Oxygen Utilization and Downward Carbon Flux in an Oxygen-Depleted Eddy in the Eastern Tropical North Atlantic

The occurrence of mesoscale eddies that develop suboxic environments at shallow depth (about 40-100 m) has recently been reported for the eastern tropical North Atlantic (ETNA). Their hydrographic structure suggests that the water mass inside the eddy is well isolated from ambient waters supporting the development of severe near-surface oxygen deficits. So far, hydrographic and biogeochemical characterization of these eddies was limited to a few autonomous surveys, with the use of moorings, under water gliders and profiling floats. In this study we present results from the first dedicated biogeochemical survey of one of these eddies conducted in March 2014 near the Cape Verde Ocean Observatory (CVOO). During the survey the eddy core showed oxygen concentrations as low as 5 µmol kg-1 with a pH of around 7.6 at approximately 100 m depth. Correspondingly, the aragonite saturation level dropped to 1 at the same depth, thereby creating unfavorable conditions for calcifying organisms. To our knowledge, such enhanced acidity within near-surface waters has never been reported before for the open Atlantic Ocean. Vertical distributions of particulate organic matter and dissolved organic matter (POM and DOM), generally showed elevated concentrations in the surface mixed layer (0-70 m), with DOM also accumulating beneath the oxygen minimum. With the use of reference data from the upwelling region where these eddies are formed, the oxygen utilization rate was calculated by determining oxygen consumption through the remineralization of organic matter. Inside the core, we found these rates were almost 1 order of magnitude higher (apparent oxygen utilization rate (aOUR); 0.26 µmol kg-1 day-1) than typical values for the open North Atlantic. Computed downward fluxes for particulate organic carbon (POC), were around 0.19 to 0.23 g C m-2 day-1 at 100 m depth, clearly exceeding fluxes typical for an oligotrophic open-ocean setting. The observations support the view that the oxygen-depleted eddies can be viewed as isolated, westwards propagating upwelling systems of their own, thereby represent re-occurring alien biogeochemical environments in the ETNA.

Seawater carbonate chemistry and calcification rate in flume experiment

Ocean acidification (OA) poses a severe threat to tropical coral reefs, yet much of what is know about these effects comes from individual corals and algae incubated in isolation under high pCO2. Studies of similar effects on coral reef communities are scarce. To investigate the response of coral reef communities to OA, we used large outdoor flumes in which communities composed of calcified algae, corals, and sediment were combined to match the percentage cover of benthic communities in the shallow back reef of Moorea, French Polynesia. Reef communities in the flumes were exposed to ambient (400 ?atm) and high pCO2 (1300 ?atm) for 8 weeks, and calcification rates measured for the constructed communities including the sediments. Community calcification was reduced by 59% under high pCO2, with sediment dissolution explaining ~ 50% of this decrease; net calcification of corals and calcified algae remained positive but was reduced by 29% under elevated pCO2. These results show that, despite the capacity of coral reef calcifiers to maintain positive net accretion of calcium carbonate under OA conditions, reef communities might transition to net dissolution as pCO2 increases, particularly at night, due to enhanced sediment dissolution.

Part of the global DOC versus AOU (dissolved organic carbon/apparent oxygen utilization) data compilation, Arabian Sea

Concentrations of total organic carbon (TOC) were determined on samples collected during six cruises in the northern Arabian Sea during the 1995 US JGOFS Arabian Sea Process Study. Total organic carbon concentrations and integrated stocks in the upper ocean varied both spatially and seasonally. Highest mixed-layer TOC concentrations (80-100 µM C) were observed near the coast when upwelling was not active, while upwelling tended to reduce local concentrations. In the open ocean, highest mixed-layer TOC concentrations (80-95 µM C) developed in winter (period of the NE Monsoon) and remained through mid summer (early to mid-SW Monsoon). Lowest open ocean mixed-layer concentrations (65-75 µM C) occurred late in the summer (late SW Monsoon) and during the Fall Intermonsoon period. The changes in TOC concentrations resulted in seasonal variations in mean TOC stocks (upper 150 m) of 1.5-2 mole C/m**2, with the lowest stocks found late in the summer during the SW Monsoon-Fall Intermonsoon transition. The seasonal accumulation of TOC north of 15°N was 31-41 x 10**12 g C, mostly taking place over the period of the NE Monsoon, and equivalent to 6-8% of annual primary production estimated for that region in the mid-1970s. A net TOC production rate of 12 mmole C/m**2/d over the period of the NE Monsoon represented ~80% of net community production. Net TOC production was nil during the SW Monsoon, so vertical export would have dominated the export terms over that period. Total organic carbon concentrations varied in vertical profiles with the vertical layering of the water masses, with the Persian Gulf Water TOC concentrations showing a clear signal. Deep water (>2000 m) TOC concentrations were uniform across the basin and over the period of the cruises, averaging 42.3±1.4 µM C.

