eastern Boundary Upwelling Systems are enhanced as a result of high supply rates from anoxic sediments. However, pronounced variations in DFe concentrations in anoxic coastal waters of the Peruvian OMZ indicate that there are factors in addition to dissolved oxygen concentrations (O2) that control Fe cycling. Our study demonstrates that sediment-derived reduced Fe (Fe(II)) forms the main DFe fraction in the anoxic/euxinic water column off Peru, which is responsible for DFe accumulations of up to 200 nmol L−1. Lowest DFe values were observed in anoxic shelf waters in the presence of nitrate and nitrite. This reflects oxidation of sediment-sourced Fe(II) associated with nitrate/nitrite reduction and subsequent removal as particulate Fe(III) oxyhydroxides. Unexpectedly, the highest DFe levels were observed in waters with elevated concentrations of hydrogen sulfide (up to 4 µmol L−1) and correspondingly depleted nitrate/nitrite concentrations (

Before the Earth's complete oxygenation (0.58 to 0.55 billion years [Ga] ago), the photic zone of the Proterozoic oceans was probably redox stratified, with a slightly aerobic, nutrient-limited upper layer above a light-limited layer that tended toward euxinia. In such oceans, cyanobacteria capable of both oxygenic and sulfide-driven anoxygenic photosynthesis played a fundamental role in the global carbon, oxygen, and sulfur cycle. We have isolated a cyanobacterium, Pseudanabaena strain FS39, in which this versatility is still conserved, and we show that the transition between the two photosynthetic modes follows a surprisingly simple kinetic regulation controlled by this organism's affinity for H 2 S. Specifically, oxygenic photosynthesis is performed in addition to anoxygenic photosynthesis only when H 2 S becomes limiting and its concentration decreases below a threshold that increases predictably with the available ambient light. The carbon-based growth rates during oxygenic and anoxygenic photosynthesis were similar. However, Pseudanabaena FS39 additionally assimilated NO 3 − during anoxygenic photosynthesis. Thus, the transition between anoxygenic and oxygenic photosynthesis was accompanied by a shift of the C/N ratio of the total bulk biomass. These mechanisms offer new insights into the way in which, despite nutrient limitation in the oxic photic zone in the mid-Proterozoic oceans, versatile cyanobacteria might have promoted oxygenic photosynthesis and total primary productivity, a key step that enabled the complete oxygenation of our planet and the subsequent diversification of life.

Whether large sulfur bacteria of the family Beggiatoaceae reduce NO3− to N2 via denitrification or to NH4+ via DNRA has been debated in the literature for more than 25 years. We resolve this debate by showing that certain members of the Beggiatoaceae use both metabolic pathways. This is important for the ecological role of these bacteria, as N2 production removes bioavailable nitrogen from the ecosystem, whereas NH4+ production retains it. For this reason, the topic of environmental controls on the competition for NO3− between N2-producing and NH4+-producing bacteria is of great scientific interest. Recent experiments on the competition between these two types of microorganisms have demonstrated that the balance between electron donor and electron acceptor availability strongly influences the end product of NO3− reduction. Our results suggest that this is also the case at the even more fundamental level of enzyme system regulation within a single organism., Filamentous large sulfur-oxidizing bacteria (FLSB) of the family Beggiatoaceae are globally distributed aquatic bacteria that can control geochemical fluxes from the sediment to the water column through their metabolic activity. FLSB mats from hydrothermal sediments of Guaymas Basin, Mexico, typically have a “fried-egg” appearance, with orange filaments dominating near the center and wider white filaments at the periphery, likely reflecting areas of higher and lower sulfide fluxes, respectively. These FLSB store large quantities of intracellular nitrate that they use to oxidize sulfide. By applying a combination of 15N-labeling techniques and genome sequence analysis, we demonstrate that the white FLSB filaments were capable of reducing their intracellular nitrate stores to both nitrogen gas and ammonium by denitrification and dissimilatory nitrate reduction to ammonium (DNRA), respectively. On the other hand, our combined results show that the orange filaments were primarily capable of DNRA. Microsensor profiles through a laboratory-incubated white FLSB mat revealed a 2- to 3-mm vertical separation between the oxic and sulfidic zones. Denitrification was most intense just below the oxic zone, as shown by the production of nitrous oxide following exposure to acetylene, which blocks nitrous oxide reduction to nitrogen gas. Below this zone, a local pH maximum coincided with sulfide oxidation, consistent with nitrate reduction by DNRA. The balance between internally and externally available electron acceptors (nitrate) and electron donors (reduced sulfur) likely controlled the end product of nitrate reduction both between orange and white FLSB mats and between different spatial and geochemical niches within the white FLSB mat. IMPORTANCE Whether large sulfur bacteria of the family Beggiatoaceae reduce NO3− to N2 via denitrification or to NH4+ via DNRA has been debated in the literature for more than 25 years. We resolve this debate by showing that certain members of the Beggiatoaceae use both metabolic pathways. This is important for the ecological role of these bacteria, as N2 production removes bioavailable nitrogen from the ecosystem, whereas NH4+ production retains it. For this reason, the topic of environmental controls on the competition for NO3− between N2-producing and NH4+-producing bacteria is of great scientific interest. Recent experiments on the competition between these two types of microorganisms have demonstrated that the balance between electron donor and electron acceptor availability strongly influences the end product of NO3− reduction. Our results suggest that this is also the case at the even more fundamental level of enzyme system regulation within a single organism.

