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The ecophysiology of complete ammonia-oxidizing bacteria (CMX) of the genus Nitrospira and their widespread occurrence in groundwater suggests that CMX bacteria have a competitive advantage over ammonia-oxidizing bacteria (AOB) and archaea (AOA) in these environments. However, the specific contribution of their activity to nitrification processes has remained unclear. We aimed to disentangle the contribution of CMX, AOA and AOB to nitrification and to identify the environmental drivers of their niche differentiation at different levels of ammonium and oxygen in oligotrophic carbonate rock aquifers. CMX ammonia monooxygenase sub-unit A (amoA) genes accounted on average for 16 to 75% of the total groundwater amoA genes detected. Nitrification rates were positively correlated to CMX clade A associated phylotypes and AOB affiliated with Nitrosomonas ureae. Short-term incubations amended with the nitrification inhibitors allylthiourea and chlorate suggested that AOB contributed a large fraction to overall ammonia oxidation, while metaproteomics analysis confirmed an active role of CMX in both ammonia and nitrite oxidation. Ecophysiological niche differentiation of CMX clades A and B, AOB and AOA was linked to their requirements for ammonium, oxygen tolerance, and metabolic versatility. Our results demonstrate that despite numerical predominance of CMX, the first step of nitrification in oligotrophic groundwater appears to be primarily governed by AOB. Higher growth yields at lower ammonia turnover rates and energy derived from nitrite oxidation most likely enable CMX to maintain consistently high populations.

Ammonia-oxidizing archaea (AOA) are one of the most abundant groups of microbes in the world’s oceans and are key players in the nitrogen cycle. Their energy metabolism—the oxidation of ammonia to nitrite—requires oxygen. Nevertheless, AOA are abundant in environments where oxygen is undetectable. By carrying out incubations for which oxygen concentrations were resolved to the nanomolar range, we show that after oxygen depletion, Nitrosopumilus maritimus produces dinitrogen and oxygen, which is used for ammonia oxidation. The pathway is not completely resolved but likely has nitric oxide and nitrous oxide as key intermediates. N. maritimus joins a handful of organisms known to produce oxygen in the dark. On the basis of this ability, we reevaluate the role of N. maritimus in oxygen-depleted marine environments.

Trichodesmium is an important dinitrogen (N2)-fixing cyanobacterium in marine ecosystems. Recent nucleic acid analyses indicate that Trichodesmium colonies with their diverse epibionts support various nitrogen (N) transformations beyond N2 fixation. However, rates of these transformations and concentration gradients of N compounds in Trichodesmium colonies remain largely unresolved. We combined isotope-tracer incubations, micro-profiling and numeric modelling to explore carbon fixation, N cycling processes as well as oxygen, ammonium and nitrate concentration gradients in individual field-sampled Trichodesmium colonies. Colonies were net-autotrophic, with carbon and N2 fixation occurring mostly during the day. Ten percent of the fixed N was released as ammonium after 12-h incubations. Nitrification was not detectable but nitrate consumption was high when nitrate was added. The consumed nitrate was partly reduced to ammonium, while denitrification was insignificant. Thus, the potential N transformation network was characterised by fixed N gain and recycling processes rather than denitrification. Oxygen concentrations within colonies were ~60–200% air-saturation. Moreover, our modelling predicted steep concentration gradients, with up to 6-fold higher ammonium concentrations, and nitrate depletion in the colony centre compared to the ambient seawater. These gradients created a chemically heterogeneous microenvironment, presumably facilitating diverse microbial metabolisms in millimetre-sized Trichodesmium colonies.

