Amazon River infl uence on nitrogen fi xation in the western tropical North Atlantic

We measured rates of N- and C-fixation with a direct tracer method in regions of the western tropical North Atlantic influenced by the Amazon River plume during the high flow period of 2010 (May–June 2010). We found distinct regional variations in N-fixation activity, with the lowest rates in the plume proper and the highest rates in the plume margins and in offshore waters. A comparison of our N- and C-fixation measurements showed that the relative contribution of N-fixation to total primary production increased from the plume core toward oceanic waters, and that most of the C-fixation in this system was supported by sources of nitrogen other than those derived from biological N-fixation, or diazotrophy. We complemented these rate experiments with measurements of the δ15N of suspended particles (δ15PN), which documented the important and often dominant role of diazotrophs in supplying nitrogen to particulate organic matter in the water column. These coupled measurements revealed that small phytoplankton contributed more new nitrogen to the particulate nitrogen pool than larger phytoplankton. We used a habitat classification method to assess the fac- tors that control diazotrophic activity and contribution to the suspended particle pool, both of which increased from the plume toward oceanic waters. Our findings provide an important constraint on the role of the Amazon plume in creating distinct niches and roles for diazotrophs in the nutrient and carbon budgets of the western tropical North Atlantic

Upward nitrate transport by phytoplankton in oceanic waters : balancing nutrient budgets in oligotrophic seas

© The Author(s), 2014. This article is distributed under the terms of the Creative Commons Attribution License. The definitive version was published in PeerJ 2 (2014): e302, doi:10.7717/peerj.302.In oceanic subtropical gyres, primary producers are numerically dominated by small (1–5 µm diameter) pro- and eukaryotic cells that primarily utilize recycled nutrients produced by rapid grazing turnover in a highly efficient microbial loop. Continuous losses of nitrogen (N) to depth by sinking, either as single cells, aggregates or fecal pellets, are balanced by both nitrate inputs at the base of the euphotic zone and N2-fixation. This input of new N to balance export losses (the biological pump) is a fundamental aspect of N cycling and central to understanding carbon fluxes in the ocean. In the Pacific Ocean, detailed N budgets at the time-series station HOT require upward transport of nitrate from the nutricline (80–100 m) into the surface layer (∼0–40 m) to balance productivity and export needs. However, concentration gradients are negligible and cannot support the fluxes. Physical processes can inject nitrate into the base of the euphotic zone, but the mechanisms for transporting this nitrate into the surface layer across many 10s of m in highly stratified systems are unknown. In these seas, vertical migration by the very largest (102–103 µm diameter) phytoplankton is common as a survival strategy to obtain N from sub-euphotic zone depths. This vertical migration is driven by buoyancy changes rather than by flagellated movement and can provide upward N transport as nitrate (mM concentrations) in the cells. However, the contribution of vertical migration to nitrate transport has been difficult to quantify over the required basin scales. In this study, we use towed optical systems and isotopic tracers to show that migrating diatom (Rhizosolenia) mats are widespread in the N. Pacific Ocean from 140°W to 175°E and together with other migrating phytoplankton (Ethmodiscus, Halosphaera, Pyrocystis, and solitary Rhizosolenia) can mediate time-averaged transport of N (235 µmol N m-2 d-1) equivalent to eddy nitrate injections (242 µmol NO3− m-2 d-1). This upward biotic transport can close N budgets in the upper 250 m of the central Pacific Ocean and together with diazotrophy creates a surface zone where biological nutrient inputs rather than physical processes dominate the new N flux. In addition to these numerically rare large migrators, there is evidence in the literature of ascending behavior in small phytoplankton that could contribute to upward flux as well. Although passive downward movement has dominated models of phytoplankton flux, there is now sufficient evidence to require a rethinking of this paradigm. Quantifying these fluxes is a challenge for the future and requires a reexamination of individual phytoplankton sinking rates as well as methods for capturing and enumerating ascending phytoplankton in the sea.This work has been funded by the National Science Foundation: OCE-0726726, OCE-0094591, OCE-9414372, OCE-9100888 and OCE-9415923 to TAV, and OCE-9423471 to CHP

Trophic relationships and the nitrogen isotopic composition of amino acids in plankton

