ALBERT

Description: The eastern tropical North Atlantic (ETNA) is characterized by a highly productive coastal upwelling system and a moderate oxygen minimum zone with lowest open ocean oxygen (O2) concentrations of around 40 μmol kg−1. Only recently, the discovery of re-occurring mesoscale eddies with sometimes close to anoxic O2 concentrations (〈1 μmol kg−1) and located just below the mixed layer challenged our understanding of O2 distribution and biogeochemical processes in this area. Here, we present the first metagenomic dataset from a deoxygenated anticyclonic modewater eddy in the open waters of the ETNA. In the eddy, we observed a significantly lower bacterial diversity compared to surrounding waters, along with a significant community shift. We detected enhanced primary productivity in the surface layer of the eddy indicated by elevated chlorophyll concentrations and increased carbon uptake rates up to three times as high as in surrounding waters. Carbon uptake below the euphotic zone correlated to the presence of a specific high-light ecotype of Prochlorococcus, which is usually underrepresented in the ETNA. Our combined data indicate that high primary production in the eddy fuels export production and the presence of a specific microbial community responsible for enhanced respiration at shallow depths, below the mixed layer base. Progressively decreasing O2 concentrations in the eddy were found to promote transcription of the key gene for denitrification, nirS, in the O2-depleted core waters. This process is usually absent from the open ETNA waters. In the light of future ocean deoxygenation our results show exemplarily that even distinct events of anoxia have the potential to alter microbial community structures and with that critically impact primary productivity and biogeochemical processes of oceanic water bodies.

Description: Mesoscale eddies play a major role in controlling ocean biogeochemistry. By impacting nutrient availability and water column ventilation, they are of critical importance for oceanic primary production. In the eastern tropical South Pacific Ocean off Peru, where a large and persistent oxygen deficient zone is present, mesoscale processes have been reported to occur frequently. However, investigations on their biological activity are mostly based on model simulations, and direct measurements of carbon and dinitrogen (N2) fixation are scarce. We examined an open ocean cyclonic eddy and two anticyclonic mode water eddies: a coastal one and an open ocean one in the waters off Peru along a section at 16° S in austral summer 2012. Molecular data and bioassay incubations point towards a difference between the active diazotrophic communities present in the cyclonic eddy and the anticyclonic mode water eddies. In the cyclonic eddy, highest rates of N2 fixation were measured in surface waters but no N2 fixation signal was detected at intermediate water depths. In contrast, both anticyclonic mode water eddies showed pronounced maxima in N2 fixation below the euphotic zone as evidenced by rate measurements and geochemical data. N2 fixation and carbon (C) fixation were higher in the young coastal mode water eddy compared to the older offshore mode water eddy. A co-occurrence between N2 fixation and biogenic N2, an indicator for N loss, indicated a link between N loss and N2 fixation in the mode water eddies, which was not observed for the cyclonic eddy. The comparison of two consecutive surveys of the coastal mode water eddy in November and December 2012 revealed also a reduction of N2 and C fixation at intermediate depths along with a reduction in chlorophyll by half, mirroring an aging effect in this eddy. Our data indicate an important role for anticyclonic mode water eddies in stimulating N2 fixation and thus supplying N offshore.

Description: The recent finding that microbial ammonia oxidation in the ocean is performed by archaea to a greater extent than by bacteria has drastically changed the view on oceanic nitrification. The numerical dominance of archaeal ammonia-oxidizers (AOA) over their bacterial counterparts (AOB) in large parts of the ocean leads to the hypothesis that AOA rather than AOB could be the key organisms for the oceanic production of the strong greenhouse gas nitrous oxide (N2O) that occurs as a by-product of nitrification. Very recently, enrichment cultures of marine ammonia-oxidizing archaea have been reported to produce N2O. Here, we demonstrate that archaeal ammonia monooxygenase genes (amoA) were detectable throughout the water column of the eastern tropical North Atlantic (ETNA) and eastern tropical South Pacific (ETSP) Oceans. Particularly in the ETNA, comparable patterns of abundance and expression of archaeal amoA genes and N2O co-occurred in the oxygen minimum, whereas the abundances of bacterial amoA genes were negligible. Moreover, selective inhibition of archaea in seawater incubations from the ETNA decreased the N2O production significantly. In studies with the only cultivated marine archaeal ammonia-oxidizer Nitrosopumilus maritimus SCM1, we provide the first direct evidence for N2O production in a pure culture of AOA, excluding the involvement of other microorganisms as possibly present in enrichments. N. maritimus showed high N2O production rates under low oxygen concentrations comparable to concentrations existing in the oxycline of the ETNA, whereas the N2O production from two AOB cultures was comparably low under similar conditions. Based on our findings, we hypothesize that the production of N2O in tropical ocean areas results mainly from archaeal nitrification and will be affected by the predicted decrease in dissolved oxygen in the ocean.