Coastal sediments are rich in conductive particles, possibly affecting microbial processes for which acetate is a central intermediate. In the methanogenic zone, acetate is consumed by methanogens and/or syntrophic acetate-oxidizing (SAO) consortia. SAO consortia live under extreme thermodynamic pressure, and their survival depends on successful partnership. Here, we demonstrate that conductive particles enable the partnership between SAO bacteria (i.e., Geobacter spp.) and methanogens (Methanosarcina spp.) from the coastal sediments of the Bothnian Bay of the Baltic Sea. Baltic methanogenic sediments were rich in conductive minerals, had an apparent isotopic fractionation characteristic of CO2-reductive methanogenesis, and were inhabited by Geobacter and Methanosarcina. As long as conductive particles were delivered, Geobacter and Methanosarcina persisted, whereas exclusion of conductive particles led to the extinction of Geobacter. Baltic Geobacter did not establish a direct electric contact with Methanosarcina, necessitating conductive particles as electrical conduits. Within SAO consortia, Geobacter was an efficient [13C]acetate utilizer, accounting for 82% of the assimilation and 27% of the breakdown of acetate. Geobacter benefits from the association with the methanogen, because in the absence of an electron acceptor it can use Methanosarcina as a terminal electron sink. Consequently, inhibition of methanogenesis constrained the SAO activity of Geobacter as well. A potential benefit for Methanosarcina partnering with Geobacter is that together they competitively exclude acetoclastic methanogens like Methanothrix from an environment rich in conductive particles. Conductive particle-mediated SAO could explain the abundance of acetate oxidizers like Geobacter in the methanogenic zone of sediments where no electron acceptors other than CO2 are available.IMPORTANCE Acetate-oxidizing bacteria are known to thrive in mutualistic consortia in which H2 or formate is shu

Nitrogen cycling is normally thought to dominate the biogeochemistry and microbial ecology of oxygen-minimum zones in marine environments. Through a combination of molecular techniques and process rate measurements, we showed that both sulfate reduction and sulfide oxidation contribute to energy flux and elemental cycling in oxygen-free waters off the coast of northern Chile. These processes may have been overlooked because in nature, the sulfide produced by sulfate reduction immediately oxidizes back to sulfate. This cryptic sulfur cycle is linked to anammox and other nitrogen cycling processes, suggesting that it may influence biogeochemical cycling in the global ocean.

Allylthiourea (ATU) and chlorate (ClO3-) are often used to selectively inhibit nitritation and nitratation. In this work we identified challenges with use of these compounds in inhibitory assays with filter material from a biological rapid sand filter for groundwater treatment. Inhibition was investigated in continuous-flow lab-scale columns, packed with filter material from a full-scale filter and supplied with NH4+ or NO2-. ATU concentrations of 0.1-0.5 mM interfered with the indophenol blue method for NH4+ quantification leading to underestimation of the measured NH4+ concentration. Interference was stronger at higher ATU levels and resulted in no NH4+ detection at 0.5 mM ATU. ClO3- at typical concentrations for inhibition assays (1-10 mM) inhibited nitratation by less than 6%, while nitritation was instead inhibited by 91% when NH4+ was supplied. On the other hand, nitratation was inhibited by 67-71% at 10-20 mM ClO3- when NO2- was supplied, suggesting significant nitratation inhibition at higher NO2- concentrations. No chlorite (ClO2-) was detected in the effluent, and thus we could not confirm that nitritation inhibition was caused by ClO3- reduction to ClO2-. In conclusion, ATU and ClO3- should be used with caution in inhibition assays, because analytical interference and poor selectivity for the targeted process may affect the experimental outcome and compromise result interpretation.

Estuarine sediments are critical for the remediation of large amounts of anthropogenic nitrogen (N) loading via production of N2 from nitrate by denitrification. However, nitrate is also recycled within sediments by dissimilatory nitrate reduction to ammonium (DNRA). Understanding the factors that influence the balance between denitrification and DNRA is thus crucial to constraining coastal N budgets. A potentially important factor is the availability of different electron donors (organic carbon, reduced iron and sulfur). Both denitrification and DNRA may be linked to ferrous iron oxidation, however the contribution of Fe(II)-fueled nitrate reduction in natural environments is practically unknown. This study investigated how nitrate-dependent Fe2+ oxidation affects the partitioning between nitrate reduction pathways using 15N-tracing methods in sediments along the salinity gradient of the periodically hypoxic Yarra River estuary, Australia. Increased dissolved Fe2+ availability resulted in significant enhancement of DNRA rates from around 10–20% total nitrate reduction in control incubations to over 40% in those with additional Fe2+, at several sites. Increases in DNRA at some locations were accompanied by reductions in denitrification. Significant correlations were observed between Fe2+ oxidation and DNRA rates, with reaction ratios corresponding to the stoichiometry of Fe2+-dependent DNRA. Our results provide experimental evidence for a direct coupling of DNRA to Fe2+ oxidation across an estuarine gradient, suggesting that Fe2+ availability may exert substantial control on the balance between retention and removal of bioavailable N. Thus, DNRA linked to Fe2+ oxidation may be of general importance to environments with Fe-rich sediments.

