Andrew P. Rees

Plankton respiration in the Eastern Atlantic Ocean

Concurrent measurements of dark community respiration (DCR), gross production (GP), size fractionated primary production (14C PP), nitrogen uptake, nutrients, chlorophyll a concentration, and heterotrophic and autotrophic bacterial abundance were collected from the upper 200 m of a latitudinal (32°S–48°N) transect in the Eastern Atlantic Ocean during May/June 1998. The mean mixed layer respiration rate was 2.5±2.1 mmol O2 m−3 d−1 (n=119) for the whole transect, 2.2±1.1 mmol O2 m−3 d−1 (n=32) in areas where chlorophyll a was <0.5 mg m−3 and 1.5±0.7 mmol O2 m−3 d−1 (n=10) where chlorophyll a was <0.2 mg m−3. These values lie within the range of published data collected in comparable waters, they co-vary with indicators of heterotrophic and autotrophic biomass (heterotrophic bacterial abundance, chlorophyll a concentration, beam attenuation and particulate organic carbon concentration) and they can be reconciled with accepted estimates of total respiratory activity. The mean and median respiratory quotient (RQ), calculated as the ratio of dissolved inorganic carbon production to dissolved oxygen consumption, was 0.8 (n=11). At the time of the study, plankton community respiration exceeded GP in the picoautotroph dominated oligotrophic regions (Eastern Tropical Atlantic [15.5°S–14.2°N] and North Atlantic Subtropical Gyre [21.5–42.5°N]), which amounted to 50% of the stations sampled along the 12,100 km transect. These regions also exhibited high heterotrophic: autotrophic biomass ratios, higher turnover rates of phytoplankton than of bacteria and low f ratios. However, the carbon supply mechanisms required to sustain the rates of respiration higher than GP could not be fully quantified. Future research should aim to determine the temporal balance of respiration and GP together with substrate supply mechanisms in these ocean regions.

Nitrogen fixation and nitrogenase (nifH) expression in tropical waters of the eastern North Atlantic

Expression of nifH in 28 surface water samples collected during fall 2007 from six stations in the vicinity of the Cape Verde Islands (north-east Atlantic) was examined using reverse transcription-polymerase chain reaction (RT-PCR)-based clone libraries and quantitative RT-PCR (RT-qPCR) analysis of seven diazotrophic phylotypes. Biological nitrogen fixation (BNF) rates and nutrient concentrations were determined for these stations, which were selected based on a range in surface chlorophyll concentrations to target a gradient of primary productivity. BNF rates greater than 6 nmolN l−1 h−1 were measured at two of the near-shore stations where high concentrations of Fe and PO43− were also measured. Six hundred and five nifH transcripts were amplified by RT-PCR, of which 76% are described by six operational taxonomic units, including Trichodesmium and the uncultivated UCYN-A, and four non-cyanobacterial diazotrophs that clustered with uncultivated Proteobacteria. Although all five cyanobacterial phylotypes quantified in RT-qPCR assays were detected at different stations in this study, UCYN-A contributed most significantly to the pool of nifH transcripts in both coastal and oligotrophic waters. A comparison of results from RT-PCR clone libraries and RT-qPCR indicated that a γ-proteobacterial phylotype was preferentially amplified in clone libraries, which underscores the need to use caution interpreting clone-library-based nifH studies, especially when considering the importance of uncultivated proteobacterial diazotrophs.

New production of the NW Iberian shelf during the upwelling season over the period 1982–1999