Part of the global DOC versus AOU (dissolved organic carbon/apparent oxygen utilization) data compilation, Pacific Ocean

Total organic carbon (TOC) was analyzed on four transects along 140°W in 1992 using a high temperature combustion/discrete injection (HTC/DI) analyzer. For two of the transects, the analyses were conducted on-board ship. Mixed-layer concentrations of organic carbon varied from about 80 µM C at either end of the transect (12°N and 12°S) to about 60 µM C at the equator. Total organic carbon concentrations decreased rapidly below the mixed-layer to about 38-40 µM C at 1000 m across the transect. Little variation was observed below this depth; deep water concentrations below 2000 m were virtually monotonic at about 36 µM C. Repeat measurements made on subsequent cruises consistently found the same concentrations at 1000 m or deeper, but substantial variations were observed in the mixed-layer and the upper water column above 400 m depth. Linear mixing models of total organic carbon versus sigmaT exhibited zones of organic carbon formation and consumption. TOC was found to be inversely correlated with apparent oxygen utilization (AOU) in the region between the mixed-layer and the oxygen minimum. In the mixed-layer, TOC concentrations varied seasonally. Part of the variations in TOC at the equator was driven by changes in the upwelling rate in response to variations in physical forcing related to an El Niño and to the passage of tropical instability waves. TOC export fluxes, calculated from simple box models, averaged 8±4 mmol C/m**2/day at the equator and also varied seasonally. These export fluxes account for 50-75% of the total carbon deficit and are consistent with other estimates and model predictions.

Part of the global DOC versus AOU (dissolved organic carbon/apparent oxygen utilization) data compilation, BATS

The contributions of total organic carbon and nitrogen to elemental cycling in the surface layer of the Sargasso Sea are evaluated using a 5-yr time-series data set (1994-1998). Surface-layer total organic carbon (TOC) and total organic nitrogen (TON) concentrations ranged from 60 to 70 µM C and 4 to 5.5 µM N seasonally, resulting in a mean C : N molar ratio of 14.4±2.2. The highest surface concentrations varied little during individual summer periods, indicating that net TOC production ceased during the highly oligotrophic summer season. Winter overturn and mixing of the water column were both the cause of concentration reductions and the trigger for net TOC production each year following nutrient entrainment and subsequent new production. The net production of TOC varied with the maximum in the winter mixed-layer depth (MLD), with greater mixing supporting the greatest net production of TOC. In winter 1995, the TOC stock increased by 1.4 mol C/m**2 in response to maximum mixing depths of 260 m. In subsequent years experiencing shallower maxima in MLD (<220 m), TOC stocks increased <0.7 mol C/m**2. Overturn of the water column served to export TOC to depth (>100 m), with the amount exported dependent on the depth of mixing (total export ranged from 0.4 to 1.4 mol C/m**2/yr). The exported TOC was comprised both of material resident in the surface layer during late summer (resident TOC) and material newly produced during the spring bloom period (fresh TOC). Export of resident TOC ranged from 0.5 to 0.8 mol C/m**2/yr, covarying with the maximum winter MLD. Export of fresh TOC varied from nil to 0.8 mol C/m**2/yr. Fresh TOC was exported only after a threshold maximum winter MLD of ~200 m was reached. In years with shallower mixing, fresh TOC export and net TOC production in the surface layer were greatly reduced. The decay rates of the exported TOC also covaried with maximum MLD. The year with deepest mixing resulted in the highest export and the highest decay rate (0.003 1/d) while shallow and low export resulted in low decay rates (0.0002 1/d), likely a consequence of the quality of material exported. The exported TOC supported oxygen utilization at dC : dO2 molar ratios ranging from 0.17 when TOC export was low to 0.47 when it was high. We estimate that exported TOC drove 15-41% of the annual oxygen utilization rates in the 100-400 m depth range. Finally, there was a lack of variability in the surface-layer TON signal during summer. The lack of a summer signal for net TON production suggests a small role for N2 fixation at the site. We hypothesize that if N2 fixation is responsible for elevated N : P ratios in the main thermocline of the Sargasso Sea, then the process must take place south of Bermuda and the signal transported north with the Gulf Stream system.

