Nitrogen is an essential component of all living organisms and the main nutrient limiting life on our planet. By far, the largest inventory of freely accessible nitrogen is atmospheric dinitrogen, but most organisms rely on more bioavailable forms of nitrogen, such as ammonium and nitrate, for growth. The availability of these substrates depends on diverse nitrogen-transforming reactions that are carried out by complex networks of metabolically versatile microorganisms. In this Review, we summarize our current understanding of the microbial nitrogen-cycling network, including novel processes, their underlying biochemical pathways, the involved microorganisms, their environmental importance and industrial applications.

Introductory Ammonia-oxidizing archaea of the phylum Thaumarchaeota are among the most abundant marine microorganisms1. These organisms thrive in the oceans despite ammonium being present at low nanomolar concentrations2,3. Some Thaumarchaeota isolates have been shown to utilize urea and cyanate as energy and N-sources through intracellular conversion to ammonium4–6. Yet, it is unclear whether patterns observed in culture extend to marine Thaumarchaeota, and whether Thaumarchaeota in the ocean directly utilize urea and cyanate or rely on co-occurring microorganisms to break these substrates down to ammonium. Urea utilization has been reported for marine ammonia-oxidizing communities7–10, but no evidence of cyanate utilization exists for marine ammonia oxidizers. Here, we demonstrate that in the Gulf of Mexico, Thaumarchaeota use urea and cyanate both directly and indirectly as energy and N-sources. We observed substantial and linear rates of nitrite production from urea and cyanate additions, which often persisted even when ammonium was added to micromolar concentrations. Furthermore, single cell analysis revealed that the Thaumarchaeota incorporated ammonium-, urea- and cyanate-derived N at significantly higher rates than most other microorganisms. Yet, no cyanases were detected in thaumarchaeal genomic data from the Gulf of Mexico. Therefore, we tested cyanate utilization in Nitrosopumilus maritimus, which also lacks a canonical cyanase, and showed that cyanate was oxidized to nitrite. Our findings demonstrate that marine Thaumarchaeota can use urea and cyanate as both an energy and N-source. Based on these results we hypothesize that urea and cyanate are substrates for ammonia-oxidizing Thaumarchaeota throughout the ocean.

Abstract. Benthic nitrogen transformation pathways were investigated in the sediment of the East China Sea (ECS) in June of 2010 using the 15N isotope pairing technique. Slurry incubations indicated that denitrification, anammox and dissimilatory nitrate reduction to ammonium (DNRA) as well as intracellular nitrate release occurred in the ECS sediments. These four processes did not exist independently, nitrate release therefore diluted the 15N labeling fraction of NO3−, and a part of the 15NH4+ derived from DNRA also formed 30N2 via anammox. Therefore, current methods of rate calculations led to over and underestimations of anammox and denitrification respectively. Following the procedure outlined in Thamdrup and Dalsgaard (2002), denitrification rates were slightly underestimated by an average 6% without regard to the effect of nitrate release, while this underestimation could be counteracted by the presence of DNRA. On the contrary, anammox rates calculated from 15NO3− experiment were significantly overestimated by 42% without considering nitrate release. In our study, this overestimation could only be compensated 14% by taking DNRA into consideration. In a parallel experiment amended with 15NH4++14NO3−, anammox rates were not significantly influenced by DNRA due to the high background of 15NH4+ addition. The significant correlation between potential denitrification rate and sediment organic matter content (r = 0.68, p < 0.001, Pearson) indicated that denitrification was regulated by organic matter, while, no such correlations were found for anammox and DNRA. The relative contribution of anammox to the total N-loss increased from 13% at the shallowest site near the Changjiang estuary to 50% at the deepest site on the outer shelf, implying the significant role of anammox in benthic nitrogen cycling in the ECS sediments, especially on the outer shelf. N-loss as N2 was the main pathway, while DNRA was also an important pathway accounting for 20–31% of benthic nitrate reduction in the ECS. Our study demonstrates the complicated interactions among different benthic nitrogen transformations and the importance of considering denitrification, DNRA, anammox and nitrate release together when designing and interpreting future studies.

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 15 N-labeled substrates under near in situ conditions. In incubations with added 15 NH 3 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 N-labeling experiments show that these subtidal permeable sediments are, in sharp contrast to common belief, a substantial source of N 2 O. 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.

Nitrogen (N) input to the coastal oceans has increased considerably because of anthropogenic activities, however, concurrent increases have not occurred in open oceans. It has been suggested that benthic denitrification in sandy coastal sediments is a sink for this N. Sandy sediments are dynamic permeable environments, where electron acceptor and donor concentrations fluctuate over short temporal and spatial scales. The response of denitrifiers to these fluctuations are largely unknown, although previous observations suggest they may denitrify under aerobic conditions. We examined the response of benthic denitrification to fluctuating oxygen concentrations, finding that denitrification not only occurred at high O concentrations but was stimulated by frequent switches between oxic and anoxic conditions. Throughout a tidal cycle, in situtranscription of genes for aerobic respiration and denitrification were positively correlated within diverse bacterial classes, regardless of O concentrations, indicating that denitrification gene transcription is not strongly regulated by O in sandy sediments. This allows microbes to respond rapidly to changing environmental conditions, but also means that denitrification is utilized as an auxiliary respiration under aerobic conditions when imbalances occur in electron donor and acceptor supply. Aerobic denitrification therefore contributes significantly to N-loss in permeable sediments making the process an important sink for anthropogenic N-inputs.

Patterns of Pore Water Chemistry which produced non-uniform groundwater flow conditions. We conclude that on mesotidal high energy beaches, the rapidly changing beach morphology produces zones with different approximations to steady-state conditions. Therefore, we suggest that zone-specific endmember sampling is the optimal strategy to reduce uncertainties of SGD-driven constituent fluxes.

Coastal zones act as a sink for riverine and atmospheric nitrogen inputs and thereby buffer the open ocean from the effects of anthropogenic activity. Recently, microbial activity in sandy permeable sediments has been identified as a dominant source of N-loss in coastal zones, namely through denitrification. Some of the highest coastal denitrification rates measured so far occur within the intertidal permeable sediments of the eutrophied Wadden Sea. Still, denitrification alone can often account for only half of the substantial nitrate (NO3 −) consumption. Therefore, to investigate alternative NO3 − sinks such as dissimilatory nitrate reduction to ammonium (DNRA), intracellular nitrate storage by eukaryotes and isotope equilibration effects we carried out 15NO3 − amendment experiments. By considering all of these sinks in combination, we could quantify the fate of the 15NO3 − added to the sediment. Denitrification was the dominant nitrate sink (50–75%), while DNRA, which recycles N to the environment accounted for 10–20% of NO3 − consumption. Intriguingly, we also observed that between 20 and 40% of 15NO3 − added to the incubations entered an intracellular pool of NO3 − and was subsequently respired when nitrate became limiting. Eukaryotes were responsible for a large proportion of intracellular nitrate storage, and it could be shown through inhibition experiments that at least a third of the stored nitrate was subsequently also respired by eukaryotes. The environmental significance of the intracellular nitrate pool was confirmed by in situ measurements which revealed that intracellular storage can accumulate nitrate at concentrations six fold higher than the surrounding porewater. This intracellular pool is so far not considered when modeling N-loss from intertidal permeable sediments; however it can act as a reservoir for nitrate during low tide. Consequently, nitrate respiration supported by intracellular nitrate storage can add an additional 20% to previous nitrate reduction estimates in intertidal sediments, further increasing their contribution to N-loss.

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