© 2002 by the Ecological Society of AmericaStable nitrogen isotope ratios of whole organisms and tissues are routinely used in studies of trophic relationships and nitrogen flow through ecosystems, yet changes underlying increases in δ¹⁵ N from food source to consumer are not completely understood. In this study, the δ¹⁵ N of 16 amino acids in marine planktonic consumers and their food sources were examined using gas chromatography/combustion/isotope ratio mass spectrometry of their N-pivaloyl-i-propyl-amino acid ester derivatives. Moderate increases in bulk δ¹⁵ N with trophic position reflect an averaging of large increases in the δ¹⁵ N of some amino acids, and little or no change in others. Amino acids showing consistently large increases (e.g., glutamic acid changes by ~7‰ between food and consumer) provide greater scope for defining trophic position than the smaller isotopic changes in bulk material. In contrast, amino acids like phenylalanine show no change in δ¹⁵ N with trophic position and therefore preserve information about nitrogen sources at the base of the food web. The ability to acquire information about both trophic level and nitrogen sources at the base of the food web from single samples of consumer tissues offers a powerful new tool for elucidating pathways of N transfer through food webs

Advancing science from plankton to whales—Celebrating the contributions of James J. McCarthy

Hailing from Sweet Home, Oregon, where his father introduced him to the fascinations of pondwater (McCarthy 2018), Jim McCarthy graduated from Gonzaga University, and in the late 1960s joined the Food Chain Research Group at the Scripps Institution of Oceanography, where he received his doctorate in 1971. The Food Chain Research Group, which was becoming recognized as the premier research group on plankton, was at that time directed by such distinguished scientists as John Strickland and Dick Eppley, among others. The goal of the Food Chain Group was to understand plankton dynamics and trophodynamics, “to a degree that will enable man to exercise satisfactory control of the environment and make useful predictions” (Institute of Marine Resources annual report, 1968, cited in Shor 1978:143) and “to predict the formation and transfer of nutrients through the full cycle of life in the ocean” (Shor 1978:140). It was there that Jim became immersed in all aspects of nutrients, plankton, and the marine food web

Vertical Marine Snow Distribution in the Stratified Hypersaline, and Anoxic Orca Basin (Gulf of Mexico)

We present a complete description of the depth distribution of marine snow in Orca Basin (Gulf of Mexico), from sea surface through the pycnocline to within 10 m of the seafloor. Orca Basin is an intriguing location for studying marine snow because of its unique geological and hydrographic setting: the deepest ~200 m of the basin are filled with anoxic hypersaline brine. A typical deep ocean profile of marine snow distribution was observed from the sea surface to the pycnocline, namely a surface maximum in total particle number and midwater minimum. However, instead of a nepheloid (particle-rich) layer positioned near the seabed, the nepheloid layer in the Orca Basin was positioned atop the brine. Within the brine, the total particle volume increased by a factor of 2–3 while the total particle number decreased, indicating accumulation and aggregation of material in the brine. From these observations we infer increased residence time and retention of material within the brine, which agrees well with laboratory results showing a 2.2–3.5-fold reduction in settling speed of laboratory-generated marine snow below the seawater-brine interface. Similarly, dissolved organic carbon concentration in the brine correlated positively with measured colored dissolved organic matter (r2 = 0.92, n = 15), with both variables following total particle volume inversely through the pycnocline. These data indicate the release of dissolved organic carbon concomitant with loss in total particle volume and increase in particle numbers at the brine-seawater interface, highlighting the importance of the Orca Basin as a carbon sink

New Perspectives on Nitrogen Fixation Measurements Using ¹⁵N₂ Gas

International audienc

New Perspectives on Nitrogen Fixation Measurements Using 15N2 Gas

Recently, the method widely used to determine 15N2 fixation rates in marine and freshwater environments was found to underestimate rates because the dissolution of the added 15N2 gas bubble in seawater takes longer than theoretically calculated. As a solution to the potential underestimate of rate measurements, the usage of the enriched water method was proposed to provide constant 15N2 enrichment. Still, the superiority of enriched water method over the previously used bubble injection remains inconclusive. To clarify this issue, we performed laboratory based experiments and implemented the results into an error analysis of 15N2 fixation rates. Moreover, we conducted a literature search on the comparison of the two methods to calculate a mean effect size using a meta-analysis approach. Our results indicate that the error potentially introduced by an equilibrium phase of the 15N2 gas is −72% at maximum for experiments with very short incubation times of 1 h. In contrast, the underestimation was negligible for incubations lasting 12–24 h (error is −0.2%). Our meta-analysis indicates that 84% of the measurements in the two groups will overlap and there is a 61% chance that a sample picked at random from the enriched water group will have a higher value than one picked at random from the bubble group. Overall, the underestimation of N2 fixation rates when using the bubble method relative to the enriched water method is highly dependent on incubation time and other experimental conditions and cannot be generalized

Nitrogen fixation rates in the Guinea Dome and the equatorial upwelling regions in the Atlantic Ocean