Impact of ocean acidification and warming on respiration, heart rate and acid-base status of Mytilus edulis from the North Sea

Anthropogenic climate change confronts marine organisms with rapid trends of concomitant warming and CO2 induced ocean acidification. The survival and distribution of species partly depend on their ability to exploit their physiological plasticity during acclimatization. Therefore, in laboratory studies the effects of simulated future ocean acidification on thermal tolerance, energy metabolism and acid-base regulation capacity of the North Sea population of the blue mussel Mytilus edulis were examined. Following one month of pre-acclimation to 10 °C and control CO2 levels, mussels were exposed for two weeks to control and projected oceanic CO2 levels (390, 750 and 1120 µatm) before being subjected to a stepwise warming protocol between 10 °C and 31 °C (+ 3 °C each night). Oxygen consumption and heart rates, anaerobic metabolite levels and haemolymph acid-base status were determined at each temperature. CO2 exposure left oxygen consumption rate unchanged at acclimation temperature but caused a somewhat stronger increase during acute warming and thus mildly higher Q10-values than seen in controls. Interestingly, the thermally induced limitation of oxygen consumption rate set in earlier in normocapnic than in hypercapnic (1120 µatm CO2) mussels (25.2 °C vs. 28.8 °C), likely due to an onset of metabolic depression in the control group following warming. However, the temperature induced increase in heart rate became limited above 25 °C in both groups indicating an unchanged pejus temperature regardless of CO2 treatment. An upper critical temperature was reached above 28 °C in both treatments indicated by the accumulation of anaerobic metabolites in the mantle tissue, paralleled by a strong increase in haemolymph PCO2 at 31 °C. Ocean acidification caused a decrease in haemolymph pH. The extracellular acidosis remained largely uncompensated despite some bicarbonate accumulation. In all treatments animals developed a progressive warming-induced extracellular acidosis. A stronger pH drop at around 25 °C was followed by stagnating heart rates. However, normocapnic mussels enhanced bicarbonate accumulation at the critical limit, a strategy no longer available to hypercapnic mussels. In conclusion, CO2 has small effects on the response patterns of mussels to warming, leaving thermal thresholds largely unaffected. High resilience of adult North Sea mussels to future ocean acidification indicates that sensitivity to thermal stress is more relevant in shaping the response to future climate change.

Enhancement of photosynthetic carbon assimilation efficiency by phytoplankton in the future coastal ocean

A mesocosm experiment was conducted to evaluate the effects of future climate conditions on photosynthesis and productivity of coastal phytoplankton. Natural phytoplankton assemblages were incubated in field mesocosms under the ambient condition (present condition: ca. 400 ppmv CO2 and ambient temp.), and two future climate conditions (acidification condition: ca. 900 ppmv CO2 and ambient temp.; greenhouse condition: ca. 900 ppmv CO2 and 3 °C warmer than ambient). Photosynthetic parameters of steady-state light responses curves (LCs; measured by PAM fluorometer) and photosynthesis-irradiance curves (P-I curves; estimated by in situ incorporation of 14C) were compared to three conditions during the experiment period. Under acidification, electron transport efficiency (alpha LC) and photosynthetic 14C assimilation efficiency (alpha) were 10% higher than those of the present condition, but maximum rates of relative electron transport (rETRm,LC) and photosynthetic 14C assimilation (PBmax) were lower than the present condition by about 19% and 7%, respectively. In addition, rETRm,LC and alpha LC were not significantly different between and greenhouse conditions, but PBmax and alpha of greenhouse conditions were higher than those of the present condition by about 9% and 30%, respectively. In particular, the greenhouse condition has drastically higher PBmax and alpha than the present condition more than 60% during the post-bloom period. According to these results, two future ocean conditions have major positive effects on the photosynthesis in terms of energy utilization efficiency for organic carbon fixation through the inorganic carbon assimilation. Despite phytoplankton taking an advantage on photosynthesis, primary production of phytoplankton was not stimulated by future conditions. In particular, biomass of phytoplankton was depressed under both acidification and greenhouse conditions after the the pre-bloom period, and more research is required to suggest that some factors such as grazing activity could be important for regulating phytoplankton bloom in the future ocean.