Dissolved iron (DFe) concentrations in oxygen minimum zones (OMZs) of Eastern Boundary Upwelling Systems are enhanced as a result of high supply rates from anoxic sediments. However, pronounced variations in DFe concentrations in anoxic coastal waters of the Peruvian OMZ indicate that there are factors in addition to dissolved oxygen concentrations (O2) that control Fe cycling. Our study demonstrates that sediment-derived reduced Fe (Fe(II)) forms the main DFe fraction in the anoxic/euxinic water column off Peru, which is responsible for DFe accumulations of up to 200 nmol L−1. Lowest DFe values were observed in anoxic shelf waters in the presence of nitrate and nitrite. This reflects oxidation of sediment-sourced Fe(II) associated with nitrate/nitrite reduction and subsequent removal as particulate Fe(III) oxyhydroxides. Unexpectedly, the highest DFe levels were observed in waters with elevated concentrations of hydrogen sulfide (up to 4 µmol L−1) and correspondingly depleted nitrate/nitrite concentrations (

The N

Summary The temperature dependency of denitrification and anaerobic ammonium oxidation (anammox) rates from Arctic fjord sediments was investigated in a temperature gradient block incubator for temperatures ranging from −1 to 40°C. Community structure in intact sediments and slurry incubations was determined using Illumina SSU rRNA gene sequencing. The optimal temperature (Topt) for denitrification was 25–27°C, whereas anammox rates were optimal at 12–17°C. Both denitrification and anammox exhibited temperature responses consistent with a psychrophilic community, but anammox bacteria may be more specialized for psychrophilic activity. Long-term (1–2 months) warming experiments indicated that temperature increases of 5–10°C above in situ had little effect on the microbial community structure or the temperature response of denitrification and anammox. Increases of 25°C shifted denitrification temperature responses to mesophilic with concurrent community shifts, and anammox activity was eliminated above 25°C. Additions of low molecular weight organic substrates (acetate and lactate) caused increases in denitrification rates, corroborating the hypothesis that the supply of organic substrates is a more dominant control of respiration rates than low temperature. These results suggest that climate-related changes in sinking particulate flux will likely alter rates of N removal more rapidly than warming.

The release of organic material upon algal cell lyses has a key role in structuring bacterial communities and affects the cycling of biolimiting elements in the marine environment. Here we show that already before cell lysis the leakage or excretion of organic matter by infected yet intact algal cells shaped North Sea bacterial community composition and enhanced bacterial substrate assimilation. Infected algal cultures of Phaeocystis globosa grown in coastal North Sea water contained gamma-and alphaproteobacterial phylotypes that were distinct from those in the non-infected control cultures 5 h after infection. The gammaproteobacterial population at this time mainly consisted of Alteromonas sp. cells that were attached to the infected but still intact host cells. Nano-scale secondary-ion mass spectrometry (nanoSIMS) showed similar to 20% transfer of organic matter derived from the infected C-13- and N-15-labelled P. globosa cells to Alteromonas sp. cells. Subsequent, viral lysis of P. globosa resulted in the formation of aggregates that were densely colonised by bacteria. Aggregate dissolution was observed after 2 days, which we attribute to bacteriophage-induced lysis of the attached bacteria. Isotope mass spectrometry analysis showed that 40% of the particulate C-13-organic carbon from the infected P. globosa culture was remineralized to dissolved inorganic carbon after 7 days. These findings reveal a novel role of viruses in the leakage or excretion of algal biomass upon infection, which provides an additional ecological niche for specific bacterial populations and potentially redirects carbon availability.

Summary Hydrothermal sediments in the Guaymas Basin are covered by microbial mats that are dominated by nitrate-respiring and sulphide-oxidizing Beggiatoa. The presence of these mats strongly correlates with sulphide- and ammonium-rich fluids venting from the subsurface. Because ammonium and oxygen form opposed gradients at the sediment surface, we hypothesized that nitrification is an active process in these Beggiatoa mats. Using biogeochemical and molecular methods, we measured nitrification and determined the diversity and abundance of nitrifiers. Nitrification rates ranged from 74 to 605 μmol N l−1 mat day−1, which exceeded those previously measured in hydrothermal plumes and other deep-sea habitats. Diversity and abundance analyses of archaeal and bacterial ammonia monooxygenase subunit A genes, archaeal 16S ribosomal RNA pyrotags and fluorescence in situ hybridization confirmed that ammonia- and nitrite-oxidizing microorganisms were associated with Beggiatoa mats. Intriguingly, we observed cells of bacterial and potential thaumarchaeotal ammonia oxidizers attached to narrow, Beggiatoa-like filaments. Such a close spatial coupling of nitrification and nitrate respiration in mats of large sulphur bacteria is novel and may facilitate mat-internal cycling of nitrogen, thereby reducing loss of bioavailable nitrogen in deep-sea sediments.