Nitrogen cycling driven by organic matter export in the South Pacific oxygen minimum zone

Oxygen minimum zones are expanding globally, and at present account for around 20-40% of oceanic nitrogen loss. Heterotrophic denitrification and anammox-anaerobic ammonium oxidation with nitrite-are responsible for most nitrogen loss in these low-oxygen waters. Anammox is particularly significant in the eastern tropical South Pacific, one of the largest oxygen minimum zones globally. However, the factors that regulate anammox-driven nitrogen loss have remained unclear. Here, we present a comprehensive nitrogen budget for the eastern tropical South Pacific oxygen minimum zone, using measurements of nutrient concentrations, experimentally determined rates of nitrogen transformation and a numerical model of export production. Anammox was the dominant mode of nitrogen loss at the time of sampling. Rates of anammox, and related nitrogen transformations, were greatest in the productive shelf waters, and tailed off with distance from the coast. Within the shelf region, anammox activity peaked in both upper and bottom waters. Overall, rates of nitrogen transformation, including anammox, were strongly correlated with the export of organic matter. We suggest that the sinking of organic matter, and thus the release of ammonium into the water column, together with benthic ammonium release, fuel nitrogen loss from oxygen minimum zones.

Net primary productivity, POC export, Th234 and U238 activities and abundance of autotrophs during U.S.C.G.C. Healy cruises HLY0802 and HLY0803

Seasonal patterns in the partitioning of phytoplankton carbon during receding sea ice conditions in the eastern Bering Sea water column are presented using rates of 14C net primary productivity (NPP), phototrophic plankton carbon content, and POC export fluxes from shelf and slope waters in the spring (March 30-May 6) and summer (July 3-30) of 2008. At ice-covered and marginal ice zone (MIZ) stations on the inner and middle shelf in spring, NPP averaged 76 ± 93 mmol C/m**2/d, and in ice-free waters on the outer shelf NPP averaged 102 ± 137 mmol C/m**2/d. In summer, rates of NPP were more uniform across the entire shelf and averaged 43 ± 23 mmol C/m**2/d over the entire shelf. A concomitant shift was observed in the phototrophic pico-, nano-, and microplankton community in the chlorophyll maximum, from a diatom dominated system (80 ± 12% autotrophic C) in ice covered and MIZ waters in spring, to a microflagellate dominated system (71 ± 31% autotrophic C) in summer. Sediment trap POC fluxes near the 1% PAR depth in ice-free slope waters increased by 70% from spring to summer, from 10 ± 7 mmol C/m**2/d to 17 ± 5 mmol C/m**2/d, respectively. Over the shelf, under-ice trap fluxes at 20 m were higher, averaging 43 ± 17 mmol C/m**2/d POC export over the shelf and slope estimated from 234Th deficits averaged 11 ± 5 mmol C/m**2/d in spring and 10 ± 2 mmol C/m**2/d in summer. Average e-ratios calculated on a station-by-station basis decreased by ~ 30% from spring to summer, from 0.46 ± 0.48 in ice-covered and MIZ waters, to 0.33 ± 0.26 in summer, though the high uncertainty prevents a statistical differentiation of these data.

Analysis of Recent benthic foraminiferal assemblages from the oxygen minimum zone of the Pakistan continental margin

Live (Rose Bengal stained) and dead benthic foraminiferal communities (hard-shelled species only) from the Pakistan continental margin oxygen minimum zone (OMZ) have been studied in order to determine the relation between faunal composition and the oxygenation of bottom waters. During R.R.S. Charles Darwin Cruises 145 and 146 (12 March to May 28 2003), 11 multicores were taken on the continental margin off Karachi, Pakistan. Two transects were sampled, constituting a composite bathymetric profile from 136 m (above the OMZ in spring 2003) down to 1870 m water depth. Cores (surface area 25.5 cm2) were processed as follows: for stations situated above, and in the upper part of the OMZ, sediment slices were taken for the 0-0.5 and 0.5-1 cm intervals, and then in 1 cm intervals down to 10 cm. For the lower part of the OMZ, the second centimetre was also sliced in half-centimetre intervals. Each sample was stored in 10 % borax-buffered formalin for further processing. Onshore, the samples were wet sieved over 63 µm, 150 µm and 300 µm sieves and the residues were stained for one week in ethanol with Rose Bengal. After staining, the residue was washed again. The stained faunas were picked wet in three granulometric fractions (63-150 µm, 150-300 µm and >300 µm), down to 10 cm depth. To gain more insight into the population dynamics we investigated the dead (unstained) foraminifera in the 2-3 cm level for the fractions 150-300 µm and >300 µm. The fractions >300 µm and 150-300 µm show nearly the same faunal distribution and therefore the results are presented here for both fractions combined (i.e. the >150 µm fraction). Live foraminiferal densities show a clear maximum in the first half centimetre of the sediment; only few specimens are found down to 4 cm depth. The faunas exhibit a clear zonation across the Pakistan margin OMZ. Down to 500 m water depth, Uvigerina ex gr. U. semiornata and Bolivina aff. B. dilatata dominate the assemblages. These taxa are largely restricted to the upper cm of the sediment. They are adapted to the very low bottom-water oxygen values (ab. 0.1 ml/l in the OMZ core) and the extremely high input of organic carbon on the upper continental slope. The lower part of the OMZ is characterized by cosmopolitan faunas, containing also some taxa that in other areas have been described in deep infaunal microhabitats.