Climate change is expected to have a pronounced effect on biogeochemical cycling in Arctic fjords, but current insight on the biogeochemical functioning of these systems is limited. Here, we present seasonal data on primary production, export of particulate organic carbon (POC), and the coupling to benthic biogeochemistry in Kobbefjord (SW Greenland). Primary production and associated POC export from the photic zone showed marked seasonality, with annual integrated values of 7.2 and 19.9 mol C m-2 yr-1, respectively. This discrepancy, the isotopic signature, and C:N ratio of the sedimentating material suggested substantial import of marine POC from outside the fjord. At least 52% of the POC export reached the sediment, but the seasonality in pelagic productivity was not reflected in the sediment biogeochemistry, showing only moderate variation. Benthic mineralization and burial of organic carbon amounted to 3.2 and 5.3 mol C m-2 yr-1, respectively. Sulfate reduction was the most prominent mineralization pathway, accounting for 69% of the benthic mineralization, while denitrification accounted for 2%. Overall, the carbon mineralization and burial in Kobbefjord were significantly higher than previously observed in other more northerly Arctic fjords. Data compilation from Arctic fjords suggests proportional increases in surface production, POC export, benthic mineralization and burial of organic material with increasing duration of the ice-free period. Thus, the projected decline in ice coverage in higher Arctic Greenlandic fjords will, as a first approximation, entail proportional increases in productivity, mineralization, and burial of organic carbon in the fjords, which will thus become similar to present-day southerly systems.

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The global marine nitrogen cycle is constrained by nitrogen fixation as a source of reactive nitrogen, and denitrification or anammox on the sink side. These processes with their respective isotope effects set the marine nitrate N-15-isotope value (delta N-15) to a relatively constant average of 5 parts per thousand. This value can be used to better assess the magnitude of these sources and sink terms, but the underlying assumption is that sedimentary denitrification and anammox, processes responsible for approximately one-third of global nitrogen removal, have little to no isotope effect on nitrate in the water column. We investigated the isotope fractionation in sediment incubations, measuring net denitrification and nitrogen and oxygen stable isotope fractionation in surface sediments from the coastal Baltic Sea (Boknis Eck, northern Germany), a site with seasonal hypoxia and dynamic nitrogen turnover. Sediment denitrification was fast, and regardless of current paradigms assuming little fractionation during sediment denitrification, we measured fractionation factors of 18.9 parts per thousand for nitrogen and 15.8 parts per thousand for oxygen in nitrate. While the input of nitrate to the water column remains speculative, these results challenge the current view of fractionation during sedimentary denitrification and imply that nitrogen budget calculations may need to consider this variability, as both preferential uptake of light nitrate and release of the remaining heavy fraction can significantly alter water column nitrate isotope values at the sediment-water interface.

Contains fulltext : 35120.pdf (publisher's version ) (Closed access), Laboratory and field studies have indicated that anaerobic ammonium oxidation (anammox) is an important process in the marine nitrogen cycle. In this study 11 additional anoxic marine sediment and water column samples were studied to substantiate this claim. In a combined approach using the molecular methods, polymerase chain reaction (PCR), qualitative and quantitative fluorescence in situ hybridization (FISH), as well as (15)N stable isotope activity measurements, it was shown that anammox bacteria were present and active in all samples investigated. The anammox activity measured in the sediment samples ranged from 0.08 fmol cell(-1) day(-1) N(2) in the Golfo Dulce (Pacific Ocean, Costa Rica) sediment to 0.98 fmol cell(-1) day(-1) N(2) in the Gullmarsfjorden (North Sea, Sweden) sediment. The percentage of anammox cell of the total population (stained with DAPI) as assessed by quantitative FISH was highest in the Barents Sea (9% +/- 4%) and in most of the samples well over 2%. Fluorescence in situ hybridization and phylogenetic analysis of the PCR products derived from the marine samples indicated the exclusive presence of members of the Candidatus'Scalindua' genus. This study showed the ubiquitous presence of anammox bacteria in anoxic marine ecosystems, supporting previous observations on the importance of anammox for N cycling in marine environments.