New production (NP) is calculated for NW Iberian shelf waters from 421 to 431N (3500 km 2 ), at the fortnight, upwelling-season (March–October) and inter-annual time-scales. The time series used are (1) upwelling rates (daily values of offshore Ekman transport from 1982 to 1999), (2) bottom shelf temperatures (twice a week values from 1987 to 1999), and (3) the nutrient–temperature relationships ofupwelled Eastern North Atlantic Central Water (ENACW) obtained during 14 hydrographic cruises to the study area (between 1977 and 1998). Marked inter-annual variability is observed, both at the fortnight and the seasonal time-scales. Average NP over the upwelling-season ranged from 330 to 815 mg C m � 2 d � 1 (mean, 4907145 mg C m � 2 d � 1 ) in the 1982–1999 period. Large inter-annual changes ofupwelling rates are the reason behind the NP fluctuations: 83% ofthe variability ofNP can be explained by the offshore Ekman Transport ð� QX Þ: NP is compared with satellite-derived net microbial community production (NCP) during the 1998– 1999 upwelling seasons, when SeaWiFS images are available. An average upwelling-season NP/NCP ratio of0.33 was obtained, indicating that 67% of NCP is respired in situ and 33% is exported off-shelf to the surrounding oligotrophic ocean.

Nitrous oxide: Estuarine sources and atmospheric flux

Nitrous oxide (N 2 O), nitrate, nitrite and ammonium concentrations were determined along axial surface water profiles of the Tamar Estuary, South West England, on five occasions. Physical variables, including suspended solids, salinity and temperature were also measured. In addition, a temporal survey was carried out in which these variables were measured at a fixed point over a tidal cycle. Nitrous oxide was supersaturated throughout the estuary on all sampling occasions, with a maximum in the low salinity region. The N 2 O maximum exhibited no significant spatial deviation with season or tidal state, in contrast to the other determined variables. Nitrous oxide supersaturation resulted predominantly from release by sediments, which was potentially enhanced by tidal resuspension and bioturbation. Nitrification in the water column associated with the turbidity maximum was considered to be a secondary seasonal source of nitrous oxide. External sources, such as terrestrial runoff and sewage input also contributed. The estuary represents a source of atmospheric nitrous oxide throughout the year.

Primary production and nutrient assimilation in the Iberian upwelling in August 1998

Primary production was measured during two Lagrangian experiments in the Iberian upwelling. The first experiment, in a body of upwelled water, measured day-to-day changes in phytoplankton activity as the water mass moved south along the shelf break. Nutrient concentrations decreased over a five day period, with concomitant increases in phytoplankton biomass. Initially the maximum phytoplankton biomass was in the upper 10m but after four days, a sub-surface chlorophyll maximum was present at 30m. Depth-integrated primary production at the beginning of the experiment was 70mmolC.m−2.d−1 (838mgC.m−2.d−1) and reached a maximum of 88mmolC.m−2.d−1 (1053mgC.m−2.d−1) on day 3. On day 1, the picoplankton fraction ( 5μm) phytoplankton, but the increase in overall production during the drift experiment was by these larger cells. Nitrate was the dominant nitrogen source. As nutrient concentrations declined, ammonium became increasingly more important as a nitrogen source and the f-ratio decreased from 0.7 to 0.5. Picoplankton cells (<2μm) were responsible for most (65–80%) of the ammonium uptake. The C:N:P uptake ratios were very close to the Redfield ratio for the first four days but as nutrients became depleted high C:N uptake ratios (11 to 43) were measured. Over the period of the experiment, nitrate concentration within the upper 40m decreased by 47.91mmolN.m−2. In vitro estimates, based on 15N nitrate uptake, accounted for 56% of the decrease in nitrate concentration observed in the drifting water mass. Ammonium uptake over the same four day period was 16.28mmolN.m−2, giving a total nitrogen uptake of 43.18mmolN.m−2. In the second experiment, an offshore filament was the focus and a water mass was sampled as it moved offshore. Nutrient concentrations were very low (nitrate was <10nmol l−1 and ammonium was 20–40nmol l−1). Primary production rate varied between 36mmolC.m−2.d−1 (436mgC.m−2.d−1) and 21mmolC.m−2.d−1 (249mgC.m−2.d−1). Picophytoplankton was the most productive fraction and was responsible for a constant proportion (ca 0.65) of the total carbon fixation. Uptake rates of both nitrate and ammonium were between 10 and 20% of those measured in the upwelling region. Urea could be a very significant nitrogen source in these waters with much higher uptake rates than nitrate or ammonium; urea turnover times were ca. one day but the source of the urea remains unknown. Urea uptake had a profound effect on calculated f ratios. If only nitrate and ammonium uptake was considered, f ratios were calculated to be 0.42–0.46 but inclusion of urea uptake reduced the f ratio to <0.1. The primary production of this oligotrophic off-shore filament was driven by regenerated nitrogen.