Description: Recent observations in the eastern tropical South Pacific (ETSP) have shown the key role of meso- and submesoscale processes (e.g. eddies) in shaping its hydrographic and biogeochemical properties. Off Peru, elevated primary production from coastal upwelling in combination with sluggish ventilation of subsurface waters fuels a prominent oxygen minimum zone (OMZ). Given that nitrous oxide (N2O) production–consumption processes in the water column are sensitive to oxygen (O2) concentrations, the ETSP is a region of particular interest to investigate its source–sink dynamics. To date, no detailed surveys linking mesoscale processes and N2O distributions as well as their relevance to nitrogen (N) cycling are available. In this study, we present the first measurements of N2O across three mesoscale eddies (two mode water or anticyclonic and one cyclonic) which were identified, tracked, and sampled during two surveys carried out in the ETSP in November–December 2012. A two-peak structure was observed for N2O, wherein the two maxima coincide with the upper and lower boundaries of the OMZ, indicating active nitrification and partial denitrification. This was further supported by the abundances of the key gene for nitrification, ammonium monooxygenase (amoA), and the gene marker for N2O production during denitrification, nitrite reductase (nirS). Conversely, we found strong N2O depletion in the core of the OMZ (O2 〈 5 μmol/L) to be consistent with nitrite (NO2-) accumulation and low levels of nitrate (NO3-), thus suggesting active denitrification. N2O depletion within the OMZ’s core was substantially higher in the centre of mode water eddies, supporting the view that eddy activity enhances N-loss processes off Peru, in particular near the shelf break where nutrient-rich, productive waters from upwelling are trapped before being transported offshore. Analysis of eddies during their propagation towards the open ocean showed that, in general, “ageing” of mesoscale eddies tends to decrease N2O concentrations through the water column in response to the reduced supply of material to fuel N loss, although the hydrographic variability might also significantly impact the pace of the production–consumption pathways for N2O. Our results evidence the relevance of mode water eddies for N2O distribution, thereby improving our understanding of the N-cycling processes, which are of crucial importance in times of climate change and ocean deoxygenation.

Description: The Peruvian upwelling system is a highly productive ecosystem with a large oxygen minimum zone (OMZ) close to the surface. In this work, we carried out a mesocosm experiment off Callao, Peru, with the addition of water masses from the regional OMZ collected at two different sites simulating two different upwelling scenarios. Here, we focus on the pelagic remineralization of organic matter by the extracellular enzyme activity of leucine aminopeptidase (LAP) and alkaline phosphatase activity (APA). After the addition of the OMZ water, dissolved inorganic nitrogen (N) was depleted, but the standing stock of phytoplankton was relatively high, even after N depletion (mostly 〉 4 µg chlorophyll a L−1). During the initial phase of the experiment, APA was 0.6 nmol L−1 h−1 even though the PO concentration was 〉 0.5 µmol L−1. Initially, the dissolved organic phosphorus (DOP) decreased, coinciding with an increase in the PO concentration that was probably linked to the APA. The LAP activity was very high, with most of the measurements in the range of 200–800 nmol L−1 h−1. This enzyme hydrolyzes terminal amino acids from larger molecules (e.g., peptides or proteins), and these high values are probably linked to the highly productive but N-limited coastal ecosystem. Moreover, the experiment took place during a rare coastal El Niño event with higher than normal surface temperatures, which could have affected enzyme activity. Using a nonparametric multidimensional scaling analysis (NMDS) with a generalized additive model (GAM), we found that biogeochemical variables (e.g., nutrient and chlorophyll-a concentrations) and phytoplankton and bacterial communities explained up to 64 % of the variability in APA. The bacterial community best explained the variability (34 %) in LAP. The high hydrolysis rates for this enzyme suggest that pelagic N remineralization, likely driven by the bacterial community, supported the high standing stock of primary producers in the mesocosms after N depletion.