Integrated values of heterotrophic picoplankton and primary production (euphotic zone) in the Antarctic polar frontal region during POLARSTERN cruises ANT-XIII/2 and ANT-XVI/3

We assessed relationships between phytoplankton standing stock, measured as chlorophyll a (Chl a), primary production (PP), and heterotrophic picoplankton production (HPP), in the epipelagic zone (0-100 m) as well as in the mesopelagic zone (100-1,000 m) in the polar frontal zone of the Atlantic sector of the Southern Ocean in austral summer (late December to January) and fall (March to early May). Integrated epipelagic HPP was positively correlated to integrated PP in summer (data for fall are not available) but not to integrated Chl a. However, integrated mesopelagic HPP was positively correlated to Chl a in summer as well as fall. The mesopelagic fraction of HPP as a percentage of total HPP was also positively correlated to Chl a, whereas the epipelagic fraction of HPP was negatively correlated to it. These results indicate that with increasing phytoplankton standing stock, constituted mainly of highly silicified diatoms, the focus of its consumption by heterotrophic picoplankton shifts from epipelagic to mesopelagic waters. With a growth efficiency of 30%, our HPP data indicate that in both the epipelagic and mesopelagic zone heterotrophic picoplankton consume 20% of PP. Mesopelagic heterotrophic picoplankton consumed around 80% of the sinking flux, measured from depletion of 234Th, which is a lower fraction than that reported from the central and subarctic Pacific. Our analysis indicates that it is important to include mesopelagic HPP in comprehensive assessments of the microbial consumption of PP, phytoplankton biomass, and particulate organic matter in cold oceanic systems with high rates of export production.

Jelly biomass sinking speed reveals a fast carbon export mechanism

Sinking of gelatinous zooplankton biomass is an important component of the biological pump removing carbon from the upper ocean. The export efficiency, e.g., how much biomass reaches the ocean interior sequestering carbon, is poorly known because of the absence of reliable sinking speed data. We measured sinking rates of gelatinous particulate organic matter (jelly-POM) from different species of scyphozoans, ctenophores, thaliaceans, and pteropods, both in the field and in the laboratory in vertical columns filled with seawater using high-quality video. Using these data, we determined taxon-specific jelly-POM export efficiencies using equations that integrate biomass decay rate, seawater temperature, and sinking speed. Two depth scenarios in several environments were considered, with jelly-POM sinking from 200 and 600 m in temperate, tropical, and polar regions. Jelly-POM sank on average between 850 and 1500 m/d (salps: 800-1200 m/d; ctenophores: 1200-1500 m/d; scyphozoans: 1000-1100 m d; pyrosomes: 1300 m/d). High latitudes represent a fast-sinking and low-remineralization corridor, regardless of species. In tropical and temperate regions, significant decomposition takes place above 1500 m unless jelly-POM sinks below the permanent thermocline. Sinking jelly-POM sequesters carbon to the deep ocean faster than anticipated, and should be incorporated into biogeochemical and modeling studies to provide more realistic quantification of export via the biological carbon pump worldwide.

Global database of surface ocean particulate organic carbon export fluxes diagnosed from the 234Th technique

Through the processes of the biological pump, carbon is exported to the deep ocean in the form of dissolved and particulate organic matter. There are several ways by which downward export fluxes can be estimated. The great attraction of the 234Th technique is that its fundamental operation allows a downward flux rate to be determined from a single water column profile of thorium coupled to an estimate of POC/234Th ratio in sinking matter. We present a database of 723 estimates of organic carbon export from the surface ocean derived from the 234Th technique. Data were collected from tables in papers published between 1985 and 2013 only. We also present sampling dates, publication dates and sampling areas. Most of the open ocean Longhurst provinces are represented by several measurements. However, the Western Pacific, the Atlantic Arctic, South Pacific and the South Indian Ocean are not well represented. There is a variety of integration depths ranging from surface to 220m. Globally the fluxes ranged from -22 to 125 mmol of C/m**2/d. We believe that this database is important for providing new global estimate of the magnitude of the biological carbon pump.

The seasonal sea-ice zone in the glacial Southern Ocean as a carbon sink, supplementary datasets

Reduced surface-deep ocean exchange and enhanced nutrient consumption by phytoplankton in the Southern Ocean have been linked to lower glacial atmospheric CO2. However, identification of the biological and physical conditions involved and the related processes remains incomplete. Here we specify Southern Ocean surface-subsurface contrasts using a new tool, the combined oxygen and silicon isotope measurement of diatom and radiolarian opal, in combination with numerical simulations. Our data do not indicate a permanent glacial halocline related to melt water from icebergs. Corroborated by numerical simulations, we find that glacial surface stratification was variable and linked to seasonal sea-ice changes. During glacial spring-summer, the mixed layer was relatively shallow, while deeper mixing occurred during fall-winter, allowing for surface-ocean refueling with nutrients from the deep reservoir, which was potentially richer in nutrients than today. This generated specific carbon and opal export regimes turning the glacial seasonal sea-ice zone into a carbon sink.