Two Lagrangian experiments in the Iberian upwelling system: tracking an upwelling event and an off-shore filament

Abstract Two Lagrangian drift experiments were carried out at the NW Iberian margin. The first tracked a body of nutrient-rich, upwelled water as it moved south along the shelf break over a 5 day period. The second experiment, of similar duration, followed a water mass as it moved into the deep ocean in an off-shelf filament. This paper describes the background to and aims of each experiment. The overall objective was to quantify chemical and biological processes relating to the additional potential for the ocean at the shelf margins to sequester atmospheric CO2 in upwelling regions. The first experiment began at a time of intense wind-driven upwelling; within 2 days, the wind speed had moderated and the system entered a relaxation period with greatly reduced upwelling. The patch of upwelled water was marked by a single buoy array and it moved south along the shelf break. Transport was initially rapid but slowed with reducing wind speed. The temperature–salinity characteristics were consistent with sampling only a single water mass throughout the experiment. A model of particle trajectories showed slight deviation from the actual movement of the marked water mass, but overall the data support the assumption that the experiment was Lagrangian. During a 5 day experimental period, nutrients were utilised with a N:P ratio of 18.3 and N:Si of 4. Nutrient concentrations first reduced in the near-surface but depletion deepened in the water column during the experiment. At the beginning of the experiment, the highest chlorophyll concentrations were in the surface 15m but this was replaced by a subsurface chlorophyll maximum at 30m. There was a shift from a small flagellate and dinoflagellate dominated photosynthetic phytoplankton assemblage to a diatom dominated assemblage. A high biomass of heterotrophic dinoflagellates and ciliates was also present. Canonical correlation analysis between environmental variables and microplankton assemblages, as defined by principal component analysis, suggested that a considerable part of DON production resulted from trophic relationships rather than direct release from phytoplankton. The second experiment followed a water mass marked with 5 Argos drifting buoys for 5 days as the water drifted off shelf in an off-shore filament. This water mass was extremely oligotrophic; nitrate concentrations were typically The data presented in this paper are a general description of the experiments and form the background to the more detailed descriptions given in the individual papers that make up this Special Issue of Progress in Oceanography.

Simultaneous high-precision measurements of methane and nitrous oxide in water and seawater by single phase equilibration gas chromatography

A novel semi-automated gas chromatographic technique for the simultaneous high-precision measurement of the partial pressures and concentrations of nitrous oxide (N2O) and methane (CH4) in discrete samples of water and air has been developed. For seawater analysis, samples collected in glass 11 volumetric flasks, with the exclusion of all air, are thermally equilibrated to 25.0 ± 0.05°C in a water bath and then rapidly equilibrated with a headspace of known N2O and CH4 compositions in a closed system at ambient pressure. The equilibrated headspace gases are separated on Porapak Q columns, followed by electron capture detection for N20 and flame ionisation detection for CH4. Complex sample purging or cold trapping procedures are avoided, eliminating the sources of major error associated with previous methods. System hardware requirements are comparatively modest, simplifying operation and maintenance, and minimal sample handling restricts the potential for sample outgassing and air contamination during analysis. Seawater samples can be routinely processed at a rate of 5–6 h−1, and air or calibration standards at a rate of ∼ 20 h- 1. For high-quality oceanographic work, individual seawater samples are each compared with two calibration standards and a sample of ambient air pumped from the ships bow. In this mode, a 12-bottle hydrocast can be processed in 3–4h. Routine analytical precisions for atmospheric air and for seawaters with saturations in the range 0–300% N2O and 0–160% CH4 relative to atmospheric air, are better than ±0.5% (100σ/x) for N2O and ±0.9% (100σ/x) for CH4. Detection limits are equivalent to ∼40pmoll−1 N2O and ∼50–150pmoll−1 CH4 for salinity 35 seawater at 20°C, although in practice the detectors always operate far above these ranges. Some N2O and CH4 data for seawater and air from the Bellingshausen Sea off Antarctica (62°–68°S, 65°–86°W) and the northwestern Indian Ocean (03°–26°N, 56°–59°E) are presented.