Biological nitrogen fixation is a key process balancing the loss of combined nitrogen in the marine nitrogen cycle. Its relevance in upwelling or high nutrient regions is still unclear, with the few available studies in these regions of the ocean reporting rates that vary widely from below detection limit to > 100 nmol N L−1 d−1. In the eastern tropical Atlantic Ocean, two open ocean upwelling systems are active in boreal summer. One is the seasonal equatorial upwelling, where the residual phosphorus associated with aged upwelled waters is suggested to enhance nitrogen fixation in this season. The other is the Guinea Dome, a thermal upwelling dome. We conducted two surveys along 23° W across the Guinea Dome and the Equator from 15° N to 5° S in September 2015 and August–September 2016 with high latitudinal resolution (20–60 nm between stations). The abundance of Trichodesmium colonies was characterized by an Underwater Vision Profiler 5 and the total biological nitrogen fixation in the euphotic layer was measured using the 15N2 technique. The highest abundances of Trichodesmium colonies were found in the area of the Guinea Dome (9°–15° N) with a maximum of 3 colonies L−1 near the surface. By contrast, colonies were almost absent in the Equatorial band between 2° N and 5° S. The highest nitrogen fixation rate was measured at the northern edge of the Guinea Dome in 2016 (ca. 31 nmol N L−1 d−1). In this region, where diazotrophs thrived on a sufficient supply of both phosphorus and iron, a patchy distribution was unveiled by our increased spatial resolution scheme. In the Equatorial band, rates were considerably lower, ranging from below detection limit to ca. 4 nmol N L−1 d−1, with a clear difference in magnitude between 2015 (rates close to zero) and 2016 (average rates around 2 nmol N L−1 d−1). This difference seemed triggered by a contrasting supply of phosphorus between years. Our study stresses the importance of surveys with sampling at fine-scale spatial resolution, and shows unexpected high variability in the rates of nitrogen fixation in the Guinea Dome, a region where diazotrophy is a significant process supplying new nitrogen into the euphotic layer.publishedVersio

Cross-shelf transport, oxygen depletion, and nitrate release within a forming mesoscale eddy in the eastern Indian Ocean

International audienceMesoscale eddies may drive a significant component of cross-shelf transport important in the ecology of shelf ecosystems and adjacent boundary currents. The Leeuwin Current in the eastern Indian Ocean becomes unstable in the austral autumn triggering the formation of eddies. We hypothesized that eddy formation represented the major driver of cross-shelf transport during the autumn. Acoustic Doppler Current Profiler profiles confirmed periodic offshore movement of 2 Sv of shelf waters into the forming eddy from the shelf, carrying a load of organic particles (>0.06 mm). The gap between inflow and outflow then closed, such that the eddy became isolated from further direct input of shelf waters. Drifter tracks supported an anticyclonic surface flow peaking at the eddy perimeter and decreasing in velocity at the eddy center. Oxygen and nutrient profiles suggested rapid remineralization of nitrate mid-depth in the isolated water mass as it rotated, with a total drawdown of oxygen of 3.6 mol m 22 to 350 m. Depletion of oxygen, and release of nitrate, occurred on the timescale of 1 week. We suggest that N supply and N turnover are rapid in this system, such that nitrate is acting primarily as a regenerated nutrient rather than as a source of new nitrogen. We hypothesize that sources of eddy particulate C and N could include particles sourced from coastal primary producers within 500 km such as macrophytes and sea-grasses known to produce copious detritus, which is prone to resuspension and offshore transport

Nitrogen sources and net growth efficiency of zooplankton in three Amazon River plume food webs

The plasticity of nitrogen specific net growth efficiency (NGE) in marine mesozooplankton is currently unresolved, with discordant lines of evidence suggesting that NGE is constant, or that it varies with nitrogen source, food availability, and food quality in marine ecosystems. Specifically, the fate of nitrogen from nitrogen fixation is poorly known. We use 15N : 14N ratios in plankton in combination with hydrological data, nutrient profiles, and nitrogen fixation rate measurements to investigate the relationship between new nitrogen sources and the nitrogen specific NGE in three plankton communities along the outer Amazon River plume. The NGE of small (200–500 μm) mesozooplankton was estimated from the δ 15N differences between particulate nitrogen and zooplankton using an open system Rayleigh fractionation model. The transfer efficiency of nitrogen among larger (\u3e 500 μm) mesozooplankton was estimated from the change in δ 15N as a function of zooplankton size. The Amazon River was not a significant source of bioavailable nitrogen anywhere in our study region, and subsurface nitrate was the primary new nitrogen source for the outer shelf community, which was dominated by diatoms. N2 fixation was the principal new nitrogen source at sites of high diatom diazotroph association abundance and at oceanic sites dominated by Trichodesmium spp. and Synechococcus spp. Although we found clear spatial differences in food quantity, food quality, and diazotroph inputs into mesozooplankton, our data show no significant differences in mesozooplankton nitrogen transfer efficiency and NGE (for latter, mean ± SD: 59 ± 10%) among sites