Temporal variability of denitrification in estuarine sediments

Abstract Sediment denitrification rates and fluxes of nitrous oxide, nitrate, nitrite and ammonium were determined at two intertidal sites in the Tamar estuary (S.W. England). High sediment nitrate uptake rates were recorded throughout the year, whereas the nitrite and ammonium fluxes were positive (from sediment to water column), with the former resulting from nitrification. Nitrous oxide flux was also positive, being largely attributable to denitrification with some contribution from nitrification or nitrification-denitrification coupling. No relationship was apparent between denitrification rate and nitrate concentration in the overlying water, invalidating the notion that denitrification automatically regulates nitrate during periods of elevated ambient concentration. However, denitrification exhibited a strong covariance with the degree of sediment bioturbation (Nereis diversicolor), which was considered to be attributable to increased transport and supply of nitrate via Nereis burrows. Denitrification accounted for 8·5% of the annual total nitrate loading to the Tamar estuary, although a maximum of 100% was observed in summer when the phytoplankton nutrient requirement would also be highest.

Biogeochemical cycling of dissolved zinc along the GEOTRACES South Atlantic transect GA10 at 40°S

The biogeochemical cycle of zinc (Zn) in the South Atlantic, at 40°S, was investigated as part of the UK GEOTRACES program. To date there is little understanding of the supply of Zn, an essential requirement for phytoplankton growth, to this highly productive region. Vertical Zn profiles displayed nutrient-like distributions with distinct gradients associated with the water masses present. Surface Zn concentrations are among the lowest reported for the worlds oceans ( 2 = 0.97, n = 460). Our results suggest that the use of a global Zn-Si relationship would lead to an underestimation of dissolved Zn in deeper waters of the South Atlantic. By utilizing Si* and a new tracer Zn* our data indicate that the preferential removal of Zn in the Southern Ocean prevented a direct return path for dissolved Zn to the surface waters of the South Atlantic at 40°S and potentially the thermocline waters of the South Atlantic subtropical gyre. The importance of Zn for phytoplankton growth was evaluated using the Zn-soluble reactive phosphorus (SRP) relationship. We hypothesize that the low Zn concentrations in the South Atlantic may select for phytoplankton cells with a lower Zn requirement. In addition, a much deeper kink at ~ 500m in the Zn:SRP ratio was observed compared to other oceanic regions. Key Points Dissolved zinc biogeochemical cycle investigated in the South Atlantic Ocean ©2014. American Geophysical Union. All Rights Reserved.

Primary production and nitrogen assimilation in the North sea during July 1987

Abstract An extensive survey of the whole of the North Sea, carried out in July 1987, is described. The study concentrated on the measurement of surface water inorganic nutrient concentrations and concomitant rates of primary production (14C) and nitrogen assimilation (15N). Primary production was investigated using size fractionation techniques. Three vertical profiles of primary production and nitrogen assimilation were also investigated. Much of the North Sea exhibed thermal stratification. Surface nutrient concentrations were low and chlorophyll concentrations typically The water column was vertically well mixed in the coastal zones. Here, inorganic nitrogen concentrations were high (e.g. up to 25 μmol NO3− with chlorophyll concentrations up to 10 mg−3, and organisms >5 μm diameter accounted for most of the primary production. As in offshore regions ammonium accounted for the major part of the nitrogen assimilated. A 115 km section obtained using an undulating oceanographic recorder showed that in certain regions of the North Sea physical features acted to increase the dependence of the phytoplankton